THE AMERICAN MIDLAND NATURALIST

Monograph No. 1

THE AMERICAN MIDLAND NATURALIST

Monograph Series

EDITORIAL STAFF

Theodor Just Botany

Editor, University of Notre Dame

Edward A. Chapin Entomology

U. S. National Museum, Washington, D. C.

Kenneth W. Cooper Cytology and Genetics

Princeton University, Princeton, New Jersey

Carroll Lane Fenton Invertebrate Paleontology

Rutgers University, New Brunswick, N. J.

John Hobart Hoskins Paleobotany

University of Cincinnati, Cincinnati, Ohio

Remington Kellogg Mammalogy

U. S. National Museum, Washington, D. C.

Jean Myron Linsdale Ornithology

Hastings Reservation, Monterey, California

George Willard Martin Mycology

State University of Iowa, Iowa City, Iowa

Karl Patterson Schmidt Ichthyology and Herpetology

Chicago Natural History Museum, Chicago, Illinois

Harley Jones Van Cleave Invertebrate Zoology

University of Illinois, Urbana, Illinois

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THE AMERICAN MIDLAND NATURALIST

Monograph No. 1

n©. /

Edited by Theodor Just

Published by the University of Notre Dame,

Notre Dame, Ind.

The Argasidae of North America,
Central America and Cuba

By R. A. COOLEY

(Senior Entomologist)

and

GLEN M. KOHLS

(Associate Entomologist)

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Contribution from the Rocky Mountain Laboratory, Hamil-
ton, Montana. Division of Infectious Diseases, National
Institute of Health, United States Public Health Service.

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The University Press

Notre Dame, Ind.

June, 1944

Copyright, 1944

by

The American Midland Naturalist

University of Notre Dame

Notre Dame, Ind.

CONTENTS

Introduction

The Medical and Veterinary Importance of the Argasidae 3

Explanation of Terms 4

Principal Characters of Argasidae. Figure 1 6

Methods 7

Family Argasidae Canestrini 1 I

Key to the Genera '-

Genus Argas Latreille ' 3

Key to Species of Argas 13

Argas reflexus (Fabricius) 14

Argas persicus (Oken) 1/

Genus Otobius Banks -'

Otobius megnini (Duges) 21

Otobius lagophihls Cooley and Kohls 32

Genus Ornilhodoros Koch 37

Key to Species of Ornilhodoros 3/

Ornilhodoros savignyi (Audouin) 38

Ornilhodoros eremicus Cooley and Kohls 40

Ornilhodoros coriaceus Koch 42

Ornilhodoros hermsi Wheeler, Herms, and Meyer 46

Ornilhodoros nicollei Mooser 50

Ornilhodoros cooleyi Mclvor 52

Ornilhodoros luricuta (Duges) 56

Ornilhodoros parf(eri Cooley 62

Ornilhodoros vumalensis Cooley and Kohls 74

Ornilhodoros brodvi Matheson 80

Ornilhodoros talaje (Guerin-Meneville) 82

Ornilhodoros concanensis Cooley and Kohls 88

Ornilhodoros stageri Cooley and Kohls "I

Ornilhodoros dveri Cooley and Kohls 95

Ornilhodoros amblus Chamberhn 99

Ornilhodoros rudis Karsch 101

Ornilhodoros dunni Matheson 103

Ornilhodoros viguerasi Cooley and Kohls 106

Ornilhodoros azteci Matheson 109

Genus Antricola Cooley and Kohls 118

Key to Species of Antricola 118

Antricola coprophilus (Mcintosh) 118

Antricola marginatum (Banks) 123

Plates 1 to 14 127

Classified List of Hosts 141

Geographical Distribution of Species 144

Bibliography 146

Index 151

58100

The Argasidac of North America, Central
America and Cuba

Introduction

The taxonomic study of the Argasidae has been beset with difficult
problems, some of which are stil! unsolved. The principal difficulty has been
and still is the lack of adequate generic characters. Some of the earlier species
were described from very few specimens and frequently with very little knowl-
edge of their biologies. By 1908, Neumann and Nuttall had notably improved
the situation by better generic definitions and the use of new specific characters.
Both retained as genera only Argas and Ornithodoros. In 1912 Banks added
the genus Otobtus, and in 1942 Antricola was added by the authors.

In 1908 only some twenty species were known. Now in North America
alone there are at least twenty-five species and about thirty new species have
been described from other countries. The characters exhibited by these new
species have served to clarify certain of the taxonomic problems; they have
made others more confusing. Some specific distinctions have found corrobora-
tive support in the biologies. With the increasing number of new species, it
has become more and more difficult to separate Argas and Ornithodoros and
we have found it necessary to change the previous conceptions of these genera.
However, no genera have been synonymized. It is probable that subsequent
workers will find others necessary but it is earnestly hoped that none will be
erected until reliable generic criteria have been clearly established. The present
names serve the purpose and should be changed only for permanent reasons.

In a previous paper1 the senior author emphasized the importance of
variation within species. This occurs principally in the ornamentation, and
the morphology is reasonably constant. In the Argasidae it is perhaps too
early to evaluate fully the variation within species, but at least it is evident
that it is of little importance in the genus Ornithodoros. Ornamentation is
absent in the known species of the Argasidae except in Ornithodoros coriaceus.

Where confusion has arisen in specific identities, it has been due largely
to overlooking characters in the mammillae and hypostome, some of which
are not very easily detected, and the very distinct ones found in the larval
stage which heretofore has not been thoroughly studied.

Host specificity varies in the numerous species. So far as known several

1 CoOLEY, R. A. The Genera Dermacenlor and Otocenlor (Ixodidae) in the United
States, with studies in variation. National Institute of Health Bulletin No. 171, Gov-
ernment Printing Office, 1938.

1

2 American Midland Naturalist Monograph No. 1

are host specific for bats though the bat tick, Ornithodoros stageri, also attacks
man readily. Nymphs and adults of O. dunni feed readily on guinea pigs and
white rats, and the larvae of O. kelleyi feed on guinea pigs. Species which are
found in the burrows and retreats of animals other than bats will apparently
attack any available host. Still others have been recorded from restricted lists.

During recent years a surprising number of new American species have
been found on bats. The original one, Antncola marginatus (Banks), was
described from Cuba in 1910. The new ones have been described during the
past six years. In 1935, Dr. Robert Matheson, of Cornell University, described
three from Panama and the Canal Zone and Mr. Allen Mcintosh, of the
Bureau of Animal Industry, United States Department of Agriculture, de-
scribed one from bat guano from Mexico and Arizona. We have recently
described a new species from Utah and Colorado, four others from Southern
States and one from Cuba. There are likely still others that infest bats in the
United States and probably in Mexico and Central America. We already
have larval specimens of undescribed species from Southwestern States of
which the adult and nymph are unknown.

Most of the material studied has been collected by staff members of the
Rocky Mountain Laboratory. Dr. Gordon E. Davis, in connection with his
studies on Ornithodoros as vectors of relapsing fever, has made many collec-
tions in the field and has furnished valuable laboratory-reared specimens.

The authors wish to express their appreciation for the assistance given by various
persons and institutions and especially the following: Dr. Jos. C. Bequaert, Harvard
University, for valuable specimens and for bringing to our attention certain items of tax-
onomy and synonymy; Dr. Robert Matheson, Cornell University, Dr. William A.
Riley, University of Minnesota, Dr. C. A. Herrick, University of Wisconsin, and Dr.
W. B. Herms, University of California, for very useful materials and helpful coopera-
tion; Drs. B. Schwartz, E. W. Price and Mr. Allen Mcintosh, Bureau of Animal
Industry, United States Department of Agriculture, for cooperation and facilities for
study while in Washington and for permission to use illustrations from the Seventeenth
Report of the Bureau of Animal Industry, for assistance in many ways while the senior
author was a visitor at his laboratories and for permission to use his drawings of Aniri-
cola coprophilus ; Drs. E. A. Chapin and H. E. Ewing for facilities and assistance while
at the National Museum; Dr. F. C. Bishopp, for assistance in locating critical material;
Dr. Barbara C. Mclvor for lending very valuable specimens; Dr. Chas. M. Wheeler,
Dr. D. E. Howell, and Mr. Thomas F. Kelley for sending ticks en various occasions;
Mr. Kenneth E. Stager, Los Angeles County Museum of History, Science and Art, for
assistance and materials; Dr. J. M. Linsdale, University of California, for opportunity to
study the collection of the Hastings Natural History Reservation; Mr. Robert Holden-
ried, for opportunity to study the collection of the Hooper Foundation; Dr. Charles T.
Vorhies and Mr. R. A. Flock, University of Arizona, for specimens and assistance;
Mr. Robert Traub, University of Illinois, for valuable specimens; Dr. I. Perez
Vigueras, for several collections of ticks from Cuba; and Dr. E. Brumpt for especially
valuable and critical materials.

In the tables of collection data the following abbreviations have been used:
R.M.L. - Rocky Mountain Laboratory; B.A.I. Bureau of Animal Indus-
try, United States Department of Agriculture; U.S.N.M. - - United States
National Museum; A - - adult; N. -- nymph; L. — larva; spec. -- specimens.

COOLEY & KOHLS: ARGASIDAE OF N. AMERICA, ETC.

The Medical and Veterinary Importance of the Argasidae

Several species of the Argasidae are of known medical or veterinary im-
portance.

Five species of Ormthodoros in this hemisphere, hermsi, tuncata, parkeri,
talaje, and rudis, are proved vectors of relapsing fever spirochetes. O. hermsi
is present in California, Nevada, Idaho, Oregon, and Colorado. O. tuncata is
found in California, Arizona, New Mexico, Texas, Oklahoma, Kansas, Colo-
rado, Utah, Florida, and in Mexico. O. parkeri is present in Washington,
Oregon, California, Idaho, Nevada, Montana, Wyoming, Utah, and Colo-
rado. O. talaje is present in California, Arizona, Nevada, Texas, Kansas,
Florida, and Central America. O. rudis is known in Panama, Mexico, and
South America.

Other species may prove to be of importance as disease vectors. Davis has
reported (Section VII, Sixth Pacific Science Congress, Berkeley, Calif., July
1939) the experimental transmission of Rocky Mountain spotted fever by O.
parkeri and has also demonstrated similar transmission of this disease and
American Q fever by O. hermsi.

O. coriaceus, of California and Mexico, while not known to be a vector of
any specific disease agent, readily attacks man. Its "bite" is quite painful.
Herms, who has studied this species, considers it to be one of the most venom-
ous of our ticks.

O. nicoltei, a Mexican species of which we have one record from the
United States (from snakes in a St. Louis, Missouri, zoo), was reported by
Mooser as occurring in native huts and feeding on man in Mexico.

Brumpt was unable to transmit iuricata strains of relapsing fever spirochetes
with nicollei but found that the species would transmit experimentally the
telapsing fever of Turkestan.

Very little is known concerning the several species in the Southwest recent-
ly described from bats and bat retreats. One of these, O. stageri, promptly
attacks man when opportunity permits. While its "bites" are rather severe, the
species is not known to be concerned in the transmission of any disease. An-
other, O. kelleyi, probably primarily associated with bats, has been found in
houses in Minnesota, Iowa, Wisconsin, Illinois, New York, and Pennsylvania
but is not known to attack man. Further settlement in Western States may
increase the possibilities of association of the various bat-infesting species
with man.

Argas persicus, of almost world-wide distribution in warm climates, is a
notorious pest of poultry. It is the vector of avian spirochetosis in many Old
World regions and in the New World the disease has been reported from
Brazil, Panama, and Cuba. Brown and Cross (1941) published results of
experiments indicating that the species may be a vector of fowl paralysis in
Texas.

Otobius megnini, the spinose ear tick, is a serious pest of cattle in most of

4 American Midland Naturalist Monograph No. ]

the Southwestern United States and parts of Mexico. While not known to be
concerned in the transmission of any disease, the habit of the immature stages
of feeding deep in the ears of the host results in intense irritation. Heavily
infested animals, especially calves, are often in poor condition and deaths of
cattle have been attributed to gross infestations of this tick. Several instances
have been reported of the occurrence of nymphs of this species in the ears of
man. Medical attention is usually required to effect the removal of the tick.

Explanation of Terms

Fig. 1

Anterior Projection: The projection of the dorsal body wall at the anterior end.
It may extend horizontally or may be curved ventrad and be continuous with or
separate from the hood.

Anus: The external evidence of the termination of the alimentary tract, located on

the venter back of the coxae. It consists of two eversible flaps enclosed in a

continuous ring or frame.
Apical Ventral Spur: A ventral spur on the distal end of the tarsus in some species.

Figure 53, D and E.
ARTICLE: A distinct articulating portion of a leg or palpus. The sequence of the articles

is indicated by Arabic numerals 1, 2, 3 and 4, 1 being proximal.

Attenuated: To become thin, fine, or less.

Basis Capituli: The basal portion of the capitulum which is movably articulated
with the body and to which the mouth parts are attached. The "basal ring" of
some authors.

Buttons: In the genus Argas, circular elevations found on the integument, each with

a central pit and often with a hair in the pit, and distinguishable from the discs

and "wrinkles."
Barbed Hairs: Tapering hairs with barbs on their sides. The barbs may be numerous

and found on all sides or few in number and present on one side only, and may

be long or very short. Fig. 52 H.

Camerostome: The depression or cavity in which the capitulum lies, and usually
less definite in much engorged specimens.

Capitulum : The movable anterior portion of the body including the mouth parts.
The "head" or "false head" of some authors.

Cheeks: Paired flaps at the sides of the camerostome, which may be either fixed or
movable; the "movable cheeks" of some writers. Fig. 1, C.

Chelicerae: Paired mouth parts for piercing the skin, lying dorsally on the hypostome
and completing the more or less cylindrical mouth parts that are inserted when
the tick feeds.

CLUBBED Hairs: Hairs which do not taper and are terminally enlarged.

Color: The color is greatly affected by ingested blood. At first there is a reddish
tinge resulting from the fresh haemoglobin but in a few hours this changes to
dark gray. During the subsequent process of depletion the changing color of the
blood gradually loses its effect and only the natural color of the body wall
remains. Color is much affected when the specimen is placed in preserving fluid;
also those preserved soon after feeding appear different from those preserved
after having been depleted for weeks or months. Thus, since the color is not
dependable, it is usually not mentioned in describing the Argasidae.

Corona: The apical portion of the hypostome which is differentiated from the
remainder by having very small denticles which may be numerous or few in
number.

Coxae : The sequence of the coxae from anterior to posterior is indicated in Roman
numerals thus, I, II, III, and IV. Fig. 1, B.

Cooley & Kohls: Argasidae of N. America, etc. 5

DENTICLES: The "teeth" on the ventral side of the hypostome. Usually arranged in
parallel, longitudinal rows or files. They may be only slight elevations or they
may be definitely raised, sharp, recurved and with distinct "over-hang" which
makes them more effective in clinging to the host.

Depletion : A term applied to the progressive changes within each stage, between
feedings, in nymphs and adults of the Argasidae. When fully fed, many species
are much distended and as depletion progresses the shape, color, depth of grooves,
elevation of folds, and the appearance of the mammillae are affected making it
difficult to recognize or adequately describe some species. The shape changes
progressively from "rotund" to "flat" without greatly changing the length and
width and the grooves become more pronounced. The dorsal surface may even
become concave. In a few species (notably Antricola coprophtlus) the shape may
become much distorted. In distended specimens the mammillae may become much
flattened and show only as sclerotized spots. Some grooves may disappear com-
pletely and reappear during depletion. The folds usually persist through complete
feeding but sometimes become difficult to see. See "Color.

Discs: Limited areas or spots which are the external evidence of modification of the
structure of the body wall at the points of attachment of the dorso-ventral
muscles. They are usually in a nearly symmetrical pattern and may or may not
be evident on the venter. They vary structurally in the numerous species and
may be superficial or depressed, faint, or distinct. The authors piefer to accept
the term "disc" as appropriate in defining any structural modification of the
body wall at the points of attachment of dorso-ventral muscles. The "obvious
discs," "foveolae," "pits," "scutella," and "patellae" of some authors. Fig. 1, A.

Dorsal Humps: Humps or elevations on the dorsal walls of the articles of the legs
and not including the subapical dorsal protuberance. Fig. 1, D and E.

Dorsal Plate: The unwrinkled spot in or near the middle of the dorsum in some
larvae. Examples are fig. 35, G and fig. 51, G.

Dorsum: The entire dorsal surface in contra-distinction to the venter.

Folds: Constant ridges of the integument found on the venter in Argasidae and, like
grooves, much influenced by the degree of engorgement. Fig. 1, B.

Frame of THE Anus: The continuous circular or oval ring which encloses the
eversible flaps of the anus.

Granulations: Irregular elevations on the surface of the integument in adults of
Otobius, in contra-distinction to tubercles in Anlricola and mammillae in Orm-
thodoros, and "elevations" or wrinkles in Argas. Also used in describing the
very small elevations, similar to micromammillae, found in some species.

GROOVES: Lineal depressions or furrows, mainly on the ventral surface. Their depths
and widths are much influenced by the degree of engorgement. See Fig. 1, B.

Hairs: Hairs found on the body or appendages of the Argasidae have been spoken
of by authors as barbed, clubbed, truncated spines, or simple hairs. In the
descriptions the authors have disregarded for the most part the character of
the hairs or spines because more useful characters are available.

HoOD : The anterior projection of the integument forming in part the walls of the
camerostome, if present. Fig. 1 , A, B and C.

HvPOSTOME : The median ventral mouth part which is immovably attached to the
basis capituli and usually bearing "teeth" or denticles. The dentition is indicated
by numerals either side of a line. Thus 3|3 means that there are three longi-
tudinal files on each half of the hypostome. Measurements are made from the
posthypostomal hairs to the anterior extremity. Fig. 1 , F.

Legs: The sequence of the legs from anterior to posterior is indicated in Roman
numerals, thus, I, II, III, and IV. Fig. 1, D and E.

Mammillae: Elevations of various forms found on the integument in OrnithoJoros
in contra-distinction to the granulations in Otobius, tubercles in Antricola, and
"elevations" or wrinkles in Argas. Previous authors have used the term to
designate the conical or hemispherical elevations found especially in such species

6 American Midland Naturalist Monograph No. 1

as savignVi and iuricaia but with the increased numbers of species now known
it is more appropriate to include under the term all similar structures on the
integument in Orniihodoros. Fig. 1, A.

MEASUREMENTS: All measurements are recorded in millimeters. The articles of the
legs are measured dorsally. The articulations "telescope" on the ventral side but
are fixed or hinged dorsally. The length of the hypostome is measured from the
,posthypostomal hairs to the tip of the hypostome.

Micromammillae : Very small, rounded elevations of nearly uniform size on the sur-
face of the legs, and sometimes on the capitulum as in Orniihodoros lalaje and
O. rudis.

MAMMILLAE

_ DORSO - VENTRAL
GROOVE

SUBAPICAL DORSAL
PROTUBERANCE

TARSUS IV

-DORSAL HUMPS

-METATARSUS IV

SUBAPICAL DORSAL jg

PROTUBERANCE *'

TARSUS I
- DORSAL HUMPS

ME TA

TA R S U S I

CAWEROSTOME
MOUTH PARTS

POSTHYPOSTOMAL
HAIRS

BA5IS CAPITULI

TROCHANTER I

^C&K-

•s aa-wc i&c-v-y: * ^^.".r

r

-FEMUR IV

TROCHANTER IV

ft

Va

_HYPOSTOME

BASIS CAPITULI
' CAMEROSTOME
.PALPUS PALPUS

HYPOSTOME
HOOD p

3ASIS CAPITULI

Fig. 1. Principal characters of Argasidae. A, Dorsal. B, Lateral and ventral. C,
ventral, anterior. D, Leg IV. E, Leg I. F, Capitulum, ventral. G, lateral, anterior.

Cooley 8c Kohls: Argasidae of N. America, etc.

OrNATE: Ornate ticks are common in Dermacenior and Ambhwmma which have a
symmetrical pattern of color usually spoken of as "white" or "gray,- "rose-
tinted," etc., superimposed over the base color of the chitin. In the Argasidae
ornamentation is lacking except in O. coriaceus in which the mammillae are
crowned with "white," "cream," or faint iridescent colors.

PANDURIFORM : Obovate, with a concavity on each side, like a violin.

PosTPALPAL Hairs: A pair of hairs placed posterior to the palpi, more or less in
line with the posthypostomal hairs.

Posthypostomal Hairs : A pair of hairs on the ventral surface at the base of the
hypostome.

Sex Opening: The external evidence of the sexual organs of either sex found on the
ventral median line posterior to the capitulum.

Similar: The sexes are similar if separable only by size and the differences in the
morphology of the sex openings, and dissimilar if there are other morphological
differences of the body or appendages, as in Antricola coprophilus. Adults and
nymphs are similar if separable only by the presence of the mature sex openings
in adults which are absent in nymphs, and dissimilar if, as in Oiobius, the
adults and nymphs have differences in body structures.

Simple Hairs: Smooth, tapering hairs which terminate in a point.

Spiracle: Paired organs, one on each side; the external openings of the respiratory
system which communicate with the tracheae.

Subapical Dorsal Protuberance: The protuberance sometimes present distad of
Haller's organ and when much drawn out produces the bifurcate termination
of the tarsus, mentioned by some authors. It should be distinguished from dorsal
humps. The protuberance appears to be different in origin or purpose from the
dorsal humps. Thus, the species of Argas and those species of Ornithodoros
which have cheeks, and micromammillae on the legs seldom have dorsal humps,
but may have the subapical dorsal protuberance. Fig. 1, D and E..

SuTURAL LlNE: A definite line of cleavage around the periphery in Argas separating
the dorsal and ventral surfaces. Fig. 2, D and Fig. 4, G.

TUBERCLES: The elevations on the integument in Anlricola comparable to the mam-
millae of Ornithodoros.

Truncated Hairs: Hairs which do not taper and are terminally truncated.

VENTER: The entire ventral surface in contra-distinction to the dorsum.

Methods

The characters in the Argasidae are definite and little difficulty should be
encountered in recognizing most species. The methods of study here outlined
are applicable also to various genera of the Ixodidae.

Dissecting microscope. — A primary need is a satisfactory dissecting micro-
scope with low, intermediate, and high power objectives. Very satisfactory
types are: first, one with a stage that can be placed directly on the surface of
the desk; second, one with a heavy base placed well away from the worker and
with a horizontal arm supporting the body of the microscope, thus leaving a
clear working space around and under the instrument.

Lighting. — Adequate lighting is about as necessary as a suitable micro-
scope. Desk and microscope lamps of numerous forms are available but a type
producing a bright beam and on a stand which can be placed well away from
the microscope is particularly useful. It should be possible to adjust the inten-
sity of the light and to avoid heating the specimen. Both aims can be accom-
plished by placing a piece of rice paper or ground glass in the beam of light.

8 American Midland Naturalist Monograph No. 1

Preserving fluid. — It is a distinct advantage to use as a preserving fluid
one that will dry quickly, thus making the surface characters visible with as
little delay as possible. Fluids containing glycerine have the disadvantage or'
leaving a "moist" or "tacky" surface which obscures the details of the surface
parts. However, glycerine can be removed by washing with 70 percent alcohol.
This takes time and subjects the specimen to some danger of injury. Carbon
tetrachloride preserves the true colors in ornate ticks to a remarkable decree
but makes the specimens very hard and brittle. At the Rocky Mountain Labor-
atory 70 percent alcohol is used. This evaporates rapidly and we have not
detected that specimens so preserved are more brittle than those placed in
fluids containing glycerine.

When ticks are kept out of the alcohol for some time and also subjected
to the heat of a light, it is very desirable to put a drop of alcohol on the
specimen occasionally.

Cleaning. — Cleaning should be avoided if possible, but is sometimes neces-
sary, since argasid ticks are often encrusted with coxal fluid which serves as
a mild glue. For cleaning it is useful to have at hand a series of small, artist's
red sable brushes, the tips of which have been cut off at different lengths, thus
affording various degrees of severity for "scrubbing." The cleaning is done
under low magnification with the tick immersed in 70 percent alcohol. Care
must be taken not to break parts or remove hairs.

In examining the hypostome in situ it is frequently desirable to remove
matter from the mouth parts with a soft brush from which the tip has not
been removed.

When especially clean specimens are needed, as for photographing, it is
preferable to use ticks that have recently molted in a clean container.

Posing. — The specimens are removed from 70 percent alcohol, pressed
lightly on soft, absorbent paper (such as is used as a substitute for handker-
chiefs) and are then transferred on a microscopical slide to the field of the
microscope for examination dorsally and ventrally. When special positions are
wanted other aids are used. When working with Dr. Nuttall in England, the
senior author saw him using small lumps of modeling clay or "plasticine"
which could be stuck to a glass slide and shaped to fit the need — a point, a
ridge, a sloping surface, or a groove. Without some such refinement of tech-
nique there is danger of failing to make some characters visible. It is common
practice in museums and laboratories to hold the tick under the microscope
between the points of forceps. This has resulted in injuring valuable specimens.
Because of this practice some types which we have seen have lost part or all
of the appendages. Parts broken off by manipulation should be put in "micro
vials," labelled, and dropped into the larger museum vial along with the dis-
membered specimen.

Dissection. — We have found that it is not always sufficient to study the
hypostome and some other parts in situ. Even with good vision and a satis-
factory microscope it is frequently impossible to see the essential characters
clearly. Dissection and mounting of parts then become necessary.

Cooley & Kohls: Argasidae of N. America, etc.

Dissections are made in a small, Syracuse watch glass (U.S. Bureau of
Plant Industry Model; diameter of 27 millimeters) under a microscope, with
the specimen immersed in alcohol. The watch glasses are improved if the
hottom is coated with collodion to prevent the dulling of sharp instruments.

The hypostome is readily removed from adults and nymphs. The specimen
is carefully held in position with slender forceps. A transverse incision is made
a short distance posterior to the posthypostomal hairs with a very small, sharp,
iris scalpel; also two longitudinal ones, one on either side of the hypostome,
each starting at the point where the palpus meets the base of the hypostome.
This usually permits the removal of a part of the basis capituli with the
hypostome attached, 'but if not yet free, insert the iris knife horizontally
between the hypostome and the stalks of the chelicerae and cut through any
pieces of chitin still attached to the base of the hypostome. It is important
that this final incision be made with the point of the knife held horizontally
to insure that the hypostome will be flat when mounted in balsam and not
raised on one side.

