The Ticks of California (Acari: Ixodida)

Home , Acari, Amblyomma, Argasidae, Argas persicus, Dermacentor, Dermacentor albipictus

BULLETIN OF THE CALIFORNIA INSECT SURVEY Volume 25

by Deane P. Furman and Edmond C. Loomis THE TICKS OF CALIFORNIA (Acari: Ixodida) BULLETIN OF THE CALIFORNIA INSECT SURVEY VOLUME 25

THE TICKS OF CALIFORNIA (Acari: Ixodida)

BY DEANE P. FURMAN AND EDMOND C. LOOMIS

UNIVERSITY OF CALIFORNIA PRESS BERKELEY LOSANGELES LONDON BULLETIN OF THE CALIFORNIA INSECT SURVEY

Editorial Board: Ted Case, John Chemsak, John Doyen, Henry Hespenheide, T.A. Miller, John Pinto, Rudolph Pipa, Jerry A. Powell, Arthur Shapiro, Robbin Thorp

VOLUME 25 Issue Date: July 1984

UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES

UNIVERSITY OF CALIFORNIA PRESS, LTD. LONWN , ENGLAND

ISBN 0-520-09685-1 LIBRARY OF CONGRESS CATALOG CARD NUMBER 83-9265

01984 BY THE REGENTS OF THE UNIVERSITY OF CALIFORNIA PRINTED IN THE UNITED STATES OF AMERICA

LIBRARY OF CONGRESS CATALOGING IN PUBLICATION DATA Furman, Deane Philip, 1915- The ticks of California (Acari: Ixodida) (Bulletin of the California Insect Survey; v. 25) Bibliography: p. 1. Ticks--California--Classification. 2. Arachnida-- Classification. 3. Arachnida--California-Xlassifica- tion. I. Loomis, Edmond C. 11. Title. 111. Series. QL475.C3C3 vol. 25 595.709794s 83-9265 rQL4581 r595.4 '23 ISBN 0-520-09685-1 (pbk.) Contents