It is necessary to sharpen the scalpel frequently on a marble stone. This
is done under the microscope. For some dissections a blunt, rather than a
pointed, scalpel may be advantageously used.

Care is needed to avoid losing dissected parts, hence the need for the very
small dishes. The parts to be mounted are readily transferred to the next
liquid by the use of a small brush of which most of the tapered portion has
been removed.

In the study of the larvae it is sometimes necessary only to clear and
mount them if the specimens are unfed, but if critical studies are required
the more difficult manipulation of separating the hypostome from the chelic-
erae becomes necessary. This dissection can usually be accomplished best by
removing first only the chelicerae, leaving the hypostome and palpi in posi-
tion. This is often difficult and is usually best done with a fine needle point
and a sharp-pointed, iris scalpel. Finally, the entire capitulum is severed,
cleared in xylol and mounted in balsam with the venter up.

Four percent potassium hydroxide is sometimes used as a clearing agent.
However, critical parts, particularly hypostomes, are sometimes obscured by
gas appearing within the specimen immediately after being placed in balsam.
This may happen even if preparation has included thorough washing to
remove the potassium hydroxide. It may be prevented by passing the specimen
through a series of xylcl-balsam mixtures of increasing density. The first should
be very fluid. This requires about twenty-four hours.

Rearing. — It is sometimes difficult or impossible to identify immature
specimens, especially earily stage nymphs. If alive, such ticks can be reared
through one or more stages as desired, feeding being permitted as necessary.

To facilitate holding living ticks in the laboratory for observation or
ecdysis various facilities have been used such as paper or wooden boxes held
over wet sand, test tubes with cotton plugs placed in covered jars, etc. The

10 American Midland Naturalist Monograph No. 1

essential conditions for keeping the ticks alive for prolonged periods are (a)
containers that permit the maintenance of a higher humidity than is usual in
room atmosphere, (b) sufficient air, and (c) avoidance of over-heating. The
container should be such as to facilitate frequent examinations of the speci-
mens. We have used two methods successfully.

For one method we employ desiccator jars that provide a space for contain-
ing a fluid below and for holding vials of ticks above. A "floor" for the upper
space can be made of cardboard saturated with melted paraffin. Pill boxes
containing ticks direct from the field may be placed temporarily in the upper
chamber. A relative humidity of 92 percent is maintained 'by a saturated
solution of ammonium chloride (NH4 CI) in the lower chamber.- Paper
pill boxes become moist and soft if held in the desiccator jars very long.
On the other hand, ticks in glass vials with cotton plugs may be held indefinite-
ly. The jars are kept at room temperature and out of the sunlight. Mold some-
times develops but causes very little difficulty if the ticks are held in glass vials
and can be further prevented or reduced by using only sterile vials.

Various modifications of the "Hixson jar" are also useful. We have found
the modification described below to be of value in holding a few living ticks in
the laboratory and also in collecting critical material in the field. Hixson (1932)
described and figured a small % inch cork-stoppered shell vial "jar" with wet
sand in the bottom. The tick or ticks are placed in a l/^ inch tube plugged with
cellu-cotton at both ends. The latter is held in a fixed position in the jar by
inserting one end in a central hole through the stopper.

For our purposes we have modified the "jar" by using a wide-mouthed
bottle, 2 inches in diameter by 4 inches tall, with plaster of paris in place of
the sand. The plaster of paris is allowed to set in the bottom of the bottle. The
jar is then placed in a warming oven with the mouth open until the plaster is
thoroughly dry. Six to ten drops of water are added to the plaster to maintain
a moderate degree of humidity. The inside tube should have a diameter of
about 1/g of an inch, should be clean and sterile and should not reach the
plaster of paris.

Critical living specimens may be placed singly in the inner glass tubes and
when ecdysis occurs the molted skin is available. The latter is often of special
value 'because the hypostome or other parts are available for comparison with
the same parts in the subsequent stage. The cast skin of the hypostome may
be placed in absolute alcohol, then transferred to xylol and when mounted in
balsam is nearly as useful as the actual tick of the stage concerned. These small
jars have several advantages when it is desirable to keep living ticks and make
daily observations and records of their development. They are also used in the
field when especially interesting lots of ticks are collected and when shipped
they are put in double mailing tubes.

2 See International Critical Tables, vol. I. p. 67. for information on maintaining
constant humidities.

Cooley dc Kohls: Argasidae of N. America, etc. 11

Family, Argasidae Canestrini, 1890

Non-scutate Ixodoidea with sexual dimorphism slight. Integument of adults
and nymphs leathery, wrinkled, granulated, mammillated or with tubercles.
Capitulum in adults and nymphs either subterminal or distant from the anterior
margin; in larvae subterminal or terminal. Capitulum, especially in depleted
adults and nymphs, in a more or less marked depression (camerostome) . Artic-
ulations of the palpi of all stages free (never fused) . Porose areas absent in both
sexes. Eyes when present placed on the supracoxal folds. Spiracles in adults and
nymphs usually placed anterior to coxae IV. Pulvilli usually rudimentary or
absent in adults and nymphs; sometimes well developed (functional) in larvae.
Nymphal stages plural and the number variable.

Type genus, Argas Latreille, 1796.

The number of genera that should be recognized in the Argasidae has long
been a matter of doubt. Nuttall et al. (1908) and Neumann (1911) recog-
nized Argas and Ormthodoros, although Nuttall stated that "we are by no
means sure that the family Argasidae contains more than one genus, Argas."
Banks (1912) erected the genus Otobius for a species formerly included in
Ormthodoros. This we recognize as valid.

Pocock (1907) advocated the revival of Carios Latreille, 1796 for Argas
vespertdioms, an Old World species associated with bats, because this species
possesses a "conspicuous transverse, lightly curved groove just behind the anus."
As pointed out by Nuttall, this structure is not peculiar to vespertilionis. It is
present in other species also, though modified, and is the character usually
referred to as the transverse postanal groove. However, the species is aberrant.
The capitulum is situated far forward and would be visible from above were it
not for the prominent hood. The sutural line on the margins of the body, a
character used by Neumann (1911) to characterize the genus Argas, is absent.
Yet by the other characters it appears to be clearly an Argas, and until further
information is available, we consider it best to refer the species to this genus.

Also in 1907 Pocock erected the genus Alectorobius for Ormthodoros talaje
on account of the presence of lateral wings (cheeks) on the camerostome. If
Alectorobius were to be reestablished, it would be desirable to find correlating
characters. While talaje and some other known species have cheeks, dorsal
humps on the tarsi, and micromammillae on the legs, certain species lack the
dorsal humps or the micromammillae, or both.

The New World species of Argas and Ornithodoros can best be distin-
guished by the presence of the sutural line in the former, its absence in the
latter. Bedford (1932) made Ormthodoros Koch a synonym of Argas, stating
that he could not consider "having the margin of the body differing in structure
from the rest of the integument" as being of generic importance. He was influ-
enced in his decision by Ornithodoros perengueyi Bedford and Hewitt, 1925.
In this species, as well as in O. dunm Matheson, 1935 and O. stageri Cooley
and Kohls, 1941, the flattened margins persist even in well fed specimens but
the sutural line is absent. While these species serve to show that the genera

12 American Midland Naturalist Monograph No. 1

grade into each other, the great majority of the species can be assigned generi-
cally with little or no difficulty.

We accept as of good standing the following genera:

1. Argas Latreille, 1795 3. Otobius Banks. 1912

2. Ornithodoros Koch, 1844 4. Anlricola Cooley and Kohls, 1942

Key to the Genera

1 . With a definite sutural line separating dorsal and ventral surfaces (see fig. 4, G)

Argas

Lacking a definite sutural line separating dorsal and ventral surfaces 2

2. Nymphs with integument beset with spines, hypostome well developed; adults with

integument granular, hypostome vestigial (see fig. 6, C) Oiobius

Integument of adults and nymphs essentially alike, mammillated or tuberculated,
and lacking spines; hypostome of various forms in nymphs and adults but not
vestigial 3

3. Hypostome broad at the base and scoop-like (see fig. 53, B) (Associated with

bats.) Antricola

Hypostome of various forms but never scoop-like (Associated with various classes
of animals and including bats.) Ornithodoros

Cooley & Kohls: Argasidae of N. America, etc. 13

Genus Argas Latreille, 1796

1796. Argas Latreille, original description, p. 18. The generic synonymy to 1850 is
reviewed in detail in Oudemans "Kritisch Histonsch Overzicht del Acarologie,"
part 2, pp. 135-137 (1929) and part 3, B, pp. 746-755 (1936). See also Nuttall
el al. (1908) pp. 4-5.

1932. Argas Latreille: Bedford, p. 275. OrnillwJoros made a synonym of Argas.

1934. Argas Latreille: Bedford, p. 60.

1936. Argas Latreille: Brumpt, p. 1186.

Sexes similar; nymphs and adults similar.

Body flattened, dorsal and ventral surfaces about equal in area; margin
distinctly flattened, made up of radial striae or quadrangular plates. Sutural
line present. Flattened margins not obliterated even when tick is fully fed.
Capitulum either distant from or near the anterior border. Integument
leathery, minutely wrinkled in folds of many shapes often intermingled with
small, rounded "buttons" each with a pit on top and often bearing a hair in
the pit. Discs present on both dorsal and ventral surfaces and placed in more
or less radial lines. Eyes absent.

Genotype, A cams reflexus Fabricius, 1794, as designated by Latreille
(1802). Nuttall et al. while admitting that reflexus had been the accepted type
preferred to take persicus as the type because of its world-wide distribution
and its having been better studied. We regard these reasons as being insuffi-
cient for changing the genotype.

Two species occur in North America, reflexus (Fabricius) 1794 (see dis-
cussion page 16) and persicus (Oken), 1818. A third species, A. brevipes
Banks 1908, described from Arizona, must remain in doubt since no specimens
so labelled 'by Banks are available and the description is inadequate.

Key to Species of Argas
(Adults and Nymphs)

Flattened margins striate; postpalpal hairs absent; hypostome apically rounded

reflexus (p. 14)

Flattened margins having quadrangular plates; post-palpal hairs present; hypostome

apically notched persicus (p. 17)

14 American Midland Naturalist Monograph No. 1

Argas REFLEXUS (Fabricius), 1794

Plate 1, Figs. 2 and 3

1793. Acarus columbarum Shaw, in Shaw and Nodder, original description, 4, pi. 128.

(nomen nudum).
1794 Acarus reflexus Fabricius, p. 426.
1802. Argas reflexus (Fabricius) : Latreille, p. 66.

1804. Rhpnchoprion columbae Hermann, p. 69.

1805. Ixodes reflexus and Ixodes colmbae Fabricius, p. 353 and 356.

1815. Argas marginaius Oken, p. 402.

1816. Rhvmchoprion marginatum Olfers, p. 75.

1827. Ixodes hispanus and Ixodes espagnol. Brebisson, p. 267.

1828. Acarus marginatus. Guerin-Meneville, p. 300.

1829. Argas reflexus. Latreille, p. 289.

1896. Argas magnus Neumann, p. 14. Degraded to Argas reflexus var. magmis,

Neumann. '1905. p. 239.
1929. Argas columbarum (Shaw): Oudemans, part 2, p. 138.
1936. Argas reflexus (Fabricius): Brumpt pp. 1187-1188.

ADULT

Body. — Oval, often distinctly narrower in front. Margin irregularly striate
with the striae varying in width in different specimens. Margin slightly turned
up (whence reflexus). Often larger than persicus and may reach 9.0 x 5.0
(Nuttall et al. 1908). Specimens may be as small as 6.25 x 3.75.

Integument. — Integument on dorsum and venter raised into irregular,
sinuous wrinkles which may be short or long. Nuttall et al. (1908) state that
the integument is much more finely wrinkled than in persicus. Specimens from
the Americas show the wrinkles to be coarser.

Discs. — Variable; may 'be either large and numerous or small and few;
when numerous arranged radially.

Buttons. — Absent or few in number, often each with a short, fine hair
arising from the pit.

Legs. — Moderate in length and size. Surface irregular — nearly granular.
Subapical dorsal protuberances well developed, about equal on all the legs.
Length of tarsus I (in larger specimens), 0.78; metatarsus, 0.78. Length of
tarsus IV, 0.96; metatarsus, 0.93.

Coxae. — Coxae I and II separated; all others contiguous. (Nuttall et al
(1908) state that coxae I and II are contiguous.)

Folds. — Coxal and supracoxal folds present; all other folds absent.

Capitulum. — Removed from the anterior margin by about its own length
(including the palpi). Basis capituli nearly twice as wide as long; lateral
margin with a group of short, fine, erect hairs at about the middle. Entire
length of palpal article 1 in contact with and overlapping the base of the
hypostome as a knife-edge flange. Postpalpal hairs absent.

Hypostome. — Mildly tapering; apex rounded. The larger denticles
arranged -/•_> with about 3 or 4 in each file. Numerous very fine denticles in
the middle arranged in diagonal rows (more numerous than in persicus)

Cooley & Kohls: Argasidae of N. America, etc.

15

Corona with only a very few fine denticles. Long posthypostomal hairs are
present. Length 0.30 to 0.39.

Camerostome. — Short and indefinite.

Sexual opening. — Placed at the level of the intervals between coxae I
and II.

Anus. — In an elliptical pattern placed nearly central on the venter.

This species is readily separated from persicus by its striated margins, its
well developed subapical dorsal protuberances on all legs, the absence of the
apical notch on the hypostome, and absence of postpalpal hairs.

DISTRIBUTION AND HOSTS

In the Old World Nuttall et al. (1908) record this species from England,
France, Germany, Italy, Russia, Rumania, and Algeria. Because of its associa-
tion with pigeons, Colamba domestka, it is commonly known as the pigeon

B

Fig. 2. Argas reflexus (Fabricius). A, Margin of adult or nymph. B, Leg I of
female. C, Leg IV of female. D, Lateral view of edge showing sutural line. E, Hypos-
tome of female. F, Hypostome of male.

16

American Midland Naturalist Monograph No. 1

tick. Other hosts mentioned are Gallus domesticus, man, and a single record
of larvae from Equus caballus reported by Starcovici in Rumania.

As for the New World, Dr. J. Bequaert has sent us specimens from Bogota,
Colombia, South Amenca, collected in chicken coops. Neumann (1911)
recorded A. reflexus magnus from California without mention of host. Osborn
(1896) stated that, "It is common, I believe as far north as St. Louis." How-
ever, no records substantiating this statement have appeared. The only United
States records we have are the following:

California. 17018, near nest of California condor, Gymnogyps caliform-
anus, September 18, 1939, and 1787, May 24, 1940, 'both lots from Santa
Paula Canyon, Ventura County, 41 nymphs and adults (Thomas F. Kelley);
17395, "nest in cave," April 8, 1939, Mt. Diablo, Contra Costa County, 1
male (D. E. Howell); 17556, nesting hole of Inyo screech owl, Otus asio
inyoensis, November 1940, Inyo County, 1 nymph, also 5 nymphs of Argas
persicus (Kenneth E. Stager).

Montana. 15195, October 1938, Haxby, 1 female. This specimen had been
sent to Dr. H. B. Mills, State Entomologist, with the information that it and
two others were "found in the bedroom on my bed." During the summer, 10
or 15 swallow nests were built under the eaves of this bedroom. It appears
likely that the swallows were the source of the infestation.

Assignment of the United States specimens to reflexus is made with some
hesitation. We believe it advisable to refer the specimens listed to this species
until more information is available. While they are smaller than Old World
and South American specimens and lack the pronounced reflexed margins of
the latter, they agree very closely in other morphological characteristics. How-
ever, it is surprising that there are no definite records of the species from
pigeons and other domestic birds in this country.

Fig. 3. Distribution of Argas reflexus (Fabricius).

Cooley & Kohls: Argasidae of N. America, etc.

17

Argas persicus (Oken), 1818

Plate I, Figs. 4 and 5

1818. Rhunchoprion persicum Oken, original description, p. 1568.

1823. Argas persicus Fischer de Waldheim, p. 282.

1844. Argas mauritianus Guerin-Meneville, p. 17.

1844. Argas chinche Goudot: Gervais, in Walckenaer, p. 462.

1844. Argas miniatus Koch, p. 219.

1872. Argas americana Packard, p. 740.

1891. Argas sanchezi Alfred Duges, p. 20.

1893. Argas radiahis Railliet, p. 718; for americana Packard, preoccupied.

1901. Argas miniatus Koch: Neumann, p. 255, " americana Packard, sanche:i Duges

and radiahis Railliet, synonyms. P. 344, - = chinche Goudot, synonym.
1901. Argas persicus Fischer: Neumann, p. 253, 256, — Argas mauritianus Guerin-

Meneville, synonym.
1905. Argas persicus var. miniatus: Neumann, pp. 240-241.
1908. Argas persicus (Oken) : Nuttall, el al., p. 9 = Argas persicus var. miniatus,

synonym.
1932. Argas persicus (Oken): Bedford, p. 281.
1934. Argas persicus (Oken): Bedford, pp. 65-69.
1936. Argas persicus (Oken): Brumpt, pp. 1189-1192.

ADULT

Body. — Usually oval, wider behind; sometimes almost elliptical. Margins
composed of irregular quadrangular plates or cells which often have one or
more circular pits. In some specimens a very small, short, fine hair is visible
arising frcm some of the pits. Like reflexus this species varies greatly in
size. Nuttall et al. give size ranging from 4.0 x 2.5 to 12.7 long. Specimens
in the collections of the Rocky Mountain Laboratory show sizes within the
range published.

Integument. — Surface of dorsum and venter with numerous rounded or
sinuous, shining wrinkles. In some specimens the sinuous elevations are unusu-
ally long.

Discs. — Numerous, large, variable in size, oval or circular, and radially
arranged. Other discs not radially arranged are found in the submarginal areas.

Buttons. — Usually numerous but faint and sparse in some specimens.

Legs. — Moderate in length and size. Some specimens from Southern
United States have the legs short and small. Surface irregular with hairs short
and fine. Subapical dorsal protuberances mild or absent on tarsi I to III;
absent on tarsus IV. Length of tarsus I, 0.57 to 0.75; metatarsus, 0.51 to 0.81.
Length of tarsus IV, 0.72 to 0.81; metatarsus, 0.66 to 0.78.

Coxae. — Coxae I and II well separated; all others contiguous; surfaces
longitudinally wrinkled.

Folds. — Coxal and supracoxal folds present.

Capitulum. — Removed from the anterior margin of the body by about the

18 American Midland Naturalist Monograph No. 1

length of the capitulum including the palpi. Basis capituli with its width
greater than its length; surface transversely wrinkled. Postpalpal hairs present,
about in line with the posthypostomal hairs. Entire length of article 1 of the
palpus in contact with the hypostome and with a knife-edge flange overlap-
ping it.

Hypostome. — Sides a little tapering; apex notched. Larger denticles few in
number and arranged -/._>; smaller ones, in middle portion, arranged 3'3-
Corona with very small denticles. Length of female hypostome 0.315 to 0.36;
male about 0.30.

Camerostome. — Short, indefinite, and affording little protection for the
mouth parts.

Sexual opening. — At the level of the intervals betweeen coxae I and II.
Anus. — In an elliptical pattern; placed nearly central.

This species is readily separated from reflexus by having the hypo-
stome terminally notched, the margins with quadrangular cells in place of
striae, and postpalpal hairs present.

GEOGRAPHICAL DISTRIBUTION

Argas persicus occurs in Europe, Asia, Africa, America, and Australia.
In general it is restricted to the warm, dry regions.

Bishopp (1927) stated that

The present distribution of the tick in the United States may be said to extend
throughout the western three-fourths of Texas, from approximately the longitude or
Dallas, westward; the southern half of New Mexico and Arizona; southwestern
Oklahoma; three-fourths of southwestern California; and the major part of Florida.
During the last few years it appears that the pest has extended ils range northward
in the Sacramento Valley of California a considerable distance, as R. W. Wells has
found it to be firmly established in Shasta County. * * * The tick has undoubtedly
been shipped with fowls into many parts of the United States and specimens have been
taken in States as far removed from the normally infested areas as Iowa. It has not
established itself generally, however, in the more humid and cooler parts of the
country. It has been thought that the tick would not become a pest in such regions
and in the higher mountain areas, but its continued spread indicates that it has consider-
able adaptability, and that it will ultimately infest a much larger part of the country
than it does at present.

The map showing the distribution of Argas persicus is after Bishopp
(1927) with the following records added:

California. 17556, nesting hole of Inyo screech owl, Olus asio inyioensis, Inyo
County, December 1940, 5 nymphs, also 1 nymph of Argas columbarum (Kenneth E.
Stager).

Georgia. 42331 B. A. I., Callus clomestitus, chicken house, Feb. 18, 1936, Atlanta,
16 specimens (W. R. Baynes).

Louisiana. 30307 B. A. I., host not stated, Oct. 27, 1940, Jeanerette, few speci-
mens (C. W. Rees).

Nevada. Many specimens without host or date, Reno (S. B. Doten). Record from
U. S. National Museum.

Cooley &. Kohls: Argasidae of N. America, etc.

19

Utah. Many specimens, said to be killing chickens near St. George, 1936 (E. W.
Davis). Record from U. S. National Museum.

Canada. 17496, golden-crowned sparrow, Zorwtrichia coronata (Pallas), May 2,
1931, Vancouver, B.C.. 4 nymphs (Hearle. 1938).

In Mexico, Hoffman (1930) stated:

It is found in all of the hot and dry States of the North and extending throughout
the regions consisting of the Central Plateau toward the south and west to the Pacific
Coast. In the more humid zone along the Gulf Coast and in the humid districts of the
south it occurs only exceptionally and then as a result of fowls being brought in from
infested places. Apparently it is unable to survive in these regions. It easily adapts
itself to the chicken houses which are protected against the cold of our Central Plateau,
for example, I have seen infested chicken houses at Tlalpam, D. F., which is some
2300 meters above sea level. (Translation.)

According to Dunn (1923) the species is "very abundant throughout
Panama, the majority of chicken houses and other places where fowls com-
monly roost in the cities of Panama and Colon, villages in the Canal Zone
and native villages in the interior being usually infested with them."

151

M8S®

E

G

Fig. 4. Argas persicus (Oken). A. Margin of adult or nymph. B, Leg I of female.
C. Leg IV of female. D, Hypostome of female. F, Hypostome of male. F, Capitulum
of larva, ventral view. G, Lateral view of edge showing sutural line.

20 American Midland Naturalist Monograph No. 1

In the West Indies the species has been found at Trinidad and at Antigua.

Vigueras (1934) stated that Argas persicus has been known in Cuba for
many years, and listed the following localities: Provinces of Havana, Matan-
zas and Santa Clara.

Koch's type of A. miniatus came from Demerara, British Guiana.

HOSTS

Argas persicus is preeminently a fowl tick. It attacks practically all species
of domestic fowls, but the chicken appears to be the preferred host. It has
been found in limited numbers on a wide variety of wild brids including quail,
wild doves, wild turkeys, vultures, golden-crowned sparrow (Z.onotricbia
coronata), and Inyo screech owl (Otus asio inyoensis) . It is said to attack
man commonly in Persia, producing serious symptoms. Hoffman (1930)
stated that in Mexico it attacks man only exceptionally, with more or less dis-
agreeable local effects, and as a rule only in houses where it cannot reach its
natural hosts.

Economic losses occasioned by this tick are considerable. Fowls are weak-
ened through loss of blood and annoyance and when attacked by large
numbers the effects are sufficient to result in death.

Fig. 5. Distribution of Argas persicus in the United States and Canada. Adapted
from Bishopp (1927), with additions.

Cooley & Kohls: Argasidae of N. America, etc. 21

Genus OTOBIUS Banks, 1912

/912 Otobius Banks, p. 99, original description.

Adults and nymphs dissimilar and with sexes similar. Adults with the
integument granulated; nymphs striated and with spines. Without change of
pattern of the integument at the sides. Capitulum distant from the anterior
margin in adults; near the margin in nymph. Hood and eyes absent. Hypo-
stome of nymphs well developed; vestigial in adults.

Genotype: Argas megnini Duges, 1884, original monotype.

Key to Species of Otobius
Adults

The numerous pits on the dorsum separated by a distance of twice or more the

diameter of one pit megnini (p. 21)

The numerous pits on the dorsum separated by a distance of the diameter or less,

of one pit lagophilus (p. 32)

Nymphs

Integument with numerous heavy spines anteriorly and lighter spines posteriorly;

hypostome with denticles 4|4; spiracles conical megnini (p. 23)

Integument with numerous spines all of one size; hypostome with denticles 3|3;

spiracles convex lagophilus (p. 34)

Otobius megnini (Duges), 1884

Plate 2, Fig. 6, 7, 8. 9 and 10

1884. Argas megnini Duges, original description, pp. 197-198, with figures.

1885. Argas megnini Alf. Duges: Megnin, redescribed, pp. 466, 472-475, with figures.
1893. "Argas americana Packard": Townsend, p. 50.

1895 Rhhnchoprium spinosum G. Marx, figures only, p. 199, Marx obituary notice.

1896. Orniihodoros megnini (Alf. Duges) : Neumann, redescribed pp. 42-44.

1901. Omithodoros megnini (Duges): Salmon and Stiles, redescribed, pp. 408-414,

with figures.
1908. Orniihodoros megnini Duges: Banks, redescribed, p. 17, with figures.
1908. Omithodoros megnini (Duges) 1883: Nuttall, Warburton, Cooper and Robinson,

redescribed, pp. 71-77, with figures.

1911. Omithodoros megnini (Alf. Dug.): Neumann, p. 125.

1912. Otobius megnini Duges: Banks, new genus proposed, p. 99.

1930. Omithodoros (Otobius) megnini Duges, 1882: Hoffman, redescribed, pp. 151 —

155, with figures.
1932. Argas megnini Duges: Bedford, p. 280.
1934. Argas megnini Duges: Bedford, pp. 77-81.
1936. Omithodorus megnini (A. Duges): Brumpt, p. 1209.

ADULT

Body. — Panduriform, rounded behind and slightly attenuated anteriorly;
broadest at legs II and III, constricted just behind leg IV. Both Salmon and
Stiles (1901) and Nuttall et at. (1908) give as the size of the female, 6.0 x
4.0 to 5.0 x 3.0. In the collections of the Rocky Mountain Laboratory we have
females as large as 8.25 x 6.00. Males are a little smaller. Specimens of adults
preserved in alcohol are enclosed by a thin, translucent covering which is easily

22

American Midland Naturalist Monograph No. 1

removed in part by the use of a needle point. The true character of the integu-
ment is better seen after the removal of the covering.

Mammillae. — True mammillae as denned in Ormthodoros by authors
appear to be absent. Dorsal and ventral surfaces have the integument granu-
lated, and with numerous circular depressions, each depression with a central
tubercle.

Discs. — The discs are easily overlooked but are present as small, slightly

iu Vu \^

Fig. 6. Otobius megnini (Duges). A, Hypostome of first-stage nymph. B. Hypostome
of second-stage nymph. C, Hypostome of adult. D, Capitulum of nymph, ventral view.
E, Leg I of adult. F, Leg IV of adult. G, Leg I of nymph. H, Leg IV of nymph.
I, Capitulum of adult, ventral view.

Cooley & Kohls: Argasidae of N. America, etc. 23

depressed areas in a symmetrical pattern; the individual discs have their
surfaces only a little changed over the general granulated surface, and are less
distinct on the ventral side.

Hairs. — A few very fine hairs, easily overlooked, arise from some of the
tubercles and are more apparent at the anterior end.

Legs. — Short and heavy. Tarsi II, III, and IV with the subapical dorsal
protuberance moderate, negligible on I. Length of tarsus I about 0.48; meta-
tarsus about 0.48. Length of tarsus IV about 0.72; metatarsus, about 0.69.
Hairs on the legs few, short, inconspicuous.

Coxae. — Each coxa with an elongated smooth sclerite which, together with
deep invaginations on the posterior side mark the position of the coxa. Coxa
IV also has such an invagination on the anterior side.

Hood and Camerostome. — Hood very short and broad — a curved eleva-
tion anterior to the mouth parts. Camerostome lined with numerous fine, long
hairs.

Capitulum. — Basis very broad and short, curved, approaching a reniform
shape with the convexity behind.

Palpi. — Short and heavy; article 1 swollen ventrally and laterally; hairs on
the palpi fine and long.

Hypostome. — Vestigial in marked contrast with the well developed hypo-
stome of the nymph. Short, broad, with the sides converging anteriorly, con-
cave dorsally and convex ventrally; without denticles and with the apical
margin curved and thin. Length in the female 0.135.

Folds. — Coxal and supracoxal folds are present but in well engorged
specimens the coxal fold is not much in evidence.

Grooves. — A short postanal groove present near the posterior end of the
body. Median postanal groove present only anterior to the postanal groove,
faint. Other grooves are negligible or absent.

Sexual opening. — At the level of the posterior ends of coxae I.

Spiracle. — Circular, mildly convex.

Eyes. — Absent.

Anus. — Circular, very small.

SECOND NYMPH

This tick is ordinarily seen only in the second nymphal stage, which is the
stage commonly found in the ears of domestic animals, and is the stage in
which the species is most easily distinguished from other ticks.

Body. — When fully fed a little larger than that of the adult. Shape of fed
specimens much as in the adult but with the lateral constrictions a little less

24

American Midland Naturalist Monograph No. 1

pronounced. Unfed specimens much narrowed posteriorly, constriction begin-
ning at about the fourth pair of legs; broadest at about the third pair of legs;
rounded anteriorly.

Discs. — The integumental markings of the nymph are very different from

Fig. 7. Otobius megnini (Duges) nymph, only a little fed; dorsal and ventral views,
showing spines. After Marx (1895).

M r

V

,4

M i i vv3

B

Fig. 8. Otobius megnini (Duges). A, Nymph, unfed, ventral view. B, Nymph.
"Portion of skin, showing spines and hairs. Greatly enlarged." From Salmon and Stiles
(1901).

Cooley & Kohls: Argasidae of N. America, etc.

those of the adult. The entire surface, shining and with fine reticulations or
striae which are continuous over depressed areas which are the counterpart of
discs. These disc areas lack spines.

Spines. — Spines conspicuous, of two kinds (see figures 7 and 8, B) ; those
with heavier bases confined to an anterior dorsal crescentic area which extends
over the sides and backward to a short distance in front of the spiracle and is
continuous with the anterior ventral area which reaches posteriorly about to
the anus; the more slender spines occupy the posterior dorsal, lateral and ven-
tral areas not detailed above. The line of demarcation between the two kinds
of spines is definite. Spines absent in the area surrounding the capitulum.

Legs. — Short and heavy. Tarsi II, III, and IV with the subapical dorsal
protuberance absent or very small; absent on I. Length of tarsus I, 0.48; meta-
tarsus, 0.42. Length of tarsus IV, 0.6; metatarsus, 0.6. Hairs on the legs a
little longer than those on the legs of the adults.

Coxae. — Indefinite; trochanters of the legs arise from a circular opening in
the body wall which is modified only by a short, V-shaped sclerite in the usual
position of the coxa.

Spiracle. — Conical.

Folds and Grooves. — Coxal and supracoxal folds are faint or absent in the
nymphs. True grooves are absent.

Capitulum. — Subrectangular, about as broad as long, ventrally tumescent;
surface smooth and shining; a few hairs at the sides in front, and with a group
of short, heavy spines on each side behind.

Hood and Camerostome. — Absent.

Hypostome. — Large, tapering, and with long, sharp denticles arranged 4 4,
with about 8 in each file. Denticles about as large and prominent in basal as
in apical portion, with no fine denticles in the corona. Posthypostomal hairs
absent. Length about 0.36.

FIRST NYMPH

The first nymphal stage is very much like the newly emerged second stage,
but is smaller, has more slender legs and the hypostome measures only 0.195.
Posthypostomal hairs absent. (See also Brumpt [1936] p. 648).

LARVA

Unfed larva measures 0.66 from tip of hypostome to posterior extremity.
Body oval; two pairs of hemispherical, ocellus-like eyes present. Integument
thin, striated, and with a few bristle-like hairs arranged symmetrically. Capitu-
lum visible in both dorsal and ventral views; hypostome and palpi very long.
Hypostome with the denticles arranged -/■_>. Palpi with articles 2 and 3 about
equal and with 1 and 4 short. Legs long; stalk of pulvillus long; pulvillus

26

American Midland Naturalist Monograph No. 1

small. Fed larva much distended, broader in front; the capitulum also dis-
tended as a conical anterior projection. Length 4.00, width 2.5.

Salmon and Stiles (1901), p. 410, state:

Some of our specimens of this species show a pupa-like stage. They are about 4 mm.

Fig. 9. Otobius megnini (Duges). A, larva, dorsal view. B, larva, ventral view. C,
Capitulum, dorsal view. D, Capitulum, ventral view. From Salmon and Stiles (1901).

Cooley & Kohls: Argasidae of N. America, etc. 27

long by about 2 mm. broad, rounded, white pyriform structures with one end elongate. . .

We have not seen this stage alive, but from the general structure it is apparent that
in changing from the hexapod to the octopod stage O. megnini passes a resting pupa-like
stage. This is by no means surprising, since the differences between the hexapod and the
octopod stages of the Argasidae are far greater than those between the corresponding
stages of the Ixodidae. This is apparently the stage which Townsend (1893) interpreted
as an egg.

Brumpt (1936), p. 647, gives the following:

From the cat (693 XV) I collected two types of nymphs (1st and 2d. n.). The
engorged nymphs in the last stage dropped on the 38th day.

The last molt, which produces the adults, is effected in the exterior environment.
From the hexapod larva to the adult according to me there certainly are three molts and
not two as claimed by others.

In February, 1940, many larvae of megnini from females, Douglas, Ari-
zona, 16128, were placed in the ears of rabbits and soon engorged. Some of
the engorged larvae were removed and held for molting while others were left
attached. Fourteen days after the rabbit was infested with larvae, a comparison
was made of the nymphs resulting from the engorged larvae removed from the
host and the nymphs that were now present on the rabbit. The two nymphs
were found to be distinctly different, showing that the nymphs on the host had
undergone a molt; thus our findings are in conformity with those of Dr.
Brumpt. The hypostome of the first nymph measured 0.195, the coxae were
smaller, and the legs not as heavy. The hypostome of the second nymph meas-
ured 0.33, the coxae were larger, and the legs heavier.

Otobius megnini has only three molts in its development from egg to adult
1 larval and 2 nymphal stages. The larva becomes very large during its
feeding and is what was spoken of as the "pupa-like stage" by Salmon and
Stiles (1901, p. 410). Whether nymph 1 ever feeds is not evident.

Nymph 2 is the stage usually seen and from it has arisen the name
"spinose ear tick." This stage is small when it emerges but during its pro-
longed feeding it becomes very large and it appears that its striated integument
is correlated with its phenomenal extensibility. When ecdysis takes place there
is a complete change of character of the integument, the spines are lost and
the large hypostome is replaced by a vestigial one.

The adult does not feed.

HOSTS

O. megnini appears to be largely restricted to domestic animals and most
of the records are from cattle and horses. Other animals known to be attacked
include mules, asses, sheep, goats, hogs, dogs, cats, coyotes, deer, mountain
sheep (Oris canadensis) , cottontail rabbit (Sylvilagus sp.), jackrabbit (Lepus
californicus walla walla), and ostrich. There are several records of occurrence
of nymphs in the ears of man.

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30

American Midland Naturalist Monograph No. 1

DISTRIBUTION

Otobius megnini (Duges) was originally described from Guanajuato,

Mexico, and is common in that country as reported by Valdez (1923),

Hooker, Bishopp and Wood (1912), and Hoffman (1930), who states

(translation) :

O. megnini is a typical inhabitant of our arid and semi-arid regions of the centra!
plateau of the southern part of the country and the Pacific Coast. In the central plateaus
I have found the species at an altitude of 2500 meters above the sea. At these elevations
they apparently develop from the eggs only in the hot months following the rainy season.

Fig. 10. Distribution of Otobius megnini (Duges).

Cooley & Kohls : Argasidae of N. America, etc. 31

In the humid regions the species does not exist since the larva is quite sensitive to
humidity.

While the species may occur in Central America, we have no records from
that region.

It has been reported in Argentina by Aragao (1935), in Brazil by Pinto
(1930), and there are specimens from Chile in the Bureau of Animal Indus-
try (40809). In South Africa it has been reported by Bedford (1912), and
Kingston (1936) reported it from India in the ear of a gelding that had been
bred and reared in Queensland, Australia.

In the United States, O. megnim appears to be rather common in parts of
Texas, New Mexico, Arizona, and southern California. We have never made
an effort to determine the definitely infested areas in the United States but
from the extensive records of occurrence in Hooker, Bishopp and Wood
(1912), the Bureau of Animal Industry, the United States National Muse-
um, as well as from the records of the Rocky Mountain Laboratory, we are
able to map the occurrence of this species in this country in a general way.
(See figure 10.) The circular spots on the map indicate records from the
Rocky Mountain Laboratory, Bureau of Animal Industry, and the United
States National Museum. The species because of its long feeding period is
particularly subject to introduction into new areas even over long distances.
Occurrences in Northern, Central, and Eastern States are probably the results
of livestock shipments.

We know that it has persisted for at least several years in eastern Mon-
tana. On April 27, 1916, Cooley wrote to Mr. Al Young, Kirby, Big Horn
County, and received his reply dated May 5, 1916, as follows:

In reply to yours of April 27 concerning ticks, in February and March 1912 a
great many cattle died in this neighborhood. While skinning these we found the ear
tick. We talked the matter (over) with our neighbors and one man said he had seen
them two years before. I only have his statement for it. But since that time (1912) I
know the ticks have been in the ears of our cattle. Ours are native cattle. The ticks
I am sending today were taken from the ear of one of my cows. Respectfully yours.

32 American Midland Naturalist Monograph No. 1

Otobius lagophilus Cooley and Kohls, 1940
Plate 3, Figs. 11 and 12

1940. Otobius lagophilus, Cooley and Kohls; original description, p. 928-933, with
figures.

ADULT

Body. — Rounded on both ends and but little constricted at the sides just
behind legs IV (less panduriform than in megnini); widest at legs II and III.
Length of female, 5.40 to 6.25; width, 3.60 to 4.00. Length of male, 4.75 to
5.00; width, 2.90 to 3.50.

Integument:! on both dorsal and ventral surfaces granular with numerous
intermingled circular depressions or pits and each pit with a single small
central elevation. Very short, fine hairs present on the elevations in the circular
depressions, less apparent on the venter except near the mouth parts.

Discs. — Indefinite, yet evident as symmetrical patterns of small depres-
sions, the floors of which are irregular.

Legs. — Short, moderately heavy, and with hairs few and small. All tarsi
with moderate subapical dorsal protuberances, more pronounced on tarsus IV.
Length of tarsus I, 0.45; metatarsus, 0.39. Length of tarsus IV, 0.54; meta-
tarsus, 0.525.

Coxae. — Coxae III and IV contiguous, the others separated. There are
deep invaginations between the coxae which are shown by dissection to be the
surface indications of large apodemes. Coxae I, II, and III each with an elon-
gated, smooth sclerite.

Hood and Camerostome. — Definite hood and camerostome are not appar-
ent, though there is a moderate swelling ventrally excavated, just anterior to
the mouth parts.

Capitulum. — Basis capituli very broid and short, curved, approaching a
reniform shape with the convex border behind. Surface irregular, with fine
hairs at the sides near the palpi and two groups, one on each side, near the
posterior margin. Palpi moderately heavy, with article 1 a little more swollen
than the others. Palpal hairs fine, moderate in length.

Hypostome. — Vestigial, in marked contrast with the well developed hypo-
stome of the nymph. Broad, short, tapering, without denticles, concave dotsal-
ly, convex ventrally; bluntly rounded or partly bilobed apically. Length about
0.10.

Folds. — Coxal and supracoxal folds present though less in evidence in well
engorged specimens.

Grooves. — A short, transverse postanal groove present near the posterior
end. Median postanal groove faint. All other grooves are absent or negligible.

3 Specimens of adults and nymphs preserved in alcohol are enclosed in a brittle,
thin, translucent covering which when dried is easily removed with a needle. The true
character of the integument of the body wall is better seen after this covering is removed.

Cooley & Kohls: Argasidae of N. America, etc.

33

Sexual opening. — In line with the posterior ends of coxae I.

Spiracle. — Ovate, with surface convex.

Eyes. — Absent.

Anus. — Small, nearly circular.

J

wj I,

/>!■/

l

Fig. II. Otobius lagophilus Cooley and Kohls. A, Hypostome of nymph. B, Hypos-
tome of adult. C, Leg I of adult. D, Leg IV of adult. E. Leg I of nymph. F. Leg IV
of nypmh. G, Spines from anterior portion of the nymph. H, Capitulum of nymph,
ventral view. I, Capitulum of adult, ventral view.

34 American Midland Naturalist Monograph No. 1

NYMPH

Shape and size about as in the adult.

The integumental markings of the nymph very different from those of the
adult. Entire surface, dorsal and ventral, smooth and shining, with fine reticu-
lations and transverse striae, similar to the body wall of larval Argasidae.

Spines. — Spines or hairs are present over the entire surface of the body
except in the area immediately surrounding the mouth parts; more abundant
and longer at the anterior end; progressively smaller toward the posterior end;
less numerous on the ventral surface.

Discs. — Definite discs are absent but corresponding depressions are some-
times evident. These depressions are free of spines and in some a few fine
punctations are visible.

Legs. — Short and moderately heavy. Subapical protuberances absent or
small on tarsi I, II, and III, but distinct on IV. Length of tarsus I, 0.36;
metatarsus, 0.30. Length of tarsus IV, 0.45; metatarsus, 0.45.

Coxae. — Present as inconspicuous sclerites.

Hood and Camerostome. — Absent.

Capitulum. — In ventral view the capitulum is in a depression formed by a
circular tumescence which makes it difficult to get a true impression of the
shape of the basis capituli unless it is dissected out. Broader than long. Palpi
moderately heavy (more slender than in megnini), and with article 1 lacking
the distinct ventral swelling found in the adult. Hairs small and few in
number.

Hypostome. — Large, with sides nearly parallel, denticles long and sharp,
in a sl$ arrangement with the denticles about equally long in the basal and
apical regions; denticles in the corona absent or few in number. Marginal den-
ticles absent in the subapical region. Length, 0.30.

Folds. — Coxal and supracoxal folds are faint or absent.

Grooves. — True grooves are absent but the preanal and median postanal
grooves are indicated by shallow, elongated depressions.

Spiracle. — Circular, mildly convex.

This species resembles the well-known spinose ear tick, O. megnini, but is
readily separated by the following characters: its smaller size; the heavy V-
shaped spines found on the anterior surfaces in megnini are replaced in
lagophilus by slender spines which are the same as those on the posterior parts;
denticles on the hypostome in a 3/3 pattern instead of 4,4; legs more slender;
spiracles of the nymph mildly convex instead of conically protuberant.

In common with megnini, the adults of lagophilus are not parasitic. In our
experience the nymphs are attached in the fur on the face of the rabbit near
the vibrissae. One living adult (17850) was found at the entrance of a rodent
burrow; another (19270) was found well down in a burrow.

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36

American Midland Naturalist Monograph No. 1

DISTRIBUTION AND HOSTS

This species is known from Alberta, Canada, and from the following
States: California, Colorado, Idaho, Montana, Nevada, Oregon, and Wyo-
ming. Except for a single record from a cat, the only known hosts are cotton-
tail rabbits, Sylvilagus sp., and jack rabbits, Lepus sp., including L. townsendu,
and L. calijornicus deserticola.

100 gpo 3Q0

p loo fto 400 too

Fig. 12. Distribution of Otobius lagophilus Cooley and Kohls.

Cooley & Kohls: Argasidae of N. America, etc. 37

Genus Ornithodoros C. L. Koch, 1844

18-44. Ornithodoros Koch, original description, p. 219.

1845. Ornithodorus Koch: Erickson, in Agassiz Nom. Zool. (spelling emended).

1877. Argas (Ornilhodoros) Murray, p. 183.

1895. Rhynchoprium Marx, p. 199. Misprint for Rhynchoprion Hermann, 1804, a
synonym of Argas Latreille.

1896. Ornilhodoros Koch: Neumann, redescribed, p. 25-26.

1907. Alectorobins Pocock, original description, p. 189 (new genus proposed for O.
lalaje) .

1908. Ornithodoros Koch: Nuttall ei a/., redescribed p. 39-40.
1911. Ornithodoros Koch: Neumann, redescribed, p. 122.
1932. Argas Latreille: Bedford, p. 280; in part.

1934. Argas Latreille: Bedford, p. 60; in part.

1936. Ornithodorus Koch: Brumpt, p. 1180 and p. 1192.

Body more or less flattened but never marginated (except dyeri) and with
the pattern of integumental structures continuous over the sides from dorsal
to ventral surfaces. Sutural line separating dorsal and ventral surfaces absent.
Usually very convex on dorsal surface when distended. Dorsal humps and
subapical dorsal protuberances on legs progressively more prominent in the
successive nymphal stages. Capitulum either subterminal or distant from the
anterior margin. Hypostome well developed and usually essentially alike in
the sexes and in nymphs and adults. Integument with discs and mammillae com-
mingling in a variety of patterns. Hood, camerostome, and cheeks present or
absent. Eyes present or absent.

Genotype: Argas savignyi Audouin, 1826, designated by Nuttall et al.
(1908) not O. coriaceus Koch designated by Oudemans (1936).

Key to Species of Ornithodoros

1 . Cheeks present, 4 dorsal humps on legs absent 2

Cheeks absent, dorsal humps on legs present or absent 13

2. Known only from bats and bat retreats 3

Known only from animals other than bats or from animal nests and burrows, or

from the open surface of the earth 11

3. Hypostome more or less notched apically 6

Hypostome not notched 4

4. Hypostome with denticles very faint 5

Hypostome with denticles distinct brodyi (p. 80)

5. Hypostome long, attenuated azteci (p. 109)

Hynostcme short, b-oad, ?nd pointed viguerasi (p. 106)

6. Body unusually long in proportion to width and with two parallel ridges follow-

ing the periphery Jljeri (p. 95)

Body not unusually long and lacking two parallel ridges on the periphery 7

7. Legs notably lone - - yumatensis (p. 74)

Legs not notably long 8

8. A small species with short legs (known from Panama) dunni (p. 103)

Species of moderate size or larger and with legs of about the usual length 9

9. Body oval, wider behind than in front - stageri (p. 91)

Body with sides about parallel 10

10. Larger species, and proportionately wider £e//e\>i (p. 113)

4 One species, stageri, lacks the cheeks in nymphs and males.

38 American Midland Naturalist Monograph No. 1

Smaller species, and proportionately narrower concanensis (p. 88)

1 1 . Discs large and noticeable 12

Discs small and inconspicuous rudis (p. 101)

12. Legs with the surfaces micromammillated talaje (p. 82)

Legs with the surfaces without micromammillae amblus (p. 99)

1 3. Dorso-ventral grooves present, capitulum subapical 14

Dorso-ventral grooves absent, capitulum distant from the anterior margin

cooleyi (p. 52)

14. Dorsal humps on tarsi I present 15

Dorsal humps on tarsi I absent hermsi (p. 46)

1 5. Eyes absent 1 6

Eyes present conaceus (p. 42)

16. Subapical dorsal protuberance on leg IV absent 17

Subapical dorsal protuberance on leg IV present nicollei (p. 50)

17. Dorsal humps absent on tarsus IV 18

Dorsal humps present on tarsus IV eremicus (p. 40)

18. Mammillae large, relatively few in number, not crowded turicala (p. 56)

Mammillae small, many, and somewhat crowded parpen (p. 62)

Ornithodoros savignyi (Audouin), 1826

Plate 4, Fig. 13
1826. Argas savignvi Audouin, p. 183, original description. (See Oudemans Part III,

B, P. 789, 1936.)

1844. Ornithodoros savignyi (Audouin) : Koch, p. 219.

1873. (?) Ornithodoros morbillosus Gerstaecker, p. 464: Nuttall et al. (1908, p. 42).

1889. Argas schinzii Berlese, p. 289.

1896. Ornithodoros savignyi (Audouin): Neumann, p. 26.

1908. Ornithodoros savignyi (Audouin) : Nuttall ei al., p. 42.

1911. Ornithodorus savignyi (Audouin): Neumann, p. 123.

1932. Argas savignyi Audouin: Bedford, p. 282.

1934. Argas savignyi Audouin: Bedford, p. 86.

1936. Ornithodorus savignyi (Audouin): Brumpt, pp. 1196-1197.
Sexes similar; adults and nymphs similar.

ADULT

Body. — Short, oval, broader behind, rounded on both ends. Size 10.0 x 7.5
to 13.0 x 9.0.

Mammillae. — Numerous, moderate in size, and of about equal size on all
parts of the tick. Individual mammillae, convex on top (some on the venter
are nearly flat on top) and with irregular, radiating ridges on their bases.
Under higher magnification some mammillae show faint, small pits on the
convex surface. Hairs smooth, tapering, arising from smallet mammillae; fewer
on the dorsum, more abundant on the venter and most abundant and longer
on the anterior and above the mouth parts.

Discs. — Distinct, small and placed in depressions; in a nearly symmetrical
pattern but more numerous than in many species of the genus.

Legs. — Short, moderately large; surface smooth (not granulated) . Hairs
on the legs few in number, longer on the dorsal surface of the articles and on
anterior legs. All femora distinctly larger distally than proximally. All tarsi
with the subapical dorsal protuberances pronounced and with 2 pronounced
dorsal humps. Metatarsi I, II, and III with 3 pronounced dorsal humps; IV
with 2. Length of tarsus I, 1.2; metatarsus, 0.96. Length of tarsus IV, 2.04;
metatarsus 1.725.

Cooley & Kohls: Argasidae of N. America, etc.

39

Coxae. Coxae contiguous and progressively reduced in sizes posteriorly.
Hood, Camerostome, and Cheeks. — Absent.

Capitulum. — Basis capituli contracted in front and behind, approaching a
sphere in shape; surface irregular and with faint transverse rugae; a group of
short spines on the sides, visible in ventral view.

Palpi. — Long, moderately heavy, article 2 much longer than article 1;
article 1 in contact with the base of the hypostome; articles 2, 3 and 4 free.
Length of article 4, three times its diameter.

Hypostome. — Short and large, rounded apically; denticles arranged ' |
but it is not easy to distinguish the longitudinal files. Denticles a little larger
in the lateral files. Length, 0.6.

Folds. — Supracoxal folds connected in front of the capitulum and termi-
nating posteriorly at preanal groove. Coxal fold extending from coxa II to
the preanal fold.

Grooves. — Dorso ventral groove present and visible from above as a notch
at the side. Preanal groove pronounced. Transverse postanal and median post-
anal grooves faint or absent.

Sexual opening. — At the level of the posterior ends of coxae I.

Eyes. — Two pairs of eyes present, all of about equal size.

Anus. — Small, in an elliptical pattern.

O. savignyi is readily distinguished from moubata, another African species,
by the absence of eyes in the latter.

HOSTS AND DISTRIBUTION

This species is found in human habitations and is said to occur in loose,
dry soil in places frequented by animals or used as resting places for caravans.
It occurs in Africa, Arabia, India, and Ceylon.

Fig. 13. Ornilhodoros savignpi (Audouin). A, Hypostome of adult. B, Leg I of
adult. C, Leg IV of adult.

40

American Midland Naturalist Monograph No. 1

Ornithodoros eremicus Cooley and Kohls, 1941
Plate 4, Figs. 14 and 16
1941(b). Ornithodoros eremicus Cooley and Kohls, original description, pp. 588-589.