Acknowledgments, viii

Introduction 1

Medical and Veterinary Importance

Biology 2

Methods 2

Preparation and Examination, 2

Simplified Identification, 3

Taxonomic Characters, 3

Key to Families (all stages) and Genera of Adults 4

Family Argasidae 5

Key to Larvae of Argasidae Species, 6

Genus Argas, 7

Key to Argas Adults and Late-Stage Nymphs, 7

Argas brevipes Banks, 8

Argas cooleyi Kohls and Hoogstraal, 9

Argas giganteus Kohls and Clifford, 10

Argas persicus (Oken) , 10

Argas sanchezi Dugbs, 11

Genus Ornithodoros. 11

Key to Ornithodoros Adults and Late-Stage Nymphs, 12

Ornithodoros concanensis Cooley and Kohls, 13

Ornithodoros coriaceus Koch, 13

Ornithodoros dyeri Cooley and Kohls, 15

Ornithodoros hermsi Wheeler, Hems and Meyer, 16

V vi Con tents

Ornithodoros kelleyi Cooley and Kohls, 17

Ornithodoros parkeri Cooley, 18

Ornithodoros rossi Kohls, Sonenshine and Clifford, 18

Ornithodoros sparnus Kohls and Clifford, 19

Ornithodoros stageri Cooley and Kohls, 19

Ornithodoros talaje (Gu6rin+l6neville), 20

Ornithodoros turicata (DugSs) , 21 Ornithodoros yumatensis Cooley and Kohls, 22

Genus Otobius, 22

Key to Otobius Adults and Late-Stage Nymphs, 22

Otobius lagophilus Cooley and Kohls, 22

Otobius megnini (Dug&), 23

Family Ixodidae 25

Key to Nymphs of Ixodidae Genera, 25

Key to Larvae of Ixodidae Genera, 26

Genus Amblyomma, 26

Amblyomma americanum (Linnaeus), 27

Genus Boophilus, 27

Boophilus annulatus (Say) , 27

Genus Dermacentor, 28

Key to Adults of Dermacentor, 28

Key to Nymphs of Dermacentor, 29

Key to Larvae of Dermacentor, 30

Dermacentor albipictus (Padcard) , 30

Dermacentor andersoni Stiles, 32

Dermacentor hunteri Bishopp, 34

Dermacentor occidentalis Marx, 37

Dermacentor parumapertus Newnann, 40

Dermacentor variabilis (Say) , 41

Genus Haemaphysalis, 43 Key to Haemaphysalis, 44 Contents vii

Haemaphysalis chordeilis (Packard) , 45

Haemaphysalis leporispalustris (Packard) , 46

Genus Ixodes , 47

Key to Females of Ixodes, 47

Key to Males of Ixodes, 50

Key to Nymphs of Ixodes, 51

Ixodes angustus Neumann, 53

Ixodes auritulus Neumann, 54

Ixodes brunneus Koch, 55

Ixodes hearlei Gregson, 56

Ixodes holdenriedi Cooley , 58

Ixodes howelli Cooley and Kohls, 58

Ixodes jellisoni Cooley and Kohls, 59

Ixodes kingi Bishopp, 60

Ixodes neotomae Cooley, 61

Ixodes ochotonae Gregson, 62

Ixodes pacificus Cooley and Kohls, 63

Ixodes peromysci Augustson, 64

Ixodes rugosus Bishopp, 65

Ixodes sculptus Neumann, 68

Ixodes signatus Birula, 70

Ixodes soricis Gregson, 71

Ixodes spinipalpis Hadwen and Nuttall, 72

Ixodes texanus Banks, 73

Ixodes uriae White, 75 _I__- --Ixodes woodi Bishopp, 76 Genus Rhipicephalus , 77

Rhipicephalus sanguineus (Latreille) , 77

Literature Cited, 81

Plates, 89 Acknowledgments

We gratefully acknowledge the following recorded at the Rocky Mountain Laboratory. individuals and institutions whose assis- Dr. H. Hoogstraal, U.S. Naval Medical tance made possible the study of institu- Research Unit Number Three, Cairo, Egypt, tional tick collections in their care: C. provided encouragement, advice and speci- M. Clifford and J. E. Keirans, Rocky Moun- mens. Scanning electron micrography was tain Laboratory, National Institute of done by Kim Hoelmer, University of Califor A1 lergy and Infectious Disease, Hamilton, nia, Berkeley, except as indicated below. Montana; D. H. Kavanaugh and V. E. Lee, We are also grateful to Dr. Clifford and California Academy of Sciences, San Francis- Dr. Keirans for providing specimens of 19 co; C. L. Hogue, Los Angeles County Museum; species of ticks used in scanning electron J. A. Powell, California Insect Survey, micrography and for numerous other courte- Department of Entomological Sciences, Uni- sies, including permission to reproduce versity of California, Berkeley; R. 0. illustrations of 6 species of Ixodes from Schuster, University of California, Davis; their publication (Keirans and Clifford, S. I. Frommer, University of California, 1978). Drawings of ticks were carefully Riverside; John Davis, Hastings Reservation, prepared by Carolyn Tibbetts. Among the University of California; R. B. Loomis, many others whose aid we gratefully acknor Department of Biology, California State ledge are A. McGowan and M. Lee for field University, Long Beach; and B. C. Nelson, and laboratory assistance, R. S. Lane, California Department of Health, Berkeley. T. Schwan and M. Buegler for numerous new Dr. Keirans and T. Gary Gautier, Chief collection records, and Ronald Davey for of the ADP Program, National Museum of unengorged nymphs of Boophilus annulatus Natural History, provided a computer print- (Say). out of all collections of California ticks

viii THE TICKS OF CALIFORNIA (Acari: Ixodida)