NYMPH

Body. — (Described from an early stage nymph.) Short oval, approaching
circular, broadly rounded on both ends, tips of mouth parts visible from above.
Size, 1.44 x 1.11.

Mammillae. — Numerous, small, and of about equal size in median and
peripheral areas on both dorsal and ventral surfaces. Individual mammillae
only a little elevated, flattened, with their surfaces smooth and shining. A few
short hairs present, those on the anterior margin and above the anterior legs
larger.

Discs. — Little in evidence, small and slightly depressed; not evident on the
venter.

Legs. — Moderate in length and size. Surface smooth. Hairs few in number.
Subapical dorsal protuberance present on tarsi I, II, and III, absent on
tarsus IV. Dorsal humps moderate; three on tarsus I, two on II, two on III,
none on IV. Metatarsi I, II, and III, each with three dorsal humps; metatar-
sus IV, with two. Length of tarsus I, 0.3; metatarsus, 0.18. Length of tarsus
IV, 0.42; metatarsus, 0.285.

Coxae. — All coxae contiguous.
Hood, Camerostome, and Cheeks. — Absent.

Hypostome. — Moderate in length, sides parallel, apex rounded. Denticles
arranged -/•> with two or three in each file and placed near the distal end.

Fig. 14. Ornithodoros erimicus Cooley and Kohls. A, Hpostome of small nymph.
B, Tarsus and metatarsus of leg I of small nymph. C, Tarsus and metatarsus of leg IV
of small nymph.

Cooley & Kohls: Argasidae of N. America, etc. 41

Length about 0.12. (Description and drawing made with the hypostome in
situ — not mounted.)

Folds. — Coxal and supracoxal fo!ds present.

Grooves. — Dorso-ventral and preanal grooves present. Median postanal
and transverse postanal grooves absent.

Eyes. — Absent.

Anus. — In a nearly circular frame.

It is to be noted that although the specimen is in an early nymphal stage
the dorsal humps on the tarsi are well developed. Since in all known American
species the dorsal humps are poorly developed in the early nymphal stages and
become progressively larger in successive stages, it appears likely either that
the adult would be of a species of small size, or if of average size, the humps
would be unusually prominent.

This tick resembles the African species, O. savignyi.

HOST AND DISTRIBUTION

Known only on Peromyscus maniculatus . The one nymph was collected
near Bluff, San Juan County, Utah, August 24, 1939.

42 American Midland Naturalist Monograph No. 1

Ornithodoros coriaceus Koch, 1844

Plate 5, Figs. 15 and 16

1844. Ornithodoros coriaceus Koch, original description, p. 219.

1847. Ornithodoros coriaceus Koch: Koch, redescribed, p. 31, with figures.

1888. Argas (Ornithodoros) coriaceus Koch: Berlese, p. 193.

1896. Ornithodoros coriaceus Koch: Neumann, redescribed, p. 31.

1901. Ornithodoros coriaceus Koch: Neumann, redescribed, p. 258.

1908. Ornithodoros coriaceus Koch: Banks, retescribed, p. 18-19, with figures.

1908. Ornithodoros coriaceus Koch: Nuttall, Warburton, Cooper, and Robinson,

redescribed, pp. 55-56, with figures.

1911. Ornithodoros coriaeceus Koch: Neumann, redescribed, p. 124.

1930. Ornithodoros coriaceus Koch: Hoffman, redescribed, pp. 163-164.

1936. Ornithodoros coriaceus Koch: Brumpt, p. 1206.

Sexes similar; nymphs and adults similar.

ADULT

Body. — Sub-oval, somewhat pointed anteriorly and with the sides nearly
parallel. Nuttall et ai. (1908) give as the size of this tick, largest female 13.8
x 8.2, smallest, 9.5 x 5.3; largest male 8.6 x 4.6, smallest, 6.4 x 3.4. Specimens
in the collections of the Rocky Mountain Laboratory are of intermediate sizes
for the most part but one male measures 5.6 x 3.4.

Mammillae. — Numerous, moderate and variable in size, irregular in shape.
In clean specimens5 the mammillae have the flattened tops ornate. Nuttall
et al. (1908) state that the color is white in living and reddish in old preserved
specimens. Living and freshly preserved specimens have the color reddish-gray,
almost iridescent, with a few punctations interrupting the color. (Color is
better seen in immersed specimens.) Interspersed in the mammillated areas
are numerous large, deep pits independent of the discs, each with a conical,
truncate elevation, bearing a hair.

Discs. — Large, depressed, with surface of the floors of the depressions
irregular, resembling the mammillated areas. Discs less definite or absent on
the ventral surface.

Legs. — Moderate in length; femur, tibia and metatarsus flared distally.
Hairs few, short, and inconspicuous. Subapical dorsal protuberances pro-
nounced on all tarsi; dorsal humps three or four on tarsi I to III: one at the
base on tarsus IV. Length of tarsus I, 1.23; metatarsus, 0.96. Length of tarsus
IV, 1.92; metatarsus, 1.5.

Coxae. — Coxae I and II a little separated; all others contiguous.

Hood. — Separated from the anterior dorsal projection of the body wall.

Camerostome. — Definite and deep.

Cheeks. — Absent.

Capitulum. — Basis capituli broader than long; surface very irregular with

5 Specimens of Ornithodoros collected in nature are often badly encrusted with dirt,
and this is particularly true of coriaceus.

Coolky & Kohls: Argasidae of N. America, etc.

43

coarse transverse wrinkles. Palpal article 1 with longitudinal wrinkles and with
a thin flange extending over the base of the hypostome.

Hypostome. — With the sides nearly parallel. Denticles arranged -j-?, with
about four large ones in each file, all about equal in size. Length in the
female, 0.57; male, 0.45.

Folds. — Coxal and supracoxal folds present.

Grooves. — Dorso-ventral, preanal, transverse postanal, and median post-
anal grooves present; dorso-venrral groove visible in dorsal view as a marginal
dent.

Sexual opening. — At the level of coxae I.

Eyes. — Two pairs of ocellus-like eyes present, those of the anterior pair
much larger.

Anus. — In an elliptical frame.

Fig. 15. Ornilhodoros coriaceus Koch. A, Leg I of adult. B, Leg IV of adult.
C, Capitulum of larva, ventral view. D, Larva, dorsal view. E. Hypostome of adult.

44

American Midland Naturalist Monograph No. 1

LARVA

Oval, a little larger in front, with the entire capitulum visible from above;
dorsal surface convex, with the sparse hairs arranged in a symmetrical pattern.
Integument striated; dorsal plate absent. Two pairs of eyes present, the anter-
ior ones about twice as large as the posterior. Length, not including the
capitulum, 0.75; width, 0.6.

Legs long and slender. Length of tarsus I, 0.3; metatarsus, 0.2. Stalk of

Fig. 16. Distribution of O. coriaceus Koch, and O. eremicus Cooley and Kohls. See
also text.

Cooley & Kohls: Argasidae of N. America, etc. 45

the pulvillus and claws long and slender. Palpi long and slender; terminal
article about three times as long as wide.

Hypostome. — Long, and with the denticles arranged -/L», lateral denticles
larger; denticles present for most of the length. Apex rounded, corona with a
few very fine denticles. Length, 0.18.

Fed larvae are short-oval, measuring about 3.75 x 2.75. Leg I as distant
from the capitulum as from leg II.

HOSTS AND DISTRIBUTION

The type specimens were from Mexico.

California. — Banks (1908) saw specimens from San Francisco and Santa
Clara County and stated that "it has been taken from cattle and cattlemen."
Nuttall, et al. (1908) mention specimens collected at Los Olivos (Santa
Barbara County) in August, 1904. Herms (1939) stated that the species
occurs in the more mountainous coastal counties and that he collected it on
Mount Hamilton (Santa Clara County) in deer beds among the low scrub
oaks (Querent diunosa) . Kohls collected 200 nymphs and adults on Mount
Hamilton, May 15, 1940.

We have seen specimens as follows: 8692 A, on ground, September 11,
1932, Mount Hamilton, Santa Clara County, 5 immature specimens (Heber
Donohoe); 8607A, deer bed ground, Mount Hamilton, July 1932, Santa
Clara County, several specimens; 12751, ground, October 1936, Monterey
District, 1 adult (Joseph W. Burke); 5 lots, dates ranging from June 10, 1938
to January 30, 1940, Hastings Natural History Reservation, Monterey Coun-
ty, several specimens of all stages, (Dr. J. M. Linsdale); 16286, Monterey
County, June 30, 1939, 1 nymph; 17338-39-41, host not stated, April, May,
June, 1940, Calaveras Dam, Alameda County, 3 nymphs (Robert Holden-
ried); 17392, Cliff swallow's nest in cave, April 8, 1932, Mt. Diablo, Contra
Costa County, 1$ (Aitken and Howell); 17955, Odocoileus hemionus,
August 8, 1940, Hastings Natural History Reservation, Monterey County, 29
nymphs (Dr. J. M. Linsdale); Odocoileus hemonicus colambianus Septem-
ber 8 and 9, 1941, Cobb Mt., Lake Co., 4 nymphs (J. E. Hare).

Mexico. — Nuttall et al. (1908) determined specimens collected in Janu-
ary 1908. They were found "whilst exploring some rock inscriptions near San
Geronimo, on a hill in a sandy plain (Isthmus of Tehauntepec) ."

Hoffman (1930) stated: "O. coriacens is a native of the hot and temper-
ate regions along the Pacific extending all the way from California to Chiapas.
As to the eastern or Gulf region of Mexico, I am as yet uninformed." (Trans-
lation.)

46 American Midland Naturalist Monograph No. 1

Ornithodoros hermsi Wheeler, Herms and Meyer, 1935

Plate 5, Figs. 17 and 18

1935. Ornilhodorus hermsi Wheeler, Herms and Meyer, not intended to be the original

May description but diagnostic differentiation is included, p. 1290-1292.

1935. Ornithodorus hermsi Wheeler, full description, pp. 435-438.
July

1936. Ornithodoros hermsi, Wheeler, Herms and Meyer: Brumpt, pp. 1206-1208.
Sexes similar; adults and nymphs similar.

ADULTS

Body. — Oval, wider behind, pointed anteriorly. The size of the female
given by Wheeler was 5.0 x 3.1, and of the male, 3.8 x 2.4. Average sizes of
numerous specimens in the Rocky Mountain Laboratory collections are, for
females, 4.85 x 3.01 and for males, 3.42 x 2.42. The largest female measures
5.37 x 3.24 and the largest male, 3.69 x 2.40.

Mammillae. — Numerous and moderate in size with the individual mam-
millae elongated, not conical, with sinuous, radiating ridges on the sides;
longer axes longitudinal in the lateral area; and in the median areas with their
longer axes tending to be concentric around the discs or groups of discs; of
about equal size throughout on the dorsal surface, smaller in the median areas
than at the sides on the ventral surface.

Hairs. — A few fine hairs usually visible.
Discs. — Definite, but shallow.

Legs. — Surface of legs smooth, not micromammillated, and with a few
hairs present. Legs short and small. Length of tarsus I, 0.42; metatarsus, 0.3.
Tarsus IV, 0.48; metatarsus, 0.36. All tarsi without dorsal humps but with
mild subapical dorsal protuberances.

Coxae. — Coxae I and II a little separated, all others contiguous.

Hood. — Well developed as an anterior projection above the capitulum
and continuous with the anterior extension of the dorsal body wall.

Camerostome. — Deep posteriorly, shallow anteriorly. In fully fed speci-
mens the camerostome is scarcely visible.

Cheeks. — Absent.

Capitulum. — Basis capituli with the length and width about equal. Surface
irregular with interrupted transverse wrinkles and with a group of short hairs
on each side at the base.

Palpi. — Surface smooth. Article 1 with a narrow, knifeedge flange extend-
ing over the base of the hypostome; articles 2, 3, and 4 fre>».

Hypostome. — Length from 0.21 to 0.27. Denticles arranged 2/2 and lim-
ited to the distal one-third, about five in each file; denticles of all files of about
equal size. Apically notched and the corona having numerous very fine denti-
cles. Posthypostomal hairs long.

Cooley & Kohls: Argasidae of N. America, etc.

47

Folds. — Coxal and supracoxal folds present.

Grooves. — Dorso-ventral groove present but faint. Preanal groove pro-
nounced, sinuous; transverse postanal groove pronounced, interrupted at the
median postanal groove which is deep and reaches from near the anus to near
the posterior border.

Sexual opening. — At the level of the interval between coxae I and II.

Eyes. — Absent.

Anus. — In a long, oval pattern.

LARVA

Oval, broader behind; mouth parts short and capitulum visible from above.
Integument thin and striated; hairs in a symmetrical pattern on dorsal and
ventral surfaces.

Hypostome. — Sides nearly parallel, apically truncated or slightly notched;
denticles 2/2 with about four large ones in each file and occupying about two-
fifths of the distal portion. Posthypostomal hairs about three-fifths the length
of the hypostome and arising from a prominent swelling. Length, about 0.08.

HOSTS

It has been assumed that O. hermsi bites man because of the known cases
of relapsing fever originating from mountain cabins in wh;ch this species has
collected. Until recently there have been no published, definite records of the
hosts of this tick, but the account of Philip and Davis (1940) gives records
of this tick biting man.

D

Fig. 17. Orniihodoros hermsi Wheeler. Herms and Meyer. A, Hypostome of adult.
B. Leg I of adult. C, Leg IV of adult. D, Capitulum of larva, ventral view.

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Cooley & Kohls: Argasidae of N. America, etc.

49

Dr. Gordon E. Davis, of the Rocky Mountain Laboratory, in connection
with his studies on relapsing fever in Colorado in 1938, collected nymphs and
adults from a chipmunk nest in an old, rotten stump of a Douglas fir tree.
There were no animal burrow holes in the ground in this vicinity. When
pushed over, the stump fell into numerous pieces and with extended search
51 specimens were recovered. The broken stump revealed a "cache" of corn
in a hollow root, and nesting material of fine, dry grass well above the ground.
The greater number of specimens was taken from the cracks and crevices of
the stump material, and many of them had recently fed While the ticks were
being collected a chipmunk hovered around "scolding." From these details it
is assumed that the chipmunk was the host animal.

Again, on June 11, 1939, Dr. Davis took 213 specimens from the cracks
and crevices of a rotten Douglas fir stump in Park County, Colorado. In this
stump there was very extensive nest material.

The records show that this species has been taken only from higher eleva-
tions. Lake Tahoe has an elevation of 6225 feet. The cabin at Moscow, Idaho,
has an elevation of about 3000 feet, and the mountainous area in Colorado
where the species was taken has an elevation of approximately 8800 feet.

Briggs (1935) states that Mark F. Boyd in a personal communication had
pointed out that O. talaje was identified as far back as 1914 from within a few
miles of Polaris (near Lake Tahoe, California) . The one remaining specimen
of the lot from which the record had been made was kindly sent to Dr. Davis
and examined by Cooley, who found it to be a specimen of O. hermsi. Thus,
this species was taken from near Lake Tahoe twenty years before the species
was described. The known records of collections of this species follow.

Fig. 18. Distribution of Ornilltodoros hermsi Wheeler. Herms and Meyer.

50 American Midland Naturalist Monograph No. 1

Ornithodoros nicollei Mooser, 1932

Plate 6, Figs. 19 and 22

1932. Ornithodorus nicollei Mooser, original description, pp. 127-131, with figures.
1936. Ornithodorus nicollei Mooser: Brumpt, pp. 1210-1211.
Sexes similar; adults and nymphs similar.

adult

Body. — Broadly rounded at both ends, hood barely visible from above.
Size of female, 7.0 x 4.9 to 10.3 x 6.1; male, 5.0 x 3.5 to 7.10 x 4.2.

Mammillae. — Numerous, moderate in size and on the dorsum about the
same in size throughout; smaller on the venter except at posterior end. Indi-
vidual mammillae sub-oval, conical, with their tops convex, smooth, and each
with a few small pits; sides with radiating striae which reach nearly to the
tops. Interspersed among the mammillae on both dorsal and ventral surfaces
are conical elevations which are truncated and each bears ' hair.

Discs. — Small, depressed and not conspicuous; absent or obscure on venter.

Legs. — Moderate in length and size, with the articles flared distally. All
tarsi with a conspicuous subapical dorsal protuberance and all except IV with
a prominent dorsal hump near the proximal end. Length of female tarsus I,
0.66; metatarsus, 0.57; length of tarsus IV, 0.72; metatarsus, 0.96.

Coxae. — Coxae I and II slightly separated, all other contiguous.

Hood. — Moderate, not continuous with the anterior extension of the
dorsal body wall.

Camerostome. — Shallow.

Cheeks . — Absent.

Capitulum. — Basis capituli wider than long, narrower in front; surface
irregular, and with a group of short, heavy spines on each side behind. Palpi
large, article 1 tumescent with a knife-edge flange overlying the base of the
hypostome and with a row of spines on the median side.

Hypostome. — Large and long, sides parallel, apex rounded; denticles in a
- ,'.) arrangement, all of about equal size and confined to about three-fifths of
the apical portion. Corona with numerous small denticles. Length, 0.6.

Folds. — Coxal and supracoxal folds present.

Grooves. — Dorso-ventral groove pronounced and visible in dorsal view.
Preanal, transverse postanal, and median postanal grooves pronounced; median
postanal groove continuous from near the anus to near the posterior margin.

Sexual opening. — Placed at the level of the intervals between coxae I
and II.

Eyes. — Absent.

Anus. — In an oval pattern.

Cooley & Kohls : Argasidae of N. America, etc.

51

NYMPH

The subapical dorsal protuberances and the dorsal humps are smaller in
the nymphs; small or negligible in the very small nymphs and become progres-
sively larger in the stages up to the adult.

HOSTS AND DISTRIBUTION

The type specimens were collected in Mexico, State of Guerrero, in native
huts.

Mexico. — Through the courtesy of Dr. Mooser, we have seen specimens,
immature and adult, collected in native huts, Balsas River villages, State of
Guerrero, April 1932.

Brumpt, Mazzotti and Brumpt (1939) mentioned specimens collected
from Neotoma (Hodotnys) alleni near Colima, State of Colima.

Missouri. — We have 1 adult and 23 nymphs collected from Florida dia-
mond back rattlesnake, St. Louis Zoo, November 1933.

Fig. 19. Ornithodoros nicollei Mooser. A, Leg IV of female. B, Leg I of female.
C, Leg IV of a small nymph. D. Leg I of a small nymph. E, Hypostome of female.

52

American Midland Naturalist Monograph No. 1

Ornithodoros cooleyi Mclvor, 1941

Plate 6. Figs. 20, 21. and 22
1941. Ornithodoros cooleyi Mclvor, original description, with figures, pp. 435-436.
Whether the sexes and adults and nymphs are similar or dissimilar is

lkn

unknown.

FEMALE

Body. — Sub-oval, sides nearly parallel, anterior end bluntly pointed, pos-
terior end broadly rounded. Length, 9.25; width, 4.5.

Mammillae. — Integument of the dorsal and the median posterior ventral
surfaces in a reticulated pattern made up of numerous deep pits surrounded
by continuous ridges of crowded mammillae of irregular shape. Surface of
the floors of the pits irregular and each pit with a hair on an elevation placed
on the anterior side of the pit. Reticulated surfaces continuous over the sides of
the body with pits more shallow on the posterior ventral surface and indistinct
or absent on the supracoxai folds, the inter-coxal area and in the vicinity of the
mouth parts. Surface wrinkled posterior to the capitulum and at the sides

Fig. 20. Ornithodoros cooleyi Mclvor. A, Capitulum of nymph, ventral view. B, Leg
I of adult. C, Leg IV of adult. D, Leg I of last stage nymph. E, Leg IV of last stage
nymph.

Cooley & Kohls: Argasidae of N. America, etc.

53

of the camerostome. Hairs absent in the median area between the coxal folds,
few in number in the postero-Iateral surfaces but more numerous and larger
on the antero-lateral surfaces.

Discs. — Reticulated dorsal surface interrupted by depressed areas which
include the discs. Discs large, distinct and their floors made irregular by
elevations similar to the mammillae. Discs visible in the median postanal
groove.

Legs. — Moderate in length and size and with a few inconspicuous hairs.
Femora a little flared distally. Subapical dorsal protuberance prominent on
all tarsi. Two dorsal humps on tarsus I; a milder one near the articulation
with the metatarsus on legs II, III, and IV. With two or three mild dorsal
humps on metatarsi I and II. Length of tarsus I, 0.72; metatarsus, 0.54.
Length of tarsus IV, 1.02; metatarsus, 0.84.

Coxae. — Coxae I and II well separated; all others contiguous.

Hood. — No obvious hood present but in its place is a mild elevation in

Fig. 21. Ornithodoros coo/epi Mclvor. Anterior portion of female, ventral view.

54

American Midland Naturalist Monograph No. 1

the shape of an inverted Y with the mouth parts within the arms and the stem
extending to the anterior margin.

Camerostome. — Distinct.

Cheeks . — Absent.

Capitulum. — Capitulum distant from the anterior margin. Basis capituli
subquadrate, slightly narrower anteriorly. Surface irregular, wrinkled and with
an elongated depression on each side parallel with the lateral 'border. With a

100 200 3Q0

Fig. 22. Distribution of Ornithodoros nicollci Mooser and O. cooleyi Mclvor.

Cooley & Kohls: Argasidae of N. America, etc. 55

group of hairs on the margin on each side back of the palpi. Palpi moderate
in length and with a few dorsal hairs. Article 1 with a narrow, knife-edge
flange extending over the base of the hypostome; articles 2, 3, and 4 free.

Hypostome. — Long, with the apex broadly pointed. Denticles arranged
2/2, with about six in each file. Posthypostomal hairs, usually found in this
genus, are absent. Length, 0.57.

Folds. — Coxal and supracoxal folds are distinct.

Grooves. — True grooves are absent though a median elongated depression
posterior to the anus suggests a median postanal groove.

Sexual opening. — Placed at the level of the posterior ends of coxae I.

Anus. — In an oval pattern.

Eyes. — Absent.

NYMPH

The single specimen from which this species was described was received
by Miss Mclvor as a living nymph. It was fed and in due course emerged as
an adult. Thus, the cast skin was available for study and reveals some nymphal
characters which differ mildly from the adult as follows: Reticulation and pits
on the dorsum more shallow and less definite. Dorsal humps and subapical
dorsal protuberances less prominent. Length of hypostome, 0.48. Length of
tarsus I, 0.6; metatarsus, 0.42; length of tarsus IV, 0.84; metatarsus, 0.6.

HOST AND DISTRIBUTION

The single specimen was found by Dr. E. Raymond Hall, Museum of
Vertebrate Zoology, University of California, Berkeley, in a shipment of skins
sent by Mr. Frank Wilson, Rox, Nevada, which contained pelts of a striped
skunk, swift foxes, and coyotes. The host of this tick is presumably one of
these three animals. The specimen was given to Miss Barbara C. Mclvor, The
Medical Center, University of California, San Francisco, California, who has
kindly given the authors full opportunity to study, figure, and photograph
the specimen.

56 American Midland Naturalist Monograph No. 1

Ornithodoros turicata (Duges), 1876
Figs. 23, 24, 27. 28, 29, 30, 31. 33 and 34

1876. Argas turicala Duges, original description in "Repertorio de Guanajuato,"
April 25.

1884. Argas turicata Duges: Duges, redescnbed pp. 195-197, with figures.

1885. Argas turicata Alf. Duges: Megnin, redescribed, pp. 463-470, with figures.

1895. Ornithodoros americanus G. Marx, figures only, p. 199, Marx obituary notice.

1896. Ornithodoros turicata (Alf. Duges): Neumann, redescribed, pp. 31-34.
1908. Ornithodoros turicata (Duges) : Banks, redescribed, p. 18, with figures.

1908. Ornithodoros turicala (A. Duges) : Nuttall, Warburton, Cooper and Robinson,

redescribed, pp. 57-59, with figures.
1911. Ornithodoros turnicata (Alf. Duges): Neumann, redescribed, p. 124.
1930. Ornithodoros turicata (Duges): Hoffman, redescribed, pp. 155-160, with figures.
1936. Ornithodorus turicata (A. Duges): Brumpt, pp. 1201-1203.

Sexes similar; nymphs and adults similar.

ADULT
Body. — Oval, slightly wider behind, anterior and posterior ends evenly
rounded, hood visible from above. Size of female from 9.90 x 6.80 to 6.20 x
4.6. Size of male from 6.3 x 4.0 to 5.4 x 3.30. Nuttall et al. (1908) give as
average size of females 6x4, and of male, 3.5 x 2.5. Thus, there is evident
an unusual range of sizes in the species.

Mammillae. — Moderately numerous and larger than in parken, distinctly
larger at the posterior margin and smallest in the mid-ventral area. The indi-
vidual mammillae are raised abruptly from the surface, nearly hemispherical
on top with small pits in the convexity and with radiating striae at the bases.
Mammillae less crowded than in parkcri, and more elevated, with the radiat-
ing ridges at the bases more distinct. Interspersed with the usual mammillae
are smaller conical, truncate elevations, each with a hair arising from a central
papilla. Similar but larger hairs occur on the anterior body wall above the
camerostome.

Discs. — Distinct, and depressed. Absent on the ventral surface except in
the grooves where they are in lineal arrangement.

Legs. — Moderate in length and size, a little larger than in parker'h with
articles mildly flared distally. Surface smooth and shining (without micro-
mammillae or granulations). Hairs on legs small, short. Tarsi I, II, and III
with a mild su'bapical dorsal protuberance; absent on tarsus IV. Tarsus I with
three or four dorsal humps; metatarsus I with four or five; absent on tarsus
IV. Length of female tarsus I, 0.72; metatarsus, 0.60. Length of female tarsus
IV, 1.08; metatarsus, 1.08. Length of male tarsus I, 0.84; metatarsus, 0.66.
Length of male tarsus IV, 0.96; metatarsus, 1.05.

Coxae. — Coxae prominent, progressively larger anteriorly. Coxae I and II
slightly separated; all others contiguous.

Hood. — Large and prominent, not continuous with the dorsal body wall.

Cheeks. — Absent.

Capitulum. — Basis capituli about as wide as long; a little smaller than in

Cooley & Kohls: Argasidae of N. America, etc.

57

parken. Surface irregular and with a few transverse wrinkles; with a few
spines in a group on each side. Palpi long, longer than in parken, with surface
of all articles smooth. Article 1 with a knife-edge flange for the entire length
on the median side extending over the base of the hypostome; articles 2, 3,
and 4 free.

Hypostome. — Long, sides nearly parallel and rounded apically. Denticles
arranged -/•>, with the large teeth limited to about the distaf two-fifths of the
length and with teeth in lateral and median files about equal in size. Length

vr<

Fig. 23. Ornithodoros turicata (Duges). A, Leg I of female. B, Leg IV of female.
C, Hypostome of female. D. Hypostome of male. E, Capitulum of larva, ventral view.
F. Larva, dorsal view.

58 American Midland Naturalist Monograph No. 1

in the female, 0.60 tc 0.69, and in the male, 0.45 to 0.525. Much longer than
in p<xrk.er\.

Folds. — Coxal and supracoxal folds present.

Grooves. — Dorso-ventral groove present and visible on the dorsal surface
at the sides. Preanal, transverse postanal and median postanal grooves pro-
nounced.

Sexual opening. — At the level of the intervals between coxae I and II.

Eyes. — Absent.

Anus. — In an oval pattern.

LARVA

Oval, slightly wider behind, mouth parts visible from above. Integument
thin, striated and with the sparse hairs in a symmetrical arrangement. Length,
not including mouth parts, 0.84, width, 0.57.

Hypostome. — Long, sides nearly parallel, and rounded apically; denticles
2/o occupying about the apical two-fifths of the length, and with the lateral
denticles larger. Posthypostomal hairs long. Length of hypostome about 0.135.

HOSTS

O. turicata attacks a wide variety of hosts, including reptiles, birds, and
mammals. Specimens have been collected from rattlesnakes, turtles, burrowing
owls, ground squirrels, prairie dogs, kangaroo rats, woodrats, rabbits, pigs,
cattle, horses, and man. In Kansas, Oklahoma, and Texas we have found it
in large numbers in rodent burrows and burrows used by burrowing owls,
Speotyto cunicularia. Davis (1936) reported the collection of eleven hundred
and ninety-seven specimens from a single sand hole which contained 1 1 terra-
pins, Terrapene ornata, in Clark County, Kansas. The U. S. Bureau of
Animal Industry has several records from pigs and from sinks frequented by
pigs in Florida and according to Hoffman (1930) there have been many
serious outbreaks of plagues among pigs in the central part of Mexico where
enormous quantities of the species have been found on the ground in pig sties.
All stages attack man readily. According to our experience the bite is pain-
less but is followed in a few hours by intense local irritation and swelling.
Subcutaneous nodules often form which, accompanied by occasional itching,
may persist for months. However, in Mexico, Hoffman (1930) states that
the bite is painful and often followed by serious secondary consequences,
characterized by the formation of gangrene in the skin. This latter effect he
attributes to the filthy habitat of the species, i.e., pig sties, etc.

This species is the only known vector of human relapsing fever in portions
of Kansas, Oklahoma, Texas, and possibly other areas in the Southwestern
United States. Hoffman believes that O. turicata has little to do with the
transmission of relapsing fever in Mexico since no cases have been found in
the region where this tick occurs.

distribution

The localities for which there are definite records of the collection of O.
turicata are shown in figure 24. In the United States the species is known
from Utah, Colorado, Kansas, Oklahoma. Texas, New Mexico, Arizona,

Cooley 8c Kohls: Argasidae of N. America, etc.

59

California, and Florida. According to Hoffman (1930), "O turicata is found
in Mexico throughout about the same regions as [O. megnint]', that is to say,
it prefers the dry sections of the Central and Western Coast States but has
seldom been seen at altitudes of more than 2,000 meters. In the Valley of
Mexico it has been found in some instances on pigs imported from the in-
fested regions, but apparently cannot exist for any length of time in the
climate of the valley. The species rather prefers the less cold regions between
1,000 and 1,800 meters." (Translation.)
The type locality is Guanajuato, Mexico.

Fig. 24. Distribution of Ornithodorob hiricaia (Duges).

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62 American Midland Naturalist Monograph No. 1

Ornithodoros parkeri Cooley, 1936
Figs. 25, 26. 27. 28, 29, 30, 32 and 34

1936. Ornithodoros parkeri Cooley, original description, pp. 431-433, with figures.

1937. Ornithodoros rvheeleri Mclvor, original description, pp. 365-367, with figures.

Sexes similar; nymphs and adults similar.

ADULT

Body. — Oval, a little broader posteriorly; hood visible from above. Size of
female 7.10 x 4.50 to 5.82 x 3.75. Size of male 5.75 x 4.00 to 5.0 x 3.25.

Mammillae. — More numerous and smaller than in tnncata, a little larger
at the sides and largest of all at the posterior end, smallest in the mid-ventral
region. The individual mammillae smooth, convex on top and in the convexity
are usually present a few very small pits. Sides of the mammillae with radiat-
ing ridges. Mammillae more crowded together, less elevated and with the radi-
ating ridges at their bases less visible than in turicata. Interspersed with the
mammillae are smaller conical elevations, each with a hair arising from an
apical pit. Hairs sparse or absent on the ventral surface.

Discs. — Discs distinct, small and depressed; absent on the ventral surface
except in the grooves where they are in lineal arrangement.

Legs. — Long and moderate in size, a little heavier than in turicata, with
articles mildly flared distally. Surface of the legs smooth (without micro-
mammillae or granulations) . Short hairs moderate in number. Tarsi I, II and
III with moderate subapical dorsal protuberances; small or absent on tarsus
IV. Tarsus I with three or four dorsal humps; metatarsus with four dorsal
humps. Tarsus IV with no dorsal humps. Length of female tarsus I, 0.69;
metatarsus, 0.57. Length of female tarsus IV, 0.96; metatarsus, 1.11. Length
of male tarsus I, 0.6; metatarsus, 0.51; length of male tarsus IV, 0.75; meta-
tarsus, 0.81.

Coxae. — Coxae prominent, progressively larger anteriorly. Coxae I and II
a little separated; all others contiguous.

Hood. — Large, prominent, not continuous with the body wall of the
dorsum.

Cheeks . — Absent.

Capitulum. — Basis capituli about as wide as long, a little larger than in
turicata; surface irregular and with transverse wrinkles; with a group of hairs
on each side. Palpi moderately long, longer than the hood; all articles smooth.
Article 1 with a thin flange on the median side extending the full length of
the article and projecting over the basal portion of the hypostome.

Hypostome. — Long, with sides about parallel, rounded apically. Denticles
in a 2/2 pattern covering about the distal half, and with teeth in lateral and

Cooley 6c Kohls: Argasidae of N. America, etc.

63

median files about equal in size. Length of female hypostome, 0.40; male, 0.33.
Shorter than in turicata.

Folds. — Coxal and supracoxal folds present.

Grooves. — Dorso-ventral groove present and visible at the sides on the
dorsal surface of the tick. Preanal, transverse postanal and median postanal
grooves are present, pronounced.

Sexual opening. — At the level of the intervals between coxae I and II.

r-\

Fig. 25. Ornithodoros parfycri Cooley. A, Leg I of female. B, Leg IV of female.
C, Hypostome of female. D, Hypostome of male. L, Capitulum of larva, ventral view.
F, Larva, dorsal view.

64

American Midland Naturalist Monograph No. 1

Eyes.-
Anus.

-Absent.
—In an oval

pattern.

LARVA

Oval, wider anteriorly, mouth parti visible from above. Integument thin,
striated; with sparse hairs in a symmetrical pattern. Length, not including
mouth parts, 0.66; width, 0.48.

Hypostome. — Hypostome moderate in length, apically rounded; denticles
2/o, occupying about the apical one-third and with the lateral teeth larger.
Posthypostomal hairs long. Length of hypostome about 0.1.

HOSTS

Ornithodoros parkeri was originally described from specimens that had
been taken from Citellus sp., Lepus sp., and Cynomys sp., forty miles south-
west of Casper, Wyoming. In our field experience it soon developed that while
scattering specimens could be collected on animals away from their burrows,
the tick was to be found more regularly and sometimes in abundance in the
burrows.

Jellison (1940) examined 18 burrows and nests of the burrowing owl,
Speotyto cunicularia, in Washington and nine were infested with O. parkeri.
In nest 106 he took 491 ticks, and in nest 108, he took 360 ticks. Some of
the nests containing fledglings had ticks that were fully fed, and microscopical
examination showed that they contained avian blood. Thus it is evident that
the burrowing owl is a natural host of O. parkeri.

Fig. 26. Distribution of Ornithodoros parfyeri Cooley.

Cooley & Kohls: Argasidae of N. America, etc. 65

It is apparent that O. parkeri is less avid in attacking man than are some
other species in the genus. Davis (1939) reported that this tick when placed
on the bare arm feeds promptly. However, neither he not his assistants, or
Jellison, were bitten while collecting this species, though engaged in the work
for hours at a time.

In summary, the known hosts of O. parkeri are: Citellus sp., C. columbi-
anus, C. nchardsoni, Cynomys sp., Cyn. leucurus, Marmota sp., Peromyscus
sp., Lepus sp., Sylvilagus sp., Mustela sp., Speotyto cunicularia, and man.

DISTRIBUTION

Ormthodoros parkeri has been collected in widely separated areas in the
following nine Western States: California, Colorado, Idaho, Montana,
Nevada, Oregon, Utah, Washington, and Wyoming. Collection localities are
shown in figure 26.

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Cooley & Kohls: Argasidae of N. America, etc.

69

COMPARISON OF O. TURICATA AND O. PARKERI

Figs. 27, 28, 29, 30, 31, 32, 33. and 34

O. turicata and O. parkeri are rather closely related and yet in most in-
stances are readily separated by their mammillae and hypostomes. In these

Fig. 27. Female of Ornithodoros luricala (Dugcs) on the left, and female of O.
parfycri on the right.

70

American Midland Naturalist Monograph No. 1

species, as well as in others of the same genus, the characters of the mammil-
lae may be less definite or distinct when the ticks are fully fed. Depleted
specimens show the mammillae closer together and their elevation is at a max-
imum. The individual mammillae are then at their best as characters and show
clearly the true shape and markings, and the folds and grooves are either
normal or exaggerated. The length and width of the tick are but little in-

Fig. 28. Male of Ornilhodoros iuricaia (Duges) on the left, and male of O. parken
Cooley on the right.

Cooley & Kohls: Argasidae of N. America, etc.

71

creased by feeding but the mammillae become much flattened and in some
species, including O. tuncata and O. parkeri, are visible only as shining
sclerites not agreeing with their descriptions. Similarly, the folds and grooves
may disappear almost completely and be visible only as faint remnants. As
specimens in Ormthodoros, the 'best ticks are those that have been depleted
until the dorsal and ventral body walls are about parallel.

i m. .

A

C

Si-

B

D

Fig. 29. A. Omilhodoros turicata (Duges), fourth nymphal stage, dorsal view.
B, Same, ventral view. C, 0. par^eri Cooley, fourth nymphal stage, dorsal view.
D, Same, ventral view.

72

American Midland Naturalist Monograph No. 1

The two species occupy overlapping ranges (see maps) . It should be
realized too, that in the area where both species occur and use the same animals
as hosts there is a possibility that O. turicata and O. parkeri may have hybrid-
ized. In fact, Dr. Davis has interbred these two species in the laboratory
(unpublished) and certain field specimens from the overlapping area show
intergraded characters.

Comparative studies have shown that other than the more definite charac-
ters which separate O. turicata and O. parkeri both as adults and nymphs,
such as hypostome and mammillae, there is little of value for separating the
two species. Both show the dorsal humps on the tarsi absent in N. 1, faint in
N. 2, and progressively more prominent in the successive stages to the adults.
O. savigny and eremicus show these humps to be very well developed in the
early nymphal stages.

Photographs of all stages of both species are shown in figures 26, 27, 28,
and 29, and pen drawings of the hypostomes and legs of all stages in figures

Fig. 30. Ornithodoros turicata (Duges) on the left, O. parkeri on the right. A, Third
stage nymph, dorsal view. B, Same, ventral view. C, Second stage nymph, dorsal view.
D, Same, ventral view. E, First nymphal stage, dorsal view. F, Same, ventral view.

Cooley & Kohls: Argasidae of N. America, etc.

73

30, 31, 32, and 33. The drawings of the hypostomes show clearly that these
characters are dependable even in the early nymphal stages. The same is true
of all other species of Ormthodoros so far as we have studied them.

Fig. 31. Ornilhodoros luricala (Duges). A, Hypostome of female. B, Hypostome
of male. C, Hypostome of fourth stage nymph. D, Hypostome of third stage nymph.
E, Hypostome of second stage nymph. F, Hypostome and palpi of first stage nymph.
G, Hypostome and palpi of larva.

74

American Midland Naturalist Monograph No. 1

Ornithodoros yumatensis Cooley and Kohls, 1941
Plate 7, Figs. 35 and 36

1941b. Ornithodoros yumatensis Cooley and Kohls, original description, pp. 592-594,
with figures.

Sexes mildly dissimilar; nymphs and adults similar.

FEMALE

Body. — Oval, sides nearly parallel, bluntly pointed in front and broadly

A ?

B o"

C

D

Fig. 32. Ornithodoros parpen Cooley. A, Hypostome of female. B, Hypostome of
male. C, Hypostome of fourth stage nymph. D, Hypostome of third stage nymph.
E, Hypostome of second stage nymph. F» Hypostome and palpi of first stage nymph.
G, Hypostome and palpi of larva.

Cooley dc Kohls: Argasidae of N. America, etc.

75

rounded behind. Size of holotype female, 4.75 x 3.25. Size of allotype male,
3.8 x 2.25. Range of sizes of females, 5.6 x 3.5 to 4.1 x 2.25. Range of sizes
of males, 4.8 x 2.40 to 3.6 x 2.0.

Mammillae. — Irregular in shape, moderate in size and number and about
equal in sizes in median and peripheral areas on both dorsal and ventral sur-
faces; close together but not crowded. Individual mammillae have irregular
surfaces on top and at the sides; those on the posterior ventral area bent back-

Fig. 33. Legs of adults. Omilhodoros turicata (Duges) on the left and O. parfferi
Cooley on the right. Leg numbers indicated by Roman numerals.

76

American Midland Naturalist Monograph No. 1

ward mildly suggesting reptilian scales. Short hairs, few in number, present
mainly on the posterior and lateral marginal areas.

Discs. — Moderate in size, superficial or mildly depressed, not conspicuous.
On the venter present in lineal arrangement in the peranal, transverse post-
anal and median postanal grooves.

Legs. — Long and slender with their surfaces made irregular by very numer-
ous granulations. Subapical dorsal protuberances and dorsal humps absent.
Leg hairs numerous and small except on ventral and distal portions of the
tarsi where they are larger. Length of female tarsus I, 1.02; metatarsus, 0.72.
Length of female tarsus IV, 1.17; metatarsus, 0.96. Length of male tarsus I,
0.66; metatarsus, 0.51. Length of male tarsus IV, 0.87; metatarsus, 0.72.

IE I^Cfe^

£F3^ ■ ^

»^c^

CiO-7^^ Pi -pes*

Fig. 34. Legs of nymphs, Ornithodoros turicata (Duges) on the left, O. parpen on
the right. Leg. numbers indicated by Roman numerals. A, Fourth nymphal stage.
B, Third nymphal stage. C, Second nymphal stage. D, First nymphal stage.

Cooley & Kohls : Argasidaf. of N. America, etc.

Coxae. — Coxae I and II a little separated, all others contiguous.

Hood. — Indefinite.

Camerostome. — Indefinite.

Cheeks. — Large, about twice as long as wide, attached along one side, and
with a very few short hairs.

Capitulum. — The capitulum is protrusile and when extended reaches
beyond the anterior end of the body. When so extended the basis capituli is
seen to be twice as long as wide and its length is about equal to the length of
the soft membrane ("neck") which unites the capitulum with the body. Living
specimens may show the capitulum either extended or withdrawn. Specimens
preserved in alcohol usually have the capitulum withdrawn and give little
evidence of extensibility. Basis capituli with the surface granulated as on the
legs, and faintly wrinkled. Numerous hairs present on the sides on the anterior
portion (not visible when the capitulum is withdrawn) . Palpal article 1
swollen and with its surface granulated. Tip of hypostome reaching to articu-
lation between articles 3 and 4.

Hypostome. — With the sides subparallel, apex mildly notched. Denticles
small or faint; larger in the lateral files and progressively smaller toward the
median line. Length, female, 0.18 to 0.20; male, 0.16 to 0.17.

Folds. — Coxal and supracoxal fold? present, the latter reaching to the
cheeks.

Grooves. — Preanal, transverse postanal, and median postanal grooves
present. Dorso-ventral groove absent.

Sexual opening. — At the level of the interval between coxae I and II.

Eyes. — Absent.

Anus. — Large, in an elliptical pattern.

MALE

i
The male differs from the female principally in the hypostome. The male
hypostome is smaller and has relatively smaller denticles.

LARVA

Large, sub oval. Length (including mouth parts), 1.20, width, 0.50.
Dorsal plate, oval. Legs long, slender. Capitulum terminal, large, its length
equal to about half the entire length of the larva. Basis capituli large, wider
behind, visible from above. Hypostome slender, long, pointed apically, and
on a conical base which is about as long as the hypostome. Denticles apically,
3/;J), then 2/2; those of the lateral (marginal) files large, those of the median
files small. Length of hypostome, 0.24.

This species is rather closely related to O. brodyi Matheson but may be
separated 'by the dentition of the adult hypostome and the character of the
mammillae.

78

American Midland Naturalist Monograph No. 1

HOSTS AND DISTRIBUTION

Known only from bats or bat retreats.

California. — 17856, from rock crevices in mine tunnel, May 20, 1940, 15
miles northeast Yuma, Arizona, 1 male, 2 nymphs.

Fig. 35. Ornithodoros yumalensis Cooley and Kohls. A, Capitulum of larva, ventral
view. B, Capitulum of male, ventral view. C, Leg I of adult. D, Leg IV of adult.
E., Hypostome of male. F, Hypostome of female. G, Larva, dorsal view.

Cooley & Kohls: Argasidae of N. America, etc.

79

Arizona. — 17881, from rock crevices in Crystal Cave, June 2, 1940, 10
miles southeast Winkelrmn, numerous adults and nymphs, few larvae; 17882
from My otis velifer velifer, in Crystal Cave, June 2, 1940, 10 miles southeast
Winkelman, 6 adults and nymphs; 17046 from Myotis velifer, Picacho Peak
near Picacho, July 23, 1940, 1 larva (R. A. Flock).

Texas. — 17252, from rock crevices in Brehmer bat cave, September 11,
1940, near New Braunfels, 32 adults and nymphs.

Fig. 36. Distribution of Ornithodoros vumalensis Cooley and Kohls.

80 American Midland Naturalist Monograph No. 1

Ornithodoros brodyi Matheson, 1935
Plate 7, Fig. 37

1935. Ornithodoros brodyi Matheson, original description, pp. 351-352, with figures.
Sexes similar; nymphs and adults similar.

ADULT

Body. — Oval, sides nearly parallel, a little pointed anteriorly and rounded
behind; hood visible from above. Length, 6.0; width, 4.0 (Matheson).

Mammillae. — Moderate in size and number and about equal in size in the
median and peripheral areas; by their shape and crowded arrangement re-
sembling the scales of reptiles. Individual mammillae varying in size, flattened,
smooth and shining; some with a few punctations in the flattened surface. A
few hairs present arising from the integument between the mammillae. Mam-
millae smaller or absent on the supracoxal folds and in the intercoxal area;
in other ventral areas, as large as on the dorsum.

Discs. — Moderate in size, superficial.

Legs. — Long and moderate in size, micromammillated and with numerous
fine hairs. Subapical dorsal protuberance moderate on tarsus I, absent on all
others; dorsal humps absent on all legs. Length of tarsus I, 1.02; metatarsus,
C.78. Length of tarsus IV, 1.2; metatarsus, 1.14.

Coxae. — Small, with the surfaces granulated. Coxae I and II separated; all
others contiguous.

Hood. — Well developed, a little depressed below the anterior extension of
the dorsal body wall.

Cheeks. — Elongated, sides subparallel, reaching anteriorly beyond the line
of attachment for about one-third the length.

Camerostome. — Well developed and together with the cheeks affording
some protection for the mouth parts.

Capitulum. — Basis wider than long, with the surface irregular and micro-
mammillated. Palpi moderate in length; article 1 micromammillated and with
a knife-edge flange extending over the base of the hypostome, 2, 3, and 4 free,
smooth.

Hypostome. — Length, 0.125. Denticles in a 2/2 pattern with the laterals
larger and greater in number than those in the median files; corona large with
numerous fine denticles, truncate or faintly notched.

Folds. — Coxal and supracoxal folds distinct.

Grooves. — Dorso-ventral groove absent. Preanal, transverse postanal, and
median postanal grooves present, the latter extending only from the anus to
the transverse postanal groove.

Sexual opening. — Placed between coxae I.

Eyes. — Absent.

Anus. — In an elliptical pattern.

Our redescriptions are from specimens kindly sent by Dr. Matheson.
Descriptions of nymph and larva are quoted from Matheson (1935).

Cooley & Kohls: Argasidae of N. America, etc.

81

Nymph
From the material before me there appears to be two nymphal stages. Fully gorged
nymph (last nymphal stage) 4.5 mm. in length by 2 mm. in width. In general shape
nymph closely resembles adult with no marked difference except size and absence of
genital opening. Capitulum practically identical with that of adult except that at base
of hypostome there are two very long spines that occupy the same position as the two
short ones do in the adult.

Larva

Fully gorged larva practically spherical in shape, the long legs and capitulum pro-
jecting far beyond margins of body; length, exclusive of capitulum 2.3 mm.; capitulum
0.64 mm.; extreme width 2.3 mm. Hypostome of larva practically identical with that
of O. azteci (Fig. 4, b). Integument of partially gorged larva with beautiful arrangement
of fine parallel lines running in various directions on body.

HOSTS AND DISTRIBUTION

Types are in the collections of the Department of Entomology, Cornell
University, Ithaca, N. Y.

Matheson (1935) states that the larvae of this species were taken on the
short-tailed bat, Hemiderma per s picillatum aztecum (Saussure), March 30,
1931, by Mr. A. L. Brody and that the same collector found nymphs and
adults in crevices of the walls and ceilings of the Chilibrillo Caves, Panama.
Eggs were also found in association with the adults. No further collections of
the species have been reported.

c . s/ / //

Fig. 37. Ornilhodoros brodm Matheson. A, Leg I of adult. B, Leg IV of adult.
C, Hypostome of adult.

82 American Midland Naturalist Monograph No. 1

Ornithodoros TALAJE (Guerin-Meneville), 1849
Plate 8, Figs. 38 and 39

1849. Argas lalaje Guerin-Meneville, original description, pp. 342-343, with figures.
1885. Argas lalaje Guerin-Meneville: Megnin, redescribed, pp. 460-463; 466, 470-472.

with figures.
1896. Ornithodoros lalaje (Guerin-Meneville): Neumann, redescribed, pp. 34-37.
1901. Ornithodoros lalaje (Guerin-Meneville): Neumann, made O. rudis Karsch 1880,

a synonym, p. 259.

1907. Alectorobius lalaje (Guerin-Meneville) : Pocock, p. 189. New genus described,
O. lalaje type species.

1908. Ornithodoros talaje (Guenn) : Banks, redescribed, p. 19, with figures.

1908. Ornithodoros talaje (Guerin-Meneville): Nuttall, Warburton, Cooper and Rob-
inson, redescribed, pp. 59-61, with figures.

1911. Ornithodoros lalaje talaje (Guer.) : Neumann, redescribed, pp. 125-126, with
figures.

1930. Ornithodoros lalaje (Guerin-Meneville) : Hoffman, redescribed, pp. 161-163,
with figures.

1936. Ornithodoros talaje (Guerin-Meneville): Brumpt, pp. 1197-1199.

Sexes similar; nymphs and adults similar.

ADULT

Body. — Oval, a little pointed anteriorly, the curve of the posterior border
slightly flattened, sides nearly parallel. Margins more incurved opposite legs
I, II, and III than in most species. Size of female from 7.25 x 4.60 to 4.20 x
2.34. Size of male, 5.8 x 3.40 to 3.36 x 1.75.

Mammillae. — Large, close but not crowded, conical, with radial ridges on
the sides reaching nearly to the tips; each mammillae witt one to three sub-
apical excavations or pits. Mammillae larger at the sides and largest on the
posterior border; small on the venter and indefinite on the supracoxal folds.
Hairs absent on dorsal and ventral surfaces and even on the hood.

Discs. — Large, distinct, in large depressed areas which occupy much of the
median area of the dorsum. Discs on the venter present in lineal arrangement
in the preanal, transverse postanal, and the median postanal grooves and in
three depressions posterior to the transverse postanal groove.

Legs. — Small, moderate in length, with the surface micromammillated
Hairs small. Tarsus I with a mild subapical dorsal protuberance; absent on
all other tarsi. Dorsal humps on the tarsi absent. Length of tarsus I, 0.72;
metatarsus, 0.54. Length of tarsus IV, 0.87; metatarsus, 0 645.

Coxae. — Coxae I and II well separated; all other contiguous. Surfaces
micromammillated and with large excrescences.

Hood. — Small, reduced to a tongue-shaped flap which is bent downward
and in contact with the cheeks with which it forms an enclosure covering the
mouth parts. In lateral profile the hood and the anterior extension of the
dorsal body wall form a distinct notch.

Cooley & Kohls: Argasidae of N. America, etc.

83

Camerostome. — Indefinite, obscured by the large cheeks.
Cheeks. — Oval, large (relatively larger than in any other known species)
and when closed often covering the mouth parts. Surface irregular.