INTRODUCTION never been isolated from ticks, man, or other animals in California, but 5 sero- Ticks are obligatory, blood-sucking logically distinct strains of spotted-fever- parasites of mammals, birds, and reptiles. group rickettsiae have been isolated from 5 There are about 850 species described for species of ticks in the State (R.S. Lane, the world, most of them in two families: personal communication, 1980). Other tick- the hard ticks, or Ixodidae, and the soft associated pathogens occurring in California ticks, Argasidae. A third family, the include Coxiella burneti, causing Q fever; Nuttalliellidae, is known only from a single Anaplasma marginale, causing anaplasmosis species in Africa. In California 7 genera in cattle; Colorado tick fever virus (CTV); containing 49 species of ticks are known to Powassan fever virus; the bacterium Franci- be established. Two additional species, sella tularensis, causing tularemia; the Amblyomma americanum and Boophilus annula- spirochetes Borrelia hermsi and E. parkeri, e,have been introduced repeatedly into causing tick-borne relapsing fever in the State and are included here in keys and humans; and the protozoan Babesia canis, a text. Other occasional introductions of cause of canine jaundice. B. microti, which exotic ticks are reported by state and has been associated in the eastern United federal agencies, but none of these have States with tick-borne babesiosis of man, a become established in California. malaria-like disease, has been recorded from This publication provides a guide to rodents and a lagomorph in southern Califor- the identification of adult and immature nia (Van Peenen and Duncan, 1968). Lyme instars of ticks occurring in California disease, often resulting in polyarthritis and information on the geographic and sea- in man, has been associated on circumstan- sonal distribution of ticks in the State tial grounds with the bite of Ixodes pacifi- and on their host associations and their -cus in California (Naversen and Gardner, importance to the health and welfare of man 1978). Epizootic bovine abortion (foothill and domestic and wild animals. Host asso- abortion) has been associated circumstan- ciation and other collection records are tially with the feeding of the argasid, summarized in tabular form for all species Ornithodoros coriaceus, in California. Tick encountered frequently. Numbers of speci- paralysis, a potentially fatal disease of mens per collection are not indicated in many animals including man, is associated the tables. Distribution maps include only with the feeding of certain female ixodids precise collection sites. and has been documented in California on a variety of livestock and wildlife (Brunetti, 1965; Loomis and Bushnell, 1968). ME;DICAL AND VETERINARY IMPORTANCE One characteristic of ticks which greatly enhances their importance as vectors and Ticks are among the most important ecto- reservoirs of pathogens is the ability to parasites affecting the health of man and transmit microorganisms transovarially from domestic and wild animals. They serve as one generation of ticks to the next one, as reservoirs and vectors of many organisms in Dermacentor -andersoni and Rocky Mountain pathogenic for humans and other animals, spotted fever, or Ornithodoros hermsi and and cause direct harm by their bites, relapsing fever. Many ticks can also trans- including irritation, anemia, toxemia, mit infections from one developmental instar allergic sensit izat ion, and paralysis. to the next, as in both the above examples. Rickettsia rickettsii, the classical agent Transmissibility is more limited in some of Rocky Mountain spotted fever in man, has cases, however, as in Colorado tick fever,