Capitulum. — Basis capituli wider than long with irregular transverse

Fig. 38. Ormthodoros talaje (Guerin-Meneville). A, Capitulum of larva, ventral
view. B, Larva, dorsal view. C, Hypostome of adult. D, Leg 1 of adult. E, Leg IV
of adult. F, Lateral view of anterior end of body. G, Hypostome and palpi of second
stage nymph.

84 American Midland Naturalist Monograph No. 1

wrinkles and micromammillae. Palpi moderate in size; article 1 micromam-
millated.

Hypostome. — Short and small, sides nearly parallel, notched apically.
Denticles arranged 2/2 with about four large ones in each file, and with those
in the median and lateral files about equal in size. Length about 0.245.

Folds. — Coxal and supracoxal folds present, the latter extending forward
tc near the hood.

Grooves. — Dorso-ventral groove absent. Preanal, transverse postanal, and
median postanal grooves present, the latter terminating at the transverse post-
anal groove.

Sexual opening. — Between coxae I.

Eyes. — Absent.

Anus. — In an oval frame.

LARVA

Unfed larva sub-oval in shape; basis capituli triangular in dorsal view.

Palpi very long, slender, and in life appressed against the hypostome. Length

of body, including capitulum, 0.66. Length of capitulum (in ventral view),
0.33.

Hypostome. — Very long and slender; denticles in a 2/L> arrangement with
the marginal denticles much larger than those in the next file which are not
easily seen, even when mounted in balsam. Length about 0.21.

O. talaje is rather closely related to O. keH?yi and O. concanensis, two
bat-infesting species. From kelleyi it may be distinguished in the adult and
late nymphal stages by a notch just above the hood when viewed in lateral
profile. In these stages, concanensis is smaller, proportionately longer, more
rounded on the posterior margin, the marginal projection opposite coxa II less
pronounced, the hood shorter, and the depressed areas occupied by the discs
less extensive. The larvae of concanensis are unknown. The larvae of talaje
have the hypostome broad, with sides rounded, while in kelleyi the
hypostome is long and narrow and the sides are nearly straight. Further, in
talaje the base of the hypostome (from the insertion of the palpi to the denti-
cles) is shorter than in kelleyi.

DISTRIBUTION

The range of O. talaje extends from California and Kansas to Argentina.
The types were from Guatemala.

In the United States it is known from California, Arizona, Nevada,
Kansas, Texas, and Florida. Hoffman (1930) stated that in Mexico this tick
is a native of the hot country and that it is found on the Peninsula of Yuca-
tan, Campeche, Tabasco, Chiapas, the Isthmus and State of Vera Cruz,
extending along the Gulf Coast toward Texas; on the Pacific Coast the species

Cooley & Kohls: Argasidae of N. America, etc.

85

thus far has not been found except to the north in the region of the Isthmus.
According to Dunn (1933) the species apparently is widely distributed in
Panama.

HOSTS

This species has been collected from a wide variety of hosts. In the United
States it has been found only on wild rodents or in association with them.

Fig. 39. Distribution of Oriiilhocloros lalaje (Guerin-Meneville) .

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88 American Midland Naturalist Monograph No. 1

The rodents concerned were Dipodomys sp., Citellns sp., and Neotoma sp.,
including N. albigida and N. floridana baileyi. While reported from houses
in Minnesota (Riley, 1935), Wisconsin (Herrick, 1935), and New York
(Matheson, 1931) the species actually concerned was O. kelleyi Cooley and
Kohls 1941.

In Mexico, Hoffman (1930) stated that the natural hosts are small, wild
rodents and probably other mammals which inhabit holes and subterranean
nests, and doubtless with the intervention of rats it is found in the houses
of man, whom it attacks with preference when once established in the habita-
tion. As a true domestic parasite it has been observed only in the south of the
Republic and never north of the region of San Andres Tuxtla. In the northern
part of the State of Vera Cruz and in Tamaulipas it is quite rare and is found
almost exclusively on wild animals (rodents). He stated that the bites were
painful but not to the extent of those by O. turicata.

Among the hosts in Panama listed by Dunn (1933) were Mus alexan-
drmus, M. norvegicus, M. rattus, several species of monkeys, dogs, cats,
chickens, and a snake, Epicrates cenchris. The ticks found were principally
larvae. Dunn stated that adults were seldom observed in dwellings and he
believed that they rarely attacked man.

Ornithodoros concanensis Cooley and Kohls, 1941
Plate 8, Fig. 40

1941 (c). Ornithodoros concanensis Cooley and Kohls, original description, pp. 910-911,
with figures.

Sexes similar; nymphs and adults similar.

ADULTS

Body. — Sub-oval, sides nearly parallel, anterior margin bluntly pointed,
posterior margin rounded. Marginal projection opposite coxa II moderate.
Size of female, 5.5 x 3.0; male, 4.20 x 2.40.

Mammillae. — Large, close but not crowded; sides and tops with a mixed
pattern of ridges with but little appearance of radiation; the majority have one
or two circular or crescentic pits on top, with a short, indistinct hair present
in some of the pits especially in the peripheral areas. Sizes about equal in
lateral and median areas of dorsum, slightly larger on posterior margin; on the
venter somewhat smaller than those on the dorsum, but larger on the posterior
margin; absent on supracoxal folds.

Discs. — Those on the dorsum depressed and with edges elevated. Venter
with the discs in lineal arrangement in the preanal and median postanal
grooves; present also in three depressions caudad of the transverse postanal
groove.

Cooley & Kohls: Argasidae of N. America, etc.

89

Legs. — Moderate in length and in diameter; surfaces micromammillated.
Tarsus I with a mild subapical dorsal protuberance; absent on all others.
Dorsal humps absent on all tarsi. Length of tarsus I, 0.6; metatarsus, 0.45;
length of tarsus IV, 0.72; metatarsus, 0.6.

Coxae. — Coxae I and II well separated; all others contiguous. Surfaces
micromammillated and also with mild excrescences.

Hood. — Limited to a short, apical elevation which is separated from the
anterior extension of the dorsal body wall by a depressed line.

Cheeks. — Oval, with the anterior end wider and free; attached 'by the side
of the narrower portion.

Fig. 40. Ornilhodoros concanensis Cooley and Kohls. A, Hypostome of adult. B,
Hypostome and palpi of second stage nymph, ventral view. C, Leg I of adult. D, Leg
IV of adult.

90 American Midland Naturalist Monograph No. 1

Capitulum. — Basis capituli about as wide as long, surface with transverse
wrinkles and numerous micromammillae; with a pair of fine hairs posterior to
the posthypostomal hairs and a group of smaller hairs on each side behind.
Article 1 of palpus micromammillated.

Hypostome. — Wider beyond the middle, moderately long, notched apical-
ly. Denticles in a - /2 pattern with about four in each file and limited to the
distal one-third. Length about 0.21.

Folds. — Coxal and supracoxal folds present. Supracoxal fold reaching
anteriorly to the hood.

Grooves. — Preanal, transverse postanal and median postanal grooves
present, the latter terminating at the transverse postanal groove. Dorso-ventral
groove absent.

Sexual opening. — Placed at the level of the intervals between coxae I
and II.

Eyes. — Absent.

Anus. — In an oval frame.

This species resembles talaje and kdleyi. From talaje it is distinguished by
being smaller, proportionately longer, more rounded on the posterior margin,
the marginal projection opposite coxa II less pronounced, the hood shorter,
and the depressed areas occupied by the discs are more shallow and less exten-
sive. From kelleyi it is distinguished by being proportionately shorter and
having the cheeks larger, as well as by having the hypostome wider beyond
the middle and the palpi having more hairs.

HOSTS AND DISTRIBUTION

This species is known from bat retreats in Arizona and Texas as follows:
Arizona. — 17875, from rock crevices in a bat-inhabited mine tunnel, May
25, 1940, Las Guijas, Pima County, 2 males, 1 nymph.

Texas. — 17261, from guano and rock crevices in bat cave, Sept. 16, 1940,
near Concan, Uvalde County, several adults and nymphs.

Cooley & Kohls: Argasidae of N. America, etc. 91

Ornithodoros stageri Cooley and Kohls, 1941

Plate 9. Figs. 41 and 42

1941 (b). Ornithodoros stageri Cooley and Kohls, original description, pp. 589-592, with
figures.

Sexes dissimilar; adults and nymphs dissimilar.

FEMALE

Body. — Oval, wider behind, approaching a point in front; tips of the palpi
often visible from above. Range of sizes from 5.3 x 3.4 to 3.3 x 2.4.

Mammillae. — Relatively large, few in number, and not crowded; irregular
in shape, their tops convex, smooth, often with radial ridges on their bases.
A few have a single, faint pit on the top which may or may not have a short,
fine hair.

Discs. — Distinct, depressed, large, mostly circular. Present also on the
venter where they are in a lineal arrangement in or near the preanal, trans-
verse postanal, and median postanal grooves.

Legs. — Moderate in length and size; surface nearly smooth, shining, with
hairs moderate in number and in length. Subapical dorsal protuberances and
dorsal humps absent. Length of female tarsus I, 0.66; metatarsus, 0.48. Length
of female tarsus IV, 0.84; metatarsus, 0.75. (Length of male tarsus I, 0.45;
metatarsus, 0.3. Length of male tarsus IV, 0.63; metatarsus, 0.54.)

Coxae I and II a little separated; all others contiguous.

Hood. — Negligible or absent.

Camerostome. — Faintly indicated.

Cheeks. — Small, and somewhat variable in shape; usually oval and attached
along one side of the broader portion. (Absent in the male.)

Capitulum. — Basis capituli about as wide as long; surface irregular with
transverse wrinkles. With two or three long, barbed hairs on each side on the
lateral walls a short distance from the insertion of the palpi, and a pair of
hairs back of the posthypostomal hairs, which are more separated and smaller.
Palpal article 1 with its surface a little roughened and with a narrow flange
projecting over the base of the hypostome. Articles 2, 3 and 4 free.

Hypostome. — Notched in both sexes, narrower in the female than in the
male, with the principal denticles arranged 2/o with the four files approximate-
ly equidistant. Length in both sexes about 0.22.

Folds. — Coxal and supracoxal folds present, the latter continuous from
side to side in front.

Grooves. — Preanal, transverse postanal, and median postanal grooves
present; dorso-ventral groove absent.

Sexual opening. — Placed at the level of the intervals between coxae I
and II.

92

American Midland Naturalist Monograph No. 1

Eyes. — Absent.

Anus. — In an elliptical pattern.

MALE

The male differs from the female as follows: The male lacks the cheeks
and has the hypostome wider and the denticles smaller with the two principal
files separated by two shorter files of smaller denticles on each side of the
median line.

E

Fig. 41. Ornithodoros slageri Cooley and Kohls. A, Male hypostome. B, Female
hypostome. C, Leg I of male. D, Leg IV of male. E, Capitulum of larva, ventral view.
F, Larva, dorsal view.

Cooley & Kohls: Argasidae of N. America, etc.

93

NYMPH

Early stage nymphs show no evidence of cheeks. Late stage nymphs show
them much smaller than in the female. Smallest nymphs measure 1.35 x 0.9®.

LARVA

Short-oval, moderate in size. Length including hypostome, 0.765; width,
0.48. Dorsal plate shining and with faint pits (visible with reflected light in
unmounted specimens). Legs about as long as the body. Capitulum terminal
and visible from above; basis broad. Hypostome lacking the long conical base
found in some species (see figure 35, A); sides a little converging anteriorly,
bluntly pointed apically. Denticles apically Aj± then 3/3, and finally 2/2 at
the base; those of the lateral files large, and those of the median files small.
Length of hypostome about 0.2.

HOSTS

Ornithodoros stageri is known only from bats and bat-inhabited caves and
mines. Adults have been found on Myotis velifer and Tadarida mexicana.
Adults and nymphs have been found in abundance in rock crevices in bat
caves and mine tunnels where bats hung and on guano deposits on the floors.

The species feeds promptly on man when it has opportunity as evidenced
by Dr. Bequaert (in letter), and both of the present authors. One of us
(G. M. K.) and his assistant received several bites while collecting the species
in bat retreats.

Fig. 42. Distribution of Ornithodoros stageri Cooley and Kohls.

94 American Midland Naturalist Monograph No. 1

distribution

California. — 17859, bat guano in Senator Mine, May 21, 1940, 21 miles
northeast of Yuma, Arizona, several adults and nymphs.

Arizona. — 17868, rock crevices in mine tunnel, May 24, 1940, Picacho
Peak near Picacho, numerous adults and nymphs; 17034, mine tunnel, July
23, 1940, Picacho Peak, near Picacho, 3 adults (R. A. Flock); 17166, Myotis
relifer, July 1940, Picacho Peak, near Picacho, 1 adult (Dr. J. C. Bequaert) .

Oklahoma. — 17017, bat cave, July 2, 1940, near Freedom, 2 nymphs, 1
larva (Dr. D. E. Howell); 17221, bat cave, August 22, 1940, Selman Ranch,
near Freedom, numerous adults and nymphs; 17393, bat cave, October 26,
1940, Weatherford, 1 d (Dr. D. E. Howell).

Texas.— 17798, Ney Cave, August 5, 1939, 20 miles north of Hondo, 1
male, 1 female, 1 nymph (Kenneth E. Stager); 17258, Ney Cave, September
14, 1940, 20 miles north of Hondo, numerous adults and nymphs; 17259,
Tadarida mexicana, in Ney Cave, September 14, 1940, 20 miles north of
Hondo, 2 adults; 17254, bat cave, September 12, 1940, near Bracken, about
150 adults and nymphs.

Cooley & Kohls: Argasidae of N. America, etc. 95

Ornithodoros dyeri Cooley and Kohls, 1940
Plate 9, Figs. 43 and 44

1940. Ornithodoros dyeri Cooley and Kohls, original description, pp. 925-928, with
figures.

Sexes similar; nymphs and adults similar.

ADULT

Body. — Length, 5.5, width, 2.1. Color (living) yellow-brown. Elongated,
sides parallel, anterior end pointed, posterior end rounded. Flattened on top
and marginated. Viewed in lateral profile the body is nearly straight. Entire
dorsum bounded by two continuous ridges which are essentially parallel and
with a third (inner) incomplete ridge on each side, plainer in early stage
nymphs. Ridges less elevated in the later stage specimens. Within the ridges
the surface is irregular due to short ridges and subcircular elevations. In the
anterior dorsal area is a distinct median, smooth hump raised above the level
of the lateral margins, and anterior to it a deep depression near the parallel
marginal ridges.

Mammillae. — Indefinite or absent though there are present small, irregular
elevations on the surface.

Hairs. — A few very fine, short hairs are scattered on dorsal and ventral
surfaces.

Legs. — Moderate in length, small, micromammillated. Fine, short hairs
scattered over the legs; those on ventral surface of the tarsi larger and in two
parallel rows. All tarsi without subapical dorsal protuberances or dorsal humps.
Stalk of the pulvillus and claws progressively longer on legs I to IV. Femur
of leg IV notably long — longer than femur III. Length of femur IV, includ-
ing trochanter, 1.08; that of leg III, 0.72. Length of tarsus I, 0.54; metatarsus,
0.48. Length of tarsus IV, 0.69; metatarsus, 0.60.

Coxae. — Coxae I and II separated; all others contiguous. Surfaces irregular
and micromammillated. Hairs absent except a row near the articulation with
trochanter on coxae II, III, and IV.

Hood. — A true hood absent but with a median ridge extending from the
mouth parts to the anterior margin.

Cheeks. — Pyriform, attached along one side, with the broader end in front
and together with the end of the ventral median ridge forming protection for
the mouth parts.

Capitulum. — Basis large, broader than long; surface having micromammil-
lae and distinct transverse wrinkles. Surface of palpal article 1 irregular and
with a narrow, knife-edge flange on the median side overlapping the base of
the hypostome; other articles free.

Hypostome. — Short, broad, truncate, with sides nearly parallel; posthypo-

96

American Midland Naturalist Monograph No. 1

stomal hairs long, faintly barbed, reaching the tip of the hypostome. There is
also a pair of shorter postpalpal hairs. Denticles short, U-shaped, arranged
5/5, covering about the distal two-thirds of the hypostome. Length, 0.21.

Folds. — Coxal and supracoxal folds present and also a supplemental pat-
tern of folds as shown in figures 43, E. Supracoxal folds not joined anteriorly.
In addition to the remarkable pattern of folds in this species is a fold or ridge
above the supracoxal fold opposite legs III and IV, which is 'bent downward

Fig. 43. Ornilhodoros Jperi Cooley and Kohls. A, Hypostome. B, Palpus. C, Leg
I. D, Leg IV. E, Diagram of venter.

Cooley & Kohls : Argasidae of N. America, etc. 97

posteriorly and becor.-.es an arc of a circle, the radial center of which would
be the insertion of leg IV. The top of this arc is smooth while the surface of
all other ridges is irregular. In living specimens leg IV in its movements is in
contact with the smooth ridge, which is enough elevated to protect the spiracle
placed just below it. Leg IV comes in contact with this smooth arc at about
the trochanter.

While in most Argasidae the spiracle is usually placed between coxae III
and IV, in this species it is situated well back of coxa IV.

Grooves. — Only the deep, short, transverse postanal groove is present.

Sexual opening. — Placed at about the level of the posterior ends of coxae
I. V-shaped depression, including the opening, present in the female, absent
in the male.

Eyes. — Absent.

Anus. — Small, in an oval pattern, distant from the transverse postanal
groove.

In this species the nymphs are remarkable in having a V-shaped depres-
sion in the position where the sexual opening is to appear, making it singular-
ly difficult to distinguish between nymphs and adults. The original descriptions
included both adults and nymphs, the authors believing that they had both
before them. Dr. Mazziotti has recently sent us a male and a female specimen,
the male being the first one we have seen. These specimens show the different
male and female sex opening quite clearly and make it evident that the speci-
mens before us at the time the species was described were nymphs. The V-
shaped depression is lacking in the male, while it is present, as in the nymph,
in the female.

The above description is made from three large females from Mr. Flock
(17189) and a male and female from Dr. Mazziotti. They differ from the
nymphs in having the continuous submarginal ridge less elevated, the dorsal
and ventral surfaces smoother and in having more spur ridges in the pattern
of the folds on the venter. The smallest nymphs we have seen are shorter in
proportion to the width than in adults and measure 1 .98 x 1.11.

This remarkable tick is easily separated from other known species by its
elongated body, the parallel marginal ridges, and the pattern of ridges on the
venter as well as by other characters.

HOSTS

While this species has never been actually taken from bats, it has been
found only on bat guano or on the walls of bat caverns.

DISTRIBUTION

Arizona. — 16083, from bat guano from mine tunnel, November 3, 1939,
Picacho Peak, near Picacho, 28 nymphs; 17863, from bat guano and rock
crevices in mine tunnel, May 23, 1940, Picacho Peak, near Picacho, several
nymphs; 17875, rock crevices in mine tunnel, May 28, 1940, Las Guijas, 1

98

American Midland Naturalist Monograph No. 1

nymph; 17189, Tucson Mountains, September 16, 1940, 3 females (R. A.
Flock).

California. — 17856, rock crevices in mine tunnel, May 20, 1940, 15 miles
northeast of Yuma, Arizona, 3 nymphs; 17839, rock crevices in bat cave,
June 14, 1940, 16 miles north of Needles, several nymphal cast skins.

Mexico. — Abandoned mine harboring numerous bats, 5 kilometers west of
Coquimatlan, Colima, November, 1940 (Mazziotti, 1941).

Fig. 44. Distribution of Ornithodoros dyeri Cooley and Kohls.

Cooley dc Kohls: Argasidae of N. America, etc. 99

Ornithodoros amblus Chamberlin, 1920
Plate 10. Fig. 45

'920. Ornithodoros amblus Chamberlin, original description, pp. 43-44, with figure.

Sexes similar; adults and nymphs similar.

ADULT

Body. — Su'boval, sides nearly parallel, anterior end a little pointed, poster-
ior end broadly rounded. Mouth parts and hood not visible from above. Size,
6.45 x 4.8 to 5.0 x 3.25. Chamberlin (1920) gave the length as 7.2.

Mammillae. — Mammillae on the dorsum large, hemispherical, close but
not crowded, those on the posterior border largest; tops convex with a few very
small pits. Less elevated on the venter and some are oval or irregular in shape;
absent or faint on the supracoxal folds.

Hairs. — Hairs few in number, more numerous on the anterior border and
placed between the mammillae.

Discus. — Large and noticeable, mostly circular, a little depressed. On the
venter present only in the grooves and in a median and two lateral depressions
posterior to the transverse postanal groove.

Legs. — Moderate in length and size in the adults, relatively larger in the
small nymphs, articles flared at their distal ends; trochanter thickened. Haller's
organ notably distant from the distal end. Subapical dorsal protuberances and
dorsal humps absent on all legs. Tarsus IV, long and tapering. Hairs few in
number and of moderate length. Length of tarsus I, 0.78; metatarsus, 0.6.
Length of tarsus IV, 1,2; metatarsus, 0.96.

Coxae. — Coxae I and II separated; all other coxae contiguous.

Hood. — Very short and continuous with a short median elevation that
connects it with anterior dorsal margin.

Camerostome. — Deep and terminating anteriorly in the short hood.

Cheeks. — Small, narrow and attached along their greatest dimension; not
overlapping the mouth parts.

Capitulum. — Basis capituli wider than long; with a group of erect hairs
on each side behind. Surface rough and with transverse wrinkles. Palpal article
1 swollen and with its surface similar to that of the basis capituli; with a knife-
edge flange overlapping the base of the hypostome. Articles 2, 3, and 4 free
and smooth.

Hypostome. — Sides nearly parallel; apex notched. Principal denticles in a
2/2 arrangement. Small denticles numerous in the corona and with a few
small ones suggesting a third file on each side. Posthypostomal hairs reaching
to about three-fourths the length of the hypostome.

Folds. — Coxal and supracoxal folds present.

Grooves. — Prenanal, transverse postanal and median postanal grooves
present, broad and deep; the median postanal terminating at the transverse
postanal groove. Dorso-ventral groove absent.

Eyes. — Absent.

Anus.- — In a nearly circular frame.

100

American Midland Naturalist Monograph No. 1

O. amblus resembles O. capensis from Africa, but the former is readily
separated by several characters including the longer legs, the more numerous
and more pointed denticles of the hypostome and in having the mammillae
more nearly hemispherical on top and more regular at their bases.

hosts and distribution

This species is known from guano islands off the coast of Peru, South
America. Chamberlin (1920) states: "Dr. Murphy notes that these ticks are
a type 'extremely abundant on all the Peruvian guano islands. They live in
the soil and the guano.' " Larval specimens were taken from the plumage of an
adult penguin (Spheniscus hwnboldti) .

Murphy (1925), p. 246, states:

Moreover, the ticks do not confine their attacks to the birds, but at times bite the
skin of the native Indian laborers, usually upon the feet or legs, producing sores which
are apt to be persistently troublesome.

Mr. Murphy in the same book (p. 245) states:

The most significant fact concerning the ground spiders is that they appear to subsist
largely upon the native ticks, which are blood-sucking parasites of the birds. The spiders'
dens are sometimes filled with shriveled remains of the ticks, for, since spiders eat only
the body juices of their quarry, they take a remarkably heavy toll. Thus they become
the chief natural agency in restricting the numbers of an important enemy of the guano
birds.

While this species has not been collected in nature in North America, it
is reported as very abundant on the guano islands and may be readily intro-
duced with shipments of bird guano. In a recent letter Bequaert states: "It
has been found recently in a port of the United States, in a shipment of
guano."

^^§?

Fig. 45. Ornithodoros amblus Chamberlin. A, Hypostome of female. B, Leg I of
female. C, Leg IV of female.

Cooley & Kohls: Argasidae of N. America, etc. 101

Ornithodoros rudis Karsch, 1880
Plate 10, Fig. 46

1880. Ornithodoros rudis Karsch, original description, p. 141.
1901. Ornithodoros talaje (Guerin-Meneville) : Neumann, p. 259.

1921. Ornilhodorus venezueliensis [sic] Brumpt, in Neveu-Lemaire. p. 348.

1922. Ornilhodorus venezuelensis Brumpt: Brumpt, redescribed, pp. 773-775, with
figures.

1928. Ornithodoros venezuelensis Brumpt: Ruge, redescribed, pp. 406-409, with figures.
1936. Ornithodoros venezuelensis Brumpt: Brumpt, redescribed, pp. 1203-1205, with

figures.
1936. Ornilhodorus migonei Brumpt, pp. 1209-1210.

Sexes similar; adults and nymphs similar.

ADULT

Body. — Long-ovate, sides nearly parallel, anterior end pointed. Size of
female, 5.5 x 3.0 to 3.95 x 2.0; male, 4.2 x 2.2 to 3.6 x 2.1.

Mammillae. — Moderate in number, convex and shining on top and with
radiating striae on their bases; with one or more crescentic pits on the margin
of each. On both the dorsal and ventral surfaces, short fine hairs arise from
the crescentic pits at the sides of the mammillae, more abundant on the sides
and at the posterior end. Mammillae a little larger in the marginal areas and
largest of all at the posterior margin; smaller and less definite on the venter.

Discs. — Superficial and faintly differentiated from the mammillae; in a
symmetrical pattern on the dorsum but on the venter visible only on the
median postanal groove.

Legs. — Moderate in length and size, with numerous short hairs, those on
the tarsi in two ventral rows. Surface of the legs micromammillated. Subapical
dorsal protuberance small on tarsus I, very small or absent on tarsus IV;
dorsal humps absent. Length of male tarsus I, 0.585; metatarsus, 0.42. Length
of male tarsus IV, 0.54; metatarsus, 0.435.

Coxae. — Short and small, I and II a little separated but others contiguous;
surfaces micromammillated.

Hood. — Continuous with anterior extension of the dorsal body wall; apical-
ly bent ventrad and together with the cheeks affords some protection for the
mouth parts.

Cheeks. — Reniform with the free edges irregular.

Capitulum. — Basis a little wider than long; surface irregular, micromam-
millated, with faint transverse wrinkles and with short, fine hairs on each
side behind. Palpi micromammillated; article 1 with a narrow flange extend-
ing over the base of the hypostome.

Hypostome. — Moderate in length, sides nearly parallel, apex notched.

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American Midland Naturalist Monograph No. 1

Denticles arranged 2/2 with only about four large ones in each file. Denticles
placed on about the distal half. Length, 0.22 to 0.30.

Folds. — Coxal and supracoxal folds present.

Grooves. — The dorso-ventral, preanal, transverse postanal and median
postanal grooves present.

Sexual opening. — At the level of the intervals between coxae I and II.

Eyes. — Absent.

Anus. — In an elliptical frame.

HOSTS AND DISTRIBUTION

The literature of this species reveals no hosts other than man and domes-
tic fowls.

Fig. 46. OrnilhoJoros rudis Karsch. A, Hypostome of adult. B, Leg I of adult.
C, Leg IV of adult. D, Capitulum of larva, ventral view. E, Larva, dorsal view.

Cooley & Kohls: Argasidae of N. America, etc. 103

The type specimens came from "Nova Granada.1' Brumpt's specimens of
venezuelensis were collected in Colombia and Venezuela; those of migonei
were from Paraguay.

We have seen living specimens of O. migonei which came originally from
Paraguay, and the Rocky Mountain Laboratory also has preserved specimens
from Paraguay sent by Dr. Brumpt. These materials have supplied an abun-
dance of specimens of all stages which have been compared with living and
preserved specimens of O. rudis Karsch. The comparison shows no specific
differences in adults, nymphs, or larvae and we find it necessary to make
migonei a synonym of rudis, which has priority.

Dunn (1927) considers "this tick (O. venezuelensis) to be the one com-
monly found in houses in various parts of Panama, Colombia, and Venezuela,
and possibly in some of the neighboring republics, and that this species accepts
man as its preferred host." In 1921 he received specimens from the native vil-
lages of Chorrera and San Juan in the Republic of Panama. In 1924, a total
of 4,880 specimens was collected in Colombia from 68 houses in 20 villages,
towns, and cities in various parts of the republic.

We have seen specimens from Panama, Canal Zone, Department of
Boyaca in Colombia, and Venezuela, as well as from Paraguay.

Ornithodoros dunni Matheson, 1935
Plate 11, Fig. 47

1935. Ornithodorus dunni Matheson, original description, pp. 347-349, with figures.
Sexes similar; adults and nymphs similar.

ADULT

Body. — Oval, a little pointed in front, wider and rounded behind; much
flattened. Hood and mouth parts not visible from above. Matheson (1935)
gives as the length 3.4; extreme width, 2.0. A single male before us measures
2.79 x 1.71.

Mammillae. — Relatively few in number, those in the margins smaller. It
is difficult to distinguish 'between the mammillae and discs in the posterior
areas near the margin on both dorsal and ventral surfaces. The individual
mammillae are smooth and shining, only a little elevated and convex on top,
usually with one or more small pits. Inconspicuous short hairs visible on the
hood and around the entire margin on both dorsal and ventral surfaces.

Discs. — Large and conspicuous as elevated, shining areas which occupy
much of the dorsal surface; a few are present also on the ventral surface, where
they are in lineal arrangement in the grooves.

Legs. — Short and heavy with numerous short hairs. Length of male tarsus

104 American Midland Naturalist Monograph No. 1

I, 0.25; metatarsus, 0.25. Length of male tarsus IV, 0.42; metatarsus, 0.32.
With a mild subapical dorsal protuberance on tarsus I, absent on all others.
Dorsal humps absent on all tarsi.

Coxae. — All coxae contiguous.

Hood. — This very small hood is separated from the anterior projection of
the dorsal body wall.

Cheeks. — Small, oval and free on their anterior ends.

Capitulum. — Basis capituli wider than long, with the surface granulated.
Article 1 of the palpus with a knife-like edge on the median side extending
over the base of the hypostome; articles 2, 3 and 4 free.

Hypostome. — Short, with the sides nearly parallel; notched apically. Den-
ticles arranged 2/2 covering the distal half, with 3 or 4 large teeth in each file.
Length about 0.18.

Folds. — The supracoxal fold extends from the hood to behind coxa IV.
Coxal fold extends from coxa III to near the posterior margin.

Grooves. — Transverse postanal groove present, all others absent.
Sexual opening. — Placed between coxae I.
Anus. — In an elliptical pattern.

NYMPH

The number of nymphal stages is not in evidence in the materials avail-
able. Matheson (1935) mentions two known nymphal stages and gives meas-
urements as 1.5 and 2.0, respectively.

LARVA

Of the larva, Matheson (1935) gives the following:

Newly hatched larva 0.64 mm. from tip of the hypostome to posterior end of body;
remarkably elongated hypostome (Fig. 4, a) 0.2 mm. in length; width of body 0.28 mm
Hypostome narrowly elongate, sharply pointed at apex with a dentition of 2|2; lateral
teeth very long and apical portion lacks denticles; base of hypostome with two short
spines; chelicerae slightly longer than hypostome; digit quite similar to that of adult
though more elongate; palps free and with few spines. Full gorged larva extremely
distended, 1 .28 mm. in length.

HOSTS AND DISTRIBUTION
Matheson states:

This tick was collected in the larval stage on the Little Bull Bat, Dirias albiventer
minor (Osgood), which was captured in a mango tree in the yard of the Panama
Hospital, Panama City, on March 17, 1931. The bat was placed in a bag and during
the next few days 32 larvae dropped, full engorged. The larvae molted, and were
carried through the nymphal stages to the adult. Larvae of this tick were also taken
on the same species of bat at Summit Canal Zone, on Sept. 30, 1932. This tick appears
to be restricted to this one species of bat.

Cooley & Kohls: Argasidae of N. America, etc. 105

In addition to Panama and the Canal Zone, the known distribution as
given by Matheson, we have received one female found "living in tree holes
with bats" collected by Mr. R. Damasceno, and kindly sent by Dr. Emmanuel
Dias. The specimen was taken on Marajo Island (mouth of Amazon), June
1941, Belem, Para, Brazil.

c

Fig. 47. Ornithodoros dunni Matheson. A, Leg I of nymph. B, Leg IV of nymph.
C, Hypostome and palpi of larva, ventral view. D, Hypostome of adult.

106 American Midland Naturalist Monograph No. 1

Ornithodoros viguerasi Cooley and Kohls, 1941
Plate II, Fig. 48

1941a. Ornithodoros viguerasi Cooley and Kohls, original description, pp. 396-399,
with figures.

Nymphs and adults dissimilar; whether sexes are similar or dissimilar is
unknown.

ADULT

Body. — Sub-oval, broadest at about the middle, narrowly rounded behind,
and a little pointed in front. Broad anterior point bent ventrad where it meets
the anterior extensions of the supracoxal folds, from which it is separated by
a depressed line. Size 3.12 x 2.1.

Mammillae. — Numerous and of various sizes and forms. Those in the
median dorsal area large, crowded, of irregular shapes, flattened on top and
each with a central pit which may bear a hair; those in the anterior marginal
areas on the dorsum much as in the median area but smaller and with hairs
more numerous; those in the dorsal posterior margin elongated, arranged
radially, and only slightly elevated. Venter with a transverse band just poster-
ior to coxae IV, extending from side to side, in which the mammillae are
unique among mammillae of all known species. This band extends over the
l?teral margins becoming visible from above and also has extensions on the
supracoxal folds and in the median area between the coxae. These mammillae
at the margins are elevated, columnar, about twice as high as their diameters,
■convex on top and each with a single fine hair; height of the mammillae dimin-
ishing progressively from margins to the median area where they are only
slightly elevated and yet retain a sharp margin of the convex top. The surface
of the derm between these modified mammillae is finely and evenly pebbled.

Discs. — Large, distinct, a little depressed and with their surfaces shining;
not apparent on the venter.

Legs. — With numerous barbed hairs which are shorter and more barbed
on the dorsal surfaces, larger and less barbed on ventral surface. Surface shin-
ing and with transverse wrinkles. Subapical dorsal protuberances and dorsal
humps absent. Length of tarsus I, 0.42; metatarsus, 0.3. Length of tarsus IV,
0.50; metatarsus, 0.4.

Coxae. — Coxae I and II a little separated; all others contiguous.

Hood. — No well developed hood is present though the anterior projection
of the dorsal body wall is bent downward and resembles a hood.

Cheeks. — Conical projections about as long as the diameter at the base
and bearing a few short, fine hairs.

Capitulum. — Basis capituli with the surface irregular but shining; with a
median longitudinal elevation. Only the anterior portion of the basis capituli
is visible in the type female and it is not evident whether the capitulum is

Cooley & Kohls: Argasidae of N. America, etc. 107

protrusile in the adult as in the nymph. Palpal article 1 long, about as long
as articles 2 and 3 combined. Article 4, conical.

Hypostome. — Small, flattened, in the shape of an inverted V; denticles
not evident when examined in situ. Posthypostomal hairs placed far behind
the insertion of the hypostome and very long. Length from hairs to apex, about
0.18. (Described in situ.)

Folds. — Coxal and supracoxal folds present. Coxal fold with a shining,
sclerotized plate bordering coxae II, III and IV. Suprocoxal fold similarly
sclerotized from opposite coxa III to the anterior end. There is also a similar
plate anterior to the sexual opening between coxae I.

Grooves. — Preanal groove distinct at the sides but interrupted in the
middle. Transverse postanal groove deep and continous from the lateral mar-
gins. Median postanal groove deep and terminating at the transverse postanal
groove. Dorso-ventral groove absent.

Sexual opening. — At the level of the intervals between coxae I and II.

Eyes. — Absent.

Anus. — In an elliptical pattern.

NYMPH

Body with shape much as in the adult. The late stage nymphs measure
3.0 x 1.95 and differ from the adult in possessing a definite sclerotized plate
posterior to the position of sexual opening and in lacking a transverse plate
between coxae I anterior to the sex opening. The late nymphs differ from the
adult also in having the mammillae in the margins of the transverse ventral
band only about as elevated as the other mammillae of this species and in
the first nymphal stage the band of modified mammillae is lacking. The
nymph lacks the definite modification of mammillae in the median posterior
margin of the dorsum as described in the adult.

In the nymphs the capitulum is very long, protrusile, and attached by a
soft "neck." When so extended the capitulum reaches beyond the anterior
point of the body.

Hypostome. — Pointed apically; the two posthypostomal hairs arising from
the tumescent base. The faint denticles scarcely visible even in mounted speci-
mens; those on the margins larger. Length about 0.15.

Nymphs are misleading in appearing to have the sex opening present. This
is due to the presence of a sclerotized semi-circular flap and just posterior to
it a large, smooth sclerotized plate. Coxal folds have oval sclerotized plates
bordering on coxae II, III and IV, but the supracoxal folds are mammillated
and lack the long, definite sclerotized plates found in adults.

LARVA

Engorged larva (not necessarily fully fed) oval, widest in front of the
middle, bluntly pointed in front, rounded behind. Length (not including the
mouth parts) 2.64; width, 1.95. Larval integument striated as in other species

108

American Midland Naturalist Monograph No. 1

but lacking a dorsal plate. Leg I distant from the insertion of the mouth parts.
Legs II, III, and IV separated by about the same distance as that from leg I
to the mouth parts.

Basis capituli long with a knob on each side and with two horns on the
posterior corners. Mouth parts very long and slender; mounted in balsam,
articulation between palpal articles 1 and 2 not visible.

Hypostome long and narrow, denticles 2/2; principal denticles in the lateral
files relatively long and sharp, except at the base where they are rounded and

Fig. 48. Ornithodoros viguerasi Cooley and Kohls. A, Capitulum of adult, ventral
view. B, Intercoxal area and coxae of nymph. C. Capitulum of larva, ventral view.
D, Leg I of nymph. E, Leg IV of nymph.

Cooley & Kohls: Argasidae of N. America, etc. 109

blunt, those of the median files small and short. Posthypostomal hairs very
small and short. Length about 0.27.

It is notable that while the large, fed larva measures 2.64 in length, the
adult measures only 3.12.

HOST AND DISTRIBUTION

This species is known only from a bat cave, Cueva Somorrostro, Jamaica,
Cuba, inhabited by Phyllonycteris poeyi Gundlach. One adult and several
nymphs were collected from the walls of the cave; three nymphs and four
larvae were from a bat. The collections were made in September, October,
and November, 1940.

Ornithodoros azteci Matheson, 1935

Plate 12, Figs. 49 and 50

1935. Ornithodorus azteci Matheson, original description, pp. 349-351, with figures.
1941. Ornithodoros anduzei Matheson, original description p. 3.

Sexes similar; adults and nymphs similar.

ADULT

Body. — Pyriform, broadly rounded behind and pointed in front, narrow-
ing gradually from behind the third pair of legs. Size 4.5 x 3.0.

Mammillae. — Large and distinct in the marginal areas, small and not well
formed in the median area. Individual large mammillae subhemispherical,
with the surfaces of the sides irregular and with a few faint, radial striae;
usually with a crater and a short hair on top. On the venter mammillae faint
or absent except in the peripheral areas; short hairs more abundant in the
posterior area.

Discs. — Round or oval, faint, superficial, moderate in size, and evident
only as modifications of the surface of the integument.

Legs. — Moderate in length and size; surface faintly micromammillated or
granular, not shining. Hairs numerous, short except on the terminal articles
where they are longer on the ventral side. Subapical dorsal protuberance mod-
erate on tarsus IV, present but smaller on III. Dorsal humps absent on all
legs. Length of tarsus I, 0.60; metatarsus, 0.54. Length of tarsus IV, 0.78;
metatarsus, 0.66.

Coxae. — Small, I and II separated, all others contiguous. Surfaces micro-
mammillated or granular.

Hood. — Absent or negligible.

Cheeks. — Large, reniform with the convex side attached and with the
anterior end free.

Camerostome. — Well formed and together with the cheeks and the over-
hanging anterior projection of the dorsal body wall forming protection for the
mouth parts.

Capitulum. — Basis capituli about as wide as long; surface irregular, micro-
mammillated or granulated. Palpi long, with article 1 very long, and with a

110

American Midland Naturalist Monograph No. 1

flange projecting over the base of the hypostome. Stalks of the chelicerae very
long and slender.

Hypostome. — Long, attenuated, pointed and with the very fine denticles
limited to near the apex; denticles not in definite files; viewed in lateral profile,
bent, with convex side ventrad.

Folds. — Coxal and supracoxal folds present.

Grooves. — Preanal, transverse postanal and median postanal grooves pres-
ent; dorso-ventral groove absent.

Sexual opening. — Placed between coxae I.

Eyes. — Absent.

Anus. — Large, in an elliptical pattern.

Matheson (1935) describes the nymph and larva as follows:

P Si

Fig. 49. Ornithodoros azteci Matheson. A, Hypostome of adult. B, Leg I of adult.
C, Leg IV of adult. D, Engorged larva, ventral view. E, Hypostome and palpi of
larva, ventral view. F, Leg I of larva. G, Leg III of larva.

Cooley & Kohls: Argasidae of N. America, etc.

Ill

Nymph

From material before me there appears to be only one nymphal stage. Body elongate,
sides subparallel, and strongly constricted in front. Length (unfed) 2.88 mm.; extreme
width 1.6 mm. Coxal and supra-coxal folds well developed; median and transverse
post-anal grooves distinct; preanal groove present but not so well marked as in adult;
integument has lost all indications of parallel transverse striations (so prominent in
larvae) except at anterior end and now appears distinctly mammillated with minute
spines between mammillae. Capitulum located in a deep camerostome as in adult ;
hypostome appears as a very thin, slightly chitinized membrane with only minute
denticles arranged in transverse rows on apical portion (fig. 4, c) ; chelicerae elongate.

Fig. 50. Distribution of Ornithodoros azleci Matheson.

112 American Midland Naturalist Monograph No. 1

sharply pointed shafts terminating in pointed digits (Fig. 2, 3) ; palps as long as
hypostome; joint 1 about twice as long as 3 ; 2 shorter than 1; 4 short, pointed, with
several apical spines.

Recently hatched larva narrowly elongate; length 0.88 mm. as measured from apex
of capitulum to posterior margin of body; capitulum measures 0.44 mm. Body bears
rather numerous stout hairs with minutely spinous margins; integument shows, under
high magnification (x400), very fine parallel lines running transversely, forming a
more or less definite pattern; striate condition appears much more distinctly in mature
larva and resembles that of O. megnini. Hypostome heavily toothed (Fig. 4, b) ;
dentition 2|2; apex with several rows of minute denticles; chelicerae as long as hypo-
stome terminating in a two-toothed article; palps as long as the hypostome; joints 1,
2 and 3 of about equal length ; joint 4 is very short and bears numerous short, pointed
spines; scattered spines present on joints 2 and 3. Fully gorged larva measures 1.68
mm. in length.

To the above it should be added that in the fed larva the capitulum is
plainly visible from above and the coxae are remote from the capitulum.
Hypostome on a conical base which is about as long as the hypostome itself.
Denticles of the lateral files much larger than the medians and both files
extend the entire length of the hypostome; basal denticles crowded and
deformed.

HOSTS AND DISTRIBUTION

From Matheson (1935) we quote as follows:

Bats, Hemiderma perspicillatum aztecum (Saussure) ; Summit, Canal Zone, Nov. 12,
1930; this bat was found to be heavily infested with larvae and the nymphs and adults
were present in the cracks and crevices of the culvert where the bats roosted in masses.
Also taken on this same bat in a cave at Taboga Island, Panama Bay and on the
vampire bat, Desmodus rotundus murinus Wagner. This tick has also been taken in the
cracks and crevices of Chilibrillo Caves and in other caves and culverts in Panama
and the Canal Zone.

Cuba. — Through the kindness of Dr. I. Perez Vigueras, University of
Havana, Havana, we have received other specimens as follows: 17158, 1
adult from walls of a bat cave, Holguin, Santiago de Cuba; 17497, 4 adults,
2 nymphs, 9 larvae, Cueva del Barro, Loma Blanquizar, Guanajay.

Venezuela. — In 1941 Dr. Matheson of Cornell University kindly sent us
specimens of this species taken from bat caves in Venezuela.

Cooley & Kohls: Argasidae of N. America, etc. 113

Ornithodoros kelleyi Cooley and Kohls, 1941
Plate 12. Figs. 51 and 52

1941c. Ornithodoros %e/Zept Cooley and Kohls, original description, pp. 912-914, with
figures.

Sexes similar; nymphs and adults similar.

ADULT5

Body. — Oval, pointed anteriorly and rounded posteriorly; sides more
curved and with the excavations at legs I, II, and III less pronounced than in
talaje. Size of female, 7.35 x 4.0.

Mammillae. — Large, close but not crowded, rounded on top and with the
radial ridges reaching nearly to the top. Equal in sizes in lateral and
median areas and larger at the posterior margin where they are elongated with
the longer axes transverse; small on the venter, and indefinite on the supra-
coxal folds. Hairs absent on both dorsal and ventral surfaces, even on the
hood.

Discs. — Large, conspicuous, in depressed areas which occupy much of the
median area of the dorsum. On the venter, discs in lineal arrangement in the
preanal and transverse postanal grooves and in three depressions caudad of
the transverse postanal groove.

Legs. — Small and moderate in length; surface micromammillated; hairs
small. Tarsus I with a mild subapical dorsal protuberance; absent on all
others; dorsal humps absent Tarsus I notably large (wide when viewed later-
ally). Length of female tarsus I, 0.66; metatarsus, 0.51. Length of tarsus IV,
0.9; metatarsus, 0.66.

Coxae. — Coxae I and II well separated; all others contiguous. Surfaces
micromammillated and with excrescences.

Hood. — Very small and short, placed close to the anterior point of the
body and separated from it only by a depressed line. Not tongue-shaped as in
talaje.

Camerostome. — Negligible and obscured by the large cheeks.

Cheeks. — Large, reniform, with the anterior portion free.

Capitulum. — Basis capituli about as wide as long; surface with irregular,
transverse, deep wrinkles and numerous micromammillae; with a pair of in-
conspicuous fine hairs posterior to the posthypostomal hairs and with a few
fine, short hairs on each lateral margin behind. Palpal article 1 micromammil-
lated. Hairs on the palpi moderate in number and in length.

5 Described from specimens from houses in New York, Minnesota, and Illinois.
The species was originally described from larvae and nymphs from bats in Utah and
Colorado.

114

American Midland Naturalist Monograph No. 1

Fig. 51. Ornithodoros £e//et)i" Cooley and Kohls. A, Lateral view of anterior end of
body. B, Hypostome of female. C, Hypostome and palpi, ventral view of second stage
nymph. D, Leg I of adult. E, Leg IV of adult. F, Capitulum of larva, ventral view.
G, Larva, dorsal view.

Cooley & Kohls: Argasidae of N. America, etc.

115

Hypostome. — Short and small, sides subparallel, a little wider at the denti-
cles than at the middle; apically notched. Denticles 2/2 with about 4 in each
file. Corona with many very fine denticles. Length of female hypostome, 0.21.

Folds. — Coxal and supracoxal folds present, the latter extending forward
to near the hood.

Grooves. — Preanal, transverse postanal, and median postanal grooves pres-
ent, the last terminating at the transverse postanal groove. Dorso-ventral
groove absent.

Sexual opening. — Between the posterior ends of coxae I.

Eyes. — Absent.

Anus. — In an oval frame.

LARVA

Unfed larva sub-oval in shape. Length of body including capitulum, 0.96;
length of capitulum (in ventral view), 0.42. Palpi very long, slender, and in
life closely appressed on the mouth parts.

Hypostome. — Long and slender and with a basal portion without denticles
about one-third as long as the apical portion with denticles; sides converging
to the pointed apex. Principal denticles 2/2, extending the full length; apically
3/3; lateral denticles much larger and sharper. Length about 0.195 (measured
from beginning of the basal teeth to the apex) .

O. kelleyi is related to talaje and concanensis. Nymphs and adults of
kelleyi may be separated from talaje by the absence of a definite notch just
above the hood, visible when viewed in lateral profile. The unfed larvae of
talaje are only about two-thirds as long as those of kelleyi. From concanensis

Fig. 52. Distribution of Ornithodoros fyelleyi Cooley and Kohls.

116 American Midland Naturalist Monograph No. 1

it is distinguished by being proportionately longer and in having the cheeks
smaller, as well as by having the sides of the adult hypostome about parallel.
O. concanensis is known only from bats and bat retreats and kelleyi from bats
and from houses, some of which harbor bats. O. talaje has not been recorded
on bats. The resemblance of this species to O. talaje led to its having been
reported as talaje in houses in New York, Wisconsin, and Minnesota, by
Matheson, 1931, Herrick, 1935, and Riley (1935), respectively. Other records
(unpublished) are from houses in Pennsylvania, Illinois, and Iowa. Collection
data are summarized in table 7.

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118 American Midland Naturalist Monograph No. 1

Genus Antricola Cooley and Kohls, 1942

Dorsal walls flattened and marginated; below the flattened dorsum the
'body convex and deep. Integument semi-translucent and the surface smooth,
shining, and with tubercles. Discs absent on the venter. Mouth parts adapted
for quick feeding and not for clinging to the host; hypostome convex ventral-
ly, concave dorsally and lacking effective denticles; chelicerae large and effec-
tive. Anal ring large. Eyes absent. Eggs small and the small larvae with
bulbous pulvillae in place of claws.

Genotype: Ormthodoros coprophtlus Mcintosh, 1935.

Key to Species of Antricola

1. Margin of body of adults and nymphs wiith finger-like projections, each having

several hairs marginatus (p. 123)

2. Margin of body of adults and nymphs lacking projections coprophilus (p. 1 18)

Antricola coprophilus (Mcintosh), 1°35

Plate 13, Figs. 53, 54, and 55

1935. Orniihodoros coprophilus Mcintosh, original description, pp. 519-522, with
figures.

Sexes dissimilar; adults and nymphs similar.

FEMALE6

Body. — Length about 6.0; greatest width about 3.3. Color light tan. Shape
in dorsal view approching pyriform, tapering to a point in front which is bent
ventrad; with two mild, marginal projections above the spiracles and two
submarginal elevations above legs II, visible from above. Lateral margins
mildly excavated back of leg IV. The marginated dorsal surface, while flat-
tened in general, is irregular and has deep, broad submarginal furrows which
unite in front. Also with deep, short, marginal furrows in the postero-lateral
areas near the excavations.

Tubercles. — Tubercles large, moderate in number, of various shapes and
sizes, those in the median area approaching a hemisphere, those on the margins
fused and directed outward at an angle; absent in the furrows except for a few
in the marginal furrows. Many of the tubercles with one to three hairs on
their curved tops; such hairs longpr in the median area. Tubercles absent on
the lateral walls above the legs but present back of the legs, on the ventral
surface, and in the inter-coxal area where they are smaller and less definite.

Discs. — Discs present in a continuous series in the deep, dorsal furrows;
absent on the venter.

Coxae. — All coxae contiguous, smooth, shining, and with a few hairs.

6 In this species the depleted adults are usually much shrunken and misshapen. Males
are less misshapen than the females. Specimens for describing were selected out of many
for being what we may call "normal.'

Cooley & Kohls: Argasidae of N. America, etc.

119

Legs. — Long and slender and with numerous barbed hairs; surface smooth,
shining. Claws large. All tarsi lacking subapical dorsal protuberances and
dorsal humps. Length of tarsus I, 0.66; metatarsus, 0.78. Length of tarsus IV,
1.02; metatarsus, 1.02.

Hood. — No true hood is present, though there is an anterior projection
of the body wall.

Fig. 53. Antricola coprophilus (Mcintosh). A, Larva, dorsal view. B, Hypostome
and palpi of larva, ventral view. C, Chelicerae of adult. D, Leg I of male. E, Leg IV
of male. F, Leg I of female. G, Leg IV of female. H, Barbed hairs. I, Capitulum of
nymph, lateral view.

120

American Midland Naturalist Monograph No. 1

Camerostome. — Well developed as a depression to receive the capitulum
when not extended.

Cheeks. — Cheeks absent but near their position are five to seven finger-like
projections each bearing one or two barbed hairs.