1 2 BULLETIN OF THE CALIFORNIA INSECT SURVEY where the virus is known to be transmissible unfed state will either seek a favorable from stage to stage in g. andersoni, but microhabitat such as under logs, in crev- transovarial transmission has not been ices, or in loose soil, to await a propi- demonstrated. A third intrinsic charac- tious time for seeking a host, or will teristic of major importance in determining climb onto grasses or shrubs to await the the role of ticks in the dissemination and passage of an animal to which they may maintenance of pathogens is the long life- attach. Thus we use terms such as wood ticks span of these Acari, commonly a year and and field ticks for many ixodids. Charac- frequently two or more years, often exceed- teristically, ixodid ticks remain attached ing that of their hosts. This provides a by their mouthparts to a suitable host for much enhanced potential in time and space several days to weeks, feeding slowly on for the transmission of pathogens to unin- lysed tissue fluids and blood at first, and fected, susceptible hosts. As noted by engorging rapidly on blood before detaching. Balashov (19681, many tick species feed on There is evidence of actual intrastadial small mammals and birds with a high rate of growth (neosomy) in some species, as well population renewal , which decreases the as stretching of soft cuticular membranes potential for a subsidence of infection due as the ixodid tick engorges. to the acquisition of immunity by suscep- Argasid tick biology differs in funda- tible vertebrates. In such situations, mental ways from that of the Ixodidae. Most ticks often become the true reservoirs of argasid ticks are rapid feeders, feeding to infection. repletion in a few minutes. Notable excep- tions are the larvae of species such as the poultry tick, Argas sanchezi, and the paja- BIOLOGY huello, Ornithodoros coriaceus, which feed on a suitable host for 5-10 days. Argasid The basic life cycle of ticks consists ticks do not become as engorged as ixodids, of egg, hexapod larva, octopod nymph, and and may attach and feed more than once dur- adult male or female. Ixodid ticks typi- ing a single life-stage. Adult females of cally have a single nymphal instar, while most argasids typically lay fewer eggs than argasids may have from 3 to 7. A decided ixodids and usually lay them in more than host preference is shown by some species, one batch, with one or more blood meals such as Boophilus annulatus, whereas others intervening. An individual argasid has more appear to be equally content with almost chance of surviving successfully from egg any terrestrial vertebrate that comes their to adult than an ixodid because most arga- way. In many tick species the hosts or sids are lair or nest inhabitants, feeding parts of hosts selected by adults are not rapidly on the host at its central homesite the same as those selected by inmature forms and then remaining at that site, where they (e.g. , Ixodes pacificus) , and the majority can feed repeatedly on the same or a related of species do not molt on the host. Those host. The remarkable ability of some arga- which remain on a host for their entire sids to survive long periods of starvation, parasitic life are known as one-host ticks, up to several years, increases their success e.g., Dermacentor albipictus and Otobius as lair inhabitants as well as the hazard megnini. they pose to hosts as dormant reservoirs of Most ixodid ticks in California are disease agents. three-host ticks: larva, nymph, and adult must each seek a host in order to obtain a blood meal, after which each form drops from METHODS the host. This means that an individual tick may be found in nature in any part of PREPARATION AND EXAMINATION its previous host's range - or, in the case of a larva, of its parental host's range. Essential equipment for definitive tick The chances of an individual tick finding a identification includes a stereoscopic suitable host on three separate occasions microscope capable of 150-200 X magnifica- under such circumstances are not good, but tion. Strong illumination is necessary, the tremendous reproductive capacity of the preferably from a cool light source such as typical ixodid tick compensates for the loss fiber optics. of developing instars. A gravid female American dog tick may lay 6-7 thousand eggs Ticks are best preserved in 70% ethyl in a single batch before dying. In general, alcohol. Nymphs and adults may be examined the various instars of ixodid ticks in the in alcohol under the stereoscopic microscope THE TICKS OF CALIFORNIA 3 or blotted until only moist, with care to body, need only be identified to genus. avoid excessive drying. A small cake of And as the pajahuello, Ornithodoros coria- plastic typewriter cleaner mounted on a =, a soft tick of the family Argasidae, microscope slide facilitates manipulation is the only one of the 6 counnon species of moist specimens; such a cake, available which lacks a dorsal scutum and has mouth- in most office supply stores, maintains its parts not visible from above, it should be flexibility for months . relatively simple to make a tentative iden- Larvae are usually examined as permanent tification. This species typically occurs whole mounts with the compound microscope. in the brushy or tree-shaded environment Preliminary clearing of the larvae may be favored