Capitulum. — Basis capituli tumescent, with the sides rounded over into
the lateral walls which are plainly visible. Surface smooth and shining and

Fig. 54. Aniricola coprophilus (Mcintosh). A, Capitulum of male, ventral aspect.
B, Anterior end of female, showing capitulum, anterior portion of hood, and finger-like
appendages of camerostome, ventral aspect. C, Distal portion of left tarsus IV of male,
lateral aspect. D, Distal portion of right tarsus I of male, ventral aspect. E, Anal valves
of male showing arrangement of hairs. F, Right stigmal plate of male and blunt hairs
surrounding same. From Mcintosh (1935).

Cooley dc Kohls: Argasidae of N. America, etc. 121

with a few scattered, long barbed hairs. Palpi of moderate length; article 1
ciosely appressed on the base of the hypostome; articles 2, 3 and 4, free.

Hypostome. — Short, broad, and rounded apically; convex ventrally and
concave dorsally ( scoop- 1 ike ) ; with very fine teeth on the anterior lateral
margins (better seen in a balsam mount under a microscope).

Chelicerae. — Very large and effective.

Folds. — Coxal and supracoxal folds present.

Grooves. — The transverse postanal groove present, short; all others indefi-
nite or absent.

Spiracles. — Spiracles an oval, globular protuberance, free on all sides
except at the smaller attached end.

Sexual opening. — At the level of the intervals between coxae I and II.

Anus. — Very large, in an elliptical frame.

MALE

Body. — Length about 4.44; greatest width (at the spiracles) about 2.75.
Color light tan. Some specimens are as small as 4.0 x 2.57 (Mcintosh) . Shape
pyriform, tapering in front to a point which is bent ventrad.

Dorsal surface in general flattened and marginated as in the female, and
with mild sub-marginal grooves which unite in front but lacking both the
excavations in the margins at the sides behind the middle and the deep mar-
ginal furrows near them.

Tubercles — As in the females, 'but with the hairs absent or few in number.

Discs. — Essentially as in the female.

Legs. — Legs shorter and larger than in the female; tarsi shorter and with
well-developed ventral spurs (absent in the female). Length of tarsus I, 0.45;
metatarsus, 0.57; length of tarsus IV, 0.54; metatarsus, 0.66.

Coxae. — As in the female.
Hood. — Absent as in the female.

Cheeks. — Cheeks absent; finger-like projections near the position of cheeks
smaller and fewer in number than in the female.

Grooves, folds, spiracles, and anus essentially as in the female.

Sexual opening. — Placed between coxae I (a little anterior to its position
in the female) .

NYMPH

With many specimens of all stages before us the sexes as well as the late
stage nymphs which are to become either male or female are readily separable.
Thus we have in this species nymphs which we may speak of as pro-male or
pro-female. Their potential sex is indicated by their size, shape, relative length
over width, legs, as well as by the absence in the pro-male of the postero-lateral
emarginations and the deep marginal grooves near them.

First and second stage nymphs measure 1.8 x 1.14 and 2.2 x 1.35, respec-

122

American Midland Naturalist Monograph No. 1

tively. The very early stage nymphs have the elongated tubercles lying hori-
zontally or at an angle at the margins and with one or more hairs projecting
from their tops, thus resembling the tubercles or "protuberances" found on
adult mar ginatus .

LARVA

Small, oval. Length (including mouth parts) 0.44, width 0.25. Legs short
and with large, bulbous pulvilli; claws absent; leg 1 the longest and only about
twice as long as the mouth parts. Capitulum terminal, with basis capituli short,
scarcely visible from above. Dorsal plate large, oval. Hypostome long, sides

Fig. 55. Anlricola coprophilus (Mcintosh). Male, ventral view. From Mcintosh

(1935).

Cooley & Kohls: Argasidae of N. America, etc. 123

subparallel, apex pointed. Denticles apically 3/3, then 2/2, those of the mar-
ginal files largest; those of the median files very small and not easily seen even
when mounted in balsam. Length of hypostome, 0.12.

HOSTS

While A. coprophilus is known to be associated with bats, the species has
not been observed reeding on these animals. The junior author when at the
Picacho Mine tunnel in Arizona, having 'both living coprophilus and living
bats, attempted to feed the ticks on bats confined in a bag but was not success-
ful. While the ticks in all stages may be found in abundance on the bat guano,
they are at times numerous on the walls of the retreat. In one instance, a
female was found laying eggs in a crevice near where the bats were hanging.

The type specimens were found in a carload of bat guano in October 1934,
originating at Linares (Nuevo Leon), Mexico, and held in quarantine at
Metamoros, Mexico; also in bat guano, May 29, 1931, at Tucson, Arizona,
by Dr. L. P. Wehrle.

DISTRIBUTION

In addition to the localities noted above, the species has been collected as
follows:

Arizona. — 16083 and 16156, bat guano, September and October, 1939,
mine tunnel, Picacho Mountain, Pinal County, several hundred nymphs and
adults (Philip, 1940); 17863, 17866, locality as above, May 23, 1940, several
hundred nymphs and adults.

Texas.— 17258, bat guano, September 14, 1940, Ney Cave, Hondo, Medi-
na County, several nymphs and adults; 17261, bat cave, September 16, 1940,
Concan, Uvalde County, numerous nymphs and adults.

Mexico. — "Cueva de la Chepa" harboring numerous bats (Leptonycteris
nivalis and Pteronotus davyi julvus), near Tuxtla Gutierrez, Chipas, October,
1940 (Mazzotti, 1940) ; abandoned mine harboring numerous bats, 5 kilo-
meters west of Coquimatlan, Colima, November, 1940 (Mazzotti, 1941).

Antricola marginatus (Banks), 1910

Plate 14, Figs. 56 and 57
1910. Ornilhodorcs marginalus Banks, original description, p. 6, with figures.

FEMALE

Body.— Size 6.62 x 4.70. Banks (1910) gave the length as 5.0 to 8.0.
Color light tan. Shape in dorsal view sub-oval, widest at about the middle,
thence tapering to the anterior point which is 'bent ventrad; broadly rounded
posteriorly. Dorsal surface flattened or even depressed (f^d specimens would
probably show the surface about level with the margin). Margin with a con-
tinuous row of large tubercles which follows the entire periphery, each bearing

124

American Midland Naturalist Monograph No. 1

a tuft of long, barbed hairs. Body wall thin, semi-translucent, smooth and
shining.

Tubercles. — In addition to the large, tufted tubercles which accentuate the
margins are numerous small ones with one to four small hairs on each, scat-
tered over the dorsal surface. They are absent or faint on the lateral walls
above the legs. There is a large one above leg I and another above leg II.
Anterior to the mouth parts on the venter, tubercles small and each with a
single hair.

Discs. — Evidenced only by faint depressions in symmetrical areas which
lack tubercles; absent on the venter.

Legs. — Long and large, with the surfaces smooth and shining and with
numerous long, barbed hairs. All tarsi lack subapical dorsal protuberances and
humps. Claws large. Length of tarsus I, 0.9; metatarsus, 1.02. Length of tarsus
IV, 1.56; metatarsus, 1.20. (Measurement from a late nymphal stage which is
about as large as the adult.)

Coxae. — All coxae contiguous, smooth, shining, and with hairs as on the
legs.

Fig. 56. Antricola marginalus (Banks). A, Tubercles from the margins. B, Leg I
of nymph. C, Leg IV of nymph. D, Barbed hair. E, Capitulum of nymph, lateral view.

Cooley dc Kohls: Argasidae of N. America, etc.

125

Hood, camerostome and cheeks. — Absent.

Capitulum. — Large, a little flattened on the ventral side. Surface smooth
and shining and with numerous long, barbed hairs. Palpi of moderate length;
articles 2, 3 and 4 free.

Hypostome. — Short, broad and rounded apically; convex ventrally and
concave dorsally, thin and scoop-like; with numerous very fine denticles near
the rounded edge.

Fig. 57. Distribution of Anlricola coprophilus (Mcintosh) and A. marginalus
(Banks).

126 American Midland Naturalist Monograph No. 1

Chelicerae. — Very large and long, with very effective digits.
Folds. — Coxal and supracoxal folds present.

Grooves. — Transverse postanal groove short and deep; all others absent or
indefinite.

Spiracle. — A flattened, globular protuberance, free on all sides except on
the attached smaller end.

Sexual opening. — Between coxae I.

Anus. — Large, in an elliptical frame placed well 'back on the venter.

nymph

With the specimens before us it is evident that the nymphs differ from the
adults only in their smaller size, absence of sex openings, and in having the
marginal tubercles longer.

Male unknown.

HOST AND DISTRIBUTION

The type specimens were from a cave in Guanajay Mountains in Cuba;
"also from a West Indian bat probably from Porto Rico." (Banks, 1910.)

Vigueras (1934) stated that Eumops glaucinus (Wagner) is a host and
reported the collection of two females from bats, Nycticeius cubanus (Gund-
lach), captured in Santiago de las Vegas.

It is evident that this species is closely related to coprophilus (Mcintosh)
but the two are readily separated by the characters mentioned in the key to
species. The original collection has been divided, part being in the National
Museum and part in the collections of the Bureau of Animal Industry, U. S.
Department of Agriculture, and neither lot contained a type label. The senior
author has seen all of the specimens. One female from these has been placed
in the collections of the Rocky Mountain Laboratory, Hamilton, Montana.
The combined lot contains 2 females and 18 nymphs of various sizes. Having
available only one sex, we cannot determine at present whether the sexes are
dissimilar in this species. In the relatively few specimens available and in view
of their relatively small size it was not possible to determine whether the even-
tual sex would be indicated in late stage nymphs.

In the lot now in the collections of the Bureau of Animal Industry there
is one small, probably a first stage, nymph which differs in having the tubercles
on the margin smaller in diameter and relatively much longer than in the
larger nymphs. It is possible either that A. marginatus makes this remarkable
change during its development, or that it represents an early stage of an
undescribed species.

The single female in the lot in the National Museum was separated and
labelled as the lectotype of the species.

Cooley 8c Kohls: Argasidae of N. Americ k, etc.

127

Plate I

Argas reflexus (Fabricius)
Argas pcrsicus (Oken)

128 American Midland Naturalist Monograph No. 1

Plate 2

Otobius megmni (Duges). Adult
Otobius megnini (Duges). Nymph

Cooley 8c Kohls: Argasidae of N. America, etc.

129

Plate 3

Otoblus lagophilus Cooley and Kohls. Adult
Otobius lagophilus Cooley and Kohls. Nymph

130

American Midland Naturalist Monograph No. 1

Plate 4

Omilhcdoros savign\)i (Audouin)
Onulhodoros eremicus Cooley and Kohls

Cooley & Kohls: Argasidae of N. America, etc.

131

Plate 5

Ornilhodoros coriaceus Koch

Ornithodoros hermsi Wheeler, Herms, and Meyer

132

American Midland Naturalist Monograph No. 1

Plate 6

Ornilhodoros nicollei Mooser
Ornilhodoros coolevi Mclvor

Cooley & Kohls: Argasidae of N. America, etc.

133

Plate 7

Ornilhodoros yumatensis Cooley and Kohls
Ornithodoros brodyi Matheson

134

American Midland Naturalist Monograph No. 1

Plate 8

Oriiithodoros talaje (Guenn-Meneville)
OrnilhoJoros concanensis Cooley and Kohls

Cooley & Kohls: Argasidae of N. America, etc.

135

Plate 9

OrnilhoJoros slageri Cooley and Kohls
OrnithoJoros dperi Cooley and Kohls

136

American Midland Naturalist Monograph No. 1

Plate 10

Ornithodoros amblus Chamberlin
Ornithodoros rudis Karsch

Cooley & Kohls: Argasidae of N. America, etc.

137

Plate 11

Ornithodoros dunni Matheson
Ornithodoros viguerasi Cooley and Kohls

138 American Midland Naturalist Monograph No. 1

Plate 12

Ornithodoros azleci Matheson
Ornithodoros }(clle\)i Cooley and Kohls

Cooley & Kohls: Argasidae of N. America, etc.

139

Plate 13

Antricola coprophilus (Mcintosh). Female
/'.t.l icAa c~prcphilus (Mcintosh). Male

140 American Midland Naturalist Monograph No. 1

Plate 14

Antricola marginalus (Banks)

Cooley & Kohls: Argasidae of N. America, etc. 141

Classified List of Hosts

Vertebrata

Reptilia

Serpentes

Boidae

Epicrates cenchris Ornithodoros talaje

Crotalidae

"Florida diamond-back rattlesnake" Ornithodoros nicollei

"Rattlesnake" Ornithodoros turicata

Testudinata
Testudinidae

Terrapene ornata Ornithodoros turicata

Terrapene sp. Ornithodoros turicata

Copherus polvphemus Ornithodoros turicata

Copherus agassizii agassizii Ornithodoros turicata

Aves

Struthioniformes
Struthionidae

Slruihio auslralis (South Africa) Argas persicus

Struthio auslralis (South Africa) Otobius megnini

Sphenisciformes
Spheniscidae

Spheniscus humboldti Ornithodoros amblus

Pelecaniformes %
Sulidae

Sula nebouxi Ornithodoros talaje

Anseriformes
Anatidae

Anser domestica (Europe) Argas reflexus

Falconiformes
Cathartidae

Cvmnogyps calif ornianus Argas reflexus

Sagittamdae

Sagittarius serpenlarius (South Africa) Argas persicus

Galliformes
Perdicidae

"Quail" Argas persicus

Phasianidae

Callus domeslicus (Europe) Argas reflexus

Callus domeslicus (Europe) Argas persicus

Callus domeslicus (Europe) Ornithodoros talaje (?)

Numididae

Numida papillosa transvaalensis

(South Africa) Argas persicus

Meleagrididae

Meleagris gallapavo Argas persicus

142

American Midland Naturalist Monograph No. 1

CoLUMBIFORMES

Columbidae

Columba livia (Europe)
Zenaidura macrura

At gas reflexus
Argas persicus

Strigiformes

Strigidae

Speotyto cunicularia

Speotyto cunicularia

Olus asio inyoensis (Nesting hole of)

Olus asio inyoensis (Nesting hole of)

Ornithodoros parpen
Ornithodoros luricata
Argas reflexus
Argas persicus

Hirundinidae

"Cliff swallow's nest in cave.

Fringillidae

Zonotrichia coronaia

Passeriformes

Ornithodoros coriaceus

Argas persicus

Didelphiidae

Didelphis marsupialis etensis

Nocrihonidae

Dirias albiventor minor

Phyllostomidae

Hemiderma perspicillatum aztecum
Hemiderma perspicillalum aztecum
Phyllonycieris poevi

Desmodontidae

Desmodus rolundus murinus

Vespertihonidae

M))olis californicus pallidus
Myotis lucifugus lucifugus
Mvotis lucifugus subsp.
Myotis velifer velifer
Myotis velifer subsp.
Pipistrellus hesperus hesperus
Pipistrellus subflavus
Nycticeius cubanus

Molossidae

Tadanda mexicana
Eumops glaucinus

Mammalia
Marsupialia

Ornithodoros ialaje

Chiroptera

Ornithodoros dunni

Mustelidae
Muslela sp.

Canidae

Canis familiaris
Canis familiaris
Canis sp. (coyote)

Felidae

Felis caius
Felis calus

Ornithodoros azteci
Ornithodoros brodyi
Ornithodoros viguerasi

Ornithodoros azteci

Ornithodoros fyelleyi
Ornithodoros l^elleyi
Ornithodoros Ifelleyi
Ornithodoros yumatensis
Ornithodoros stageri
Ornithodoros fyelleyi
Ornithodoros fyelleyi
Antricola marginatus

Ornithodoros stageri
Antricola marginatus

Carnivora

Ornithodoros parfyeri

Otobius megnini
Ornithodoros ialaje
Otobius megnini

Otobius lagophilus
Ornithodoros talaje

Cooley & Kohls: Argasidae of N. America, etc.

143

"White faced monkey"

Alouattidae

Aloualla palliata palliata

Cebidae

Cebus capucinus capucinus
Cebus capucinus imitator

Saimiridae

Saimiri orstedii orstedii

Hominidae

Homo sapiens

Primates

Ornilhodoros ialaje

Ornithodoros talaje

Ornilhodoros Ialaje

Ornilhodoros Ialaje

Ornilhodoros talaje

Sciundae

Marmota sp.
Citellus sp.
Citellus sp.
Citellus sp.
Citellus beecheyi
Citellus beecheyi fisheri
Citellus columbianus
Citellus richardsoni
Cynomys sp.
Cynomys sp.
Cynomys leucurus
Eutamias sp. (nest of)

Heteromyidae

Dipodomys sp.

Dipodomys sp. (burrows of)

Dipodomys sp. (burrows of)
Cricetidae

Peromyscus sp.

Peromyscus maniculaius

Neoloma floridana baileyi

Neoloma sp.

Neoloma sp. (den of)

Hodomys alleni
Mundae

Rattus nor\>egicus

Ratlus ratlus

Raiius rattus alexandrinus

Argas reflexus

Argas persicus

Otobius megnini

Ornilhodoros

coriaceus

Ornilhodoros

hermsi

Ornilhodoros

nicollei

Ornilhodoros

parfyeri

Ornilhodoros

rudis

Ornilhodoros

slageri

Ornilhodoros

talaje

Ornithodoros

turicala

RoDENTIA

Ornithodoros

parfceri

Ornithodoros

ialaje

Ornilhodoros

turicala

Ornithodoros

parfyeri

Ornilhodoros

talaje

Ornithodoros

turicala

Ornithodoros

parfyeri

Ornithodoros

parlferi

Ornithodoros

parl(eri

Ornithodoros

turicala

Ornithodoros

parfferi

Ornilhodoros

hermsi

Ornithodoros ialaje
Ornithodoros turicala
Ornithodoros parfyeri

Ornithodoros par}(cri
Ornithodoros eremicus
Ornilhodoros talaje
Ornithodoros Ialaje
Ornilhodoros turicaia
Ornithodoros nicollei

Ornithodoros Ialaje
Ornithodoros talaje
Ornithodoros talaje

Lepondae

Lepus sp. (jack rabbits)

Lepus sp. (jack rabbits)

Lepus sp. (jack rabbits)

Lagomorpha

Ornithodoros turicala
Otobius lagophilus
Otobius megnini

144

American Midland Naturalist Monograph No. 1

Lepus californicus

Olobius lagophilus

Lepus californicus

desert

\icola

Otobius lagophilus

Lepus californicus

toe

dla

to alia

Olobius megnini

Lepus totonsendii

Olobius lagophilus

S\)lvilagus sp.

Otobius lagophilus

Sylvilagus sp.

Olobius megnini

Sylvilagus sp.

Ornithodoros parfyeri

Svfoilagus sp.

Ornithodoros turicata

"Rabbit"

Olobius lagophilus

Artiodactyla

Suiidae

Sus scrofa
Cervidae

Odocoileus hemionus subsp.

Odocoileus hemionus subsp.

Odocoileus hemionus columbianus

Odocoileus virginianus

"Deer"
Bovidae

Dos taurus

Dos taurus

Dos laurus

Ovi's aries

Ovis canadensis

Capra hircus

Ornilhodoros turicata

Otobius megnini
Ornithodoros coriaceus
Ornilhodoros coriaceus
Otobius megnini
Otobius megnini

Otobius megnini
Ornithodoros turicala
Ornithodoros coriaceus
Olobius megnini
Olobius megnini
Otobius megnini

Perissodactyla

Equidae

Equus caballus

Equus caballus

Equus caballus (Rumania)

Equus asinus

Mule

Olobius megnini
Ornithodoros turicata
Argas reflexus
Olobius megnini
Otobius megnini

Geographical Distribution of Species

Canada

Argas persicus : British Columbia

Olobius lagophilus : Alberta

Otobius megnini: British Columbia

United States

Argas reflexus California, Montana

Argas persicus : Widespread

Otobius lagophilus : California, Colorado, Idaho, Montana, Ne-
vada, Oregon, Wyoming.

Otobius megnini : Widespread

Ornithodoros amblus : In "seaport" in guano shipment from Peruv-
ian islands.

Ornithodoros concanensis : Arizona, Texas

Ornithodoros cooleyi : Nevada

Ornilhodoros coriaceus : California

Ornithodoros dyeri : Arizona, California

Ornithodoros eremicus : Utah

Cooley & Kohls: Argasidae of N. America, etc. 145

Omithodoros hcrmsi : California, Colorado, Idaho, Nevada, Oregon.

Ornithodoros Ij;elle\)i : Utah, Colorado, Illinois, Iowa, Minnesota,

New York, Pennsylvania, Wisconsin.

Ornithodoros nicollei: Missouri (in Zoo).

Ornithodoros parl(eri : California, Colorado, Idaho, Montana, Ne-
vada, Oregon, Utah, Washington, Wyoming.

Ornithodoros stageri : Arizona, California, Oklahoma, Texas.

Ornithodoros talaje: Arizona, California, Florida, Kansas, Nevada,

Texas.

Ornithodoros luricata: Arizona, California, Colorado, Florida, Kan-
sas, New Mexico, Oklahoma, Texas, Utah.

Ornithodoros yumalensis : Arizona, California, Texas.

Antncola coprophilus : Arizona, Texas.

Mexico

Argas persicus : Widespread.

Otohius megnini : Widespread.

Ornithodoros coriaceus : Widespread.

Ornithodoros dyeri: State of Colima.

Ornithodoros nicollei: States of Guerrero and Colima.

Ornithodoros talaje : Widespread.

Ornithodoros luricata : Widespread.

Antricola coprophilus: States of Nuevo Leon, Chiapas and Colima.

Guatemala
Ornithodoros talaje: Probably widespread.

Panama

Argas persicus : Widespread.

Ornithodoros azleci : Widespread.

Ornithodoros hrodyi : Chilibrillo Caves.

Ornithodoros dunni : Panama City; Summit, Canal Zone.

Ornithodoros rudis : Widespread.

Ornithodoros talaje : Widespread.

Cuba

Argas persicus: Provinces of Havana. Matanzas, and Santa

Clara.

Otohius megnini: Province of Havana.

Ornithodoros azteci : Holguin, Guanajay.

Ornithodoros viguerasi : Cueva Somorrostro (near Jamaica).

Antricola marginalus : Guanajay Mts.; Santiago de las Vegas.

146 American Midland Naturalist Monograph No. 1

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Cooley & Kohls: Argasidae of N. America, etc.

151

INDEX

Synonyms

abbreviations 2

Acarus (Argas) 14

Alectorobius (Ornithodoros) 37, 82

amblus (Ornithodoros) 99

American Q fever 3

americana (Argas) 17, 21

americanus (Ornithodoros) 56

anduzci (Ornithodoros) 109

anterior projection 4

Antricola 1 18

key to species of Antricola 1 18

anus 4

frame of 5

apical ventral spur 4

Argas - 13

key to species of Argas 13

Argas (Ornithodoros) 37, 56, 82

Argasidae 1 1

key to genera of Argasidae 12

article 4

attenuated - 4

azteci (Ornithodoros) 109

barbed hairs 4

basis capituli 4

brevipes (Argas) 13

brodyi (Ornithodoros) 80

buttons 4

camerostome 4

capituli, basis 4

capitulum 4

characters (Fig. 1) 6

cheeks 4

chehcerae 4

chinche (Argas) 17

cleaning 8

clubbed hairs 4

color 4

columbarum (Argas) 14

concanensis (Ornithodoros) 88

cooleyi (Ornithodoros) 52

coprophilus (Antricola) 118

conaceus (Ornithodoros) 42

corona 4

coxae 4

denticles 5

depletion 5

discs 5

dissecting microscope 7

dissection 8

dorsal humps 5

dorsal plate 5

in italics

dorsum 5

dunni (Ornithodoros) 103

dyeri (Ornithodoros) 95

eremicus (Ornithodoros) 40

espagnol (Ixodes) 14

explanation of terms 4

folds 5

frame of the anus 5

granulations 5

grooves 5

hairs 5

barbed hairs 4

clubbed hairs 4

hermsi (Ornithodoros) 46

hood 5

hosts, classified list of 141

humps, dorsal 5

hypostome 5

introduction 1

Ixodes (Argas) 14

kelleyi (Ornithodoros) 113

lagophilus (Otobius) 32

legs 5

lighting 7

magnus (Argas) 14

mammillae 5

marginatus (Argas) 14

marginatus (Acarus) 14

marginatus (Antricola) 123

mauritianus (Argas) 17

measurements 6

medical importance 3

megnini (Otobius) 21

methods 7

micromammillae 6

microscope, dissecting 7

migonei (Ornithodoros) 101

millimeters (see measurements) 6

miniatus (Argas) 17

morbillosus (Ornithodoros) 38

nicollei (Ornithodoros) 50

ornate 7

Ornithodoros 37

key to species of Ornithodoros 37

152

American Midland Naturalist Monograph No. 1

Ornithodoros (Otobius) 21

Otobius 21

key to species of Otobius 21

panduriform 7

paralysis of fowls 3

parkeri (Ornithodoros) 62

parkeri and turicata compared 69

persicus (Argas) 17

pest of cattle 3

pest of poultry 3

plates '27

protuberance, dorsal subapical 7

posing o

posthypostomal hairs - 7

postpalpal hairs 7

preserving fluid 0

projection, anterior 4

radiatus (Argas) 17

rearing "

reflexus (Argas) 14

relapsing fever 3

Rhynchoprion (Argas) 14, 17, 37

Rhynchoprium (Ornithodoros) 21, 37

Rocky Mountain spotted fever 3

rudis (Ornithodoros) 101

sanchezi (Argas) 17

savignyi (Ornithodoros) 38

schinzii (Argas) 38

sex opening 7

similar '

simple hairs 7

spinosum (Rhynchoprium) 21

spiracle /

spirochetes of relapsing fever 3

spirochetosis, avian 3

spur, apical ventral 4

stageri (Ornithodoros) 91

subapical dorsal protuberance 7

sutural line 7

talaje (Ornithodoros) 82

terms, explanation of 4

truncated hairs _ 7

tubercles '

turicata (Ornithodoros) 56

venezuelensis (Ornithodoros) 101

venter '

veterinary importance 3

viguerasi (Ornithodoros) 106

rvheeleri (Ornithodoros) 62

yumatensis (Ornithodoros) 74