as resting sites by deer and cattle. necessary in preparing mounts; one to three When man also frequents such sites, he is minutes' submersion in steaming lactic acid subject to attack by the resident ticks. is usually sufficient to clear most speci- Host of the ticks in California are mens. After being washed in two or more normally associated with mammalian or avian changes of water, each specimen can be hosts other than man or his domestic pets, mounted on a slide in a drop of Hoyer's and are encountered by man only when he mounting medium and covered with a cover- works with those hosts or invades their slip. Mounted specimens continue to clear ecological niche - e-g., the winter tick, if placed in a warming oven at 40°C for Dermacentor albipictus, on horses and deer, 24-48 hours. After drying, permanent slide or Ornithodoros hermsi, on chipmunks invad- mounts should be ringed with a commercial ing cabins in the high mountains. slide-ringing compound or clear fingernail It is essential in using a key to deter- pol ish. mine if the specimen is a larva, nymph, or adult. Larvae are distinguished by three SIMPLIFIED IDENTIFICATION pairs of legs, in contrast to the four pairs seen in nymphs and adults. Nymphs lack the Identification of the many kinds of genital aperture, which occurs in the adult ticks occurring in California may appear a in the ventral median area between the formidable chore. Fortunately, most of second to fourth pairs of coxae. Nymphs of those encountered by the general public the family Ixodidae have a scutum of the belong to only 6 species, whose habits bring female type, but the basis capituli lacks them into contact with man or his domestic the porose areas seen only in adult females. pets. Of these 6 common ticks 4 are usually TAXONOMIC CHARACTERS acquired by man or his pets through contact- ing infested brush or grass. One of these, Familiarity with tick morphology and Ixodes pacificus, is a small, dark-legged with the terminology used in taxonomic diag- form which need only be identified to genus nosis is necessary for proper tick identifi- for reasonable reliability. The other 3 cation. Specialized terms used in charac- species in this group are the Pacific Coast terizing ticks are defined in the following tick, Dermacentor occidentalis, the American list; see also the labeled illustrations of dog tick, E. variabilis, and the Rocky Moun- key characters on Plates 1-4. tain wood tick, E. andersoni. In most cases the adults of these 3 species are easily Accessory shields: Paired shields outside identified by the characters given in the adanal shields in males of Rhipicepha- key. Identification of E. andersoni is lus. simplified by its restricted range in Cali- Adanrplates: Paired plates lateral to anal fornia, but there are limited areas of over- plate in males of Ixodes. lapping geographic distribution with E. Adanal shields: Paired shields near the anus occidentalis in northeastern California, in males of Rhipicephalus. where intergradation of morphology appears Aeropyles: See Goblets. to occur. Ambulacrum (prew: The terminal segment Its habit of breeding in and around dog of the leg, composed of a basal stalk, kennels or in homes containing dogs makes paired claws, and membranous pulvillus. the brown dog tick, Rhipicephalus sangui- Anal groove: A groove of the cuticle par- e,a common pest in both urban and rural tially circumscribing the anus. situations. Adults of this species, charac- Anal plate: a sclerite surrounding the anus terized by a short capitulum, hexagonal of males of Ixodes. basis capituli, inornate scutum with eyes, Auriculae: Ventrolateral paired extensions and festoons at the posterior end of the of the basis capituli. 4 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Basis capituli: The basal part of the capi- subapical dorsal surface of the tarsi tulum which bears the hypostome, cheli- of the anterior pair of legs. cerae, and palpi. Hypostome: The ventral, anteromedian projec- Camerostome: In Argasidae, the anteroventral tion of the basis capituli, typically cavity in which the capitulum lies. with ventral longitudinal files of Capitulum (gnathosoma) : The anterior part recurved denticles. of a tick including hypostome, cheli- Idiosoma: All of the body exclusive of the cerae, palpi, and basis capituli. capitulum. Cervical grooves: Paired grooves in the Lateral carinae: Anterolateral longitudinal scutum. ridges of the scutum. Cheeks: Paired flaps at the sides of the Lateral grooves: Dorsal grooves at sides of camerostome of some Argasidae. scutum of male ixodids. Chelicerae: Paired mouthparts armed with Marginal grooves: Dorsal grooves at sides laterally directed denticles. of body behind the scutum in female Cornua: Dorsal, posterior projections of ixodids. the posterolateral corners of the basis Median ventral plate: A sclerite, OT plate, capituli of ixodids. on the ventral surface of male Ixodes, Corona: Distal portion of the hypostome, situated between the genital and anal typically armed with minute denticles. apertures. Coxa: Basal segment of leg. Porose areas: Paired, depressed, finely Denticles: The "teeth" on the ventral side pitted areas on the dorsal surface of of the hypostome, usually arranged in the basis capituli of female ixodids. parallel, longitudinal files. Pseudoecutum: The anterior portion of the Dentition: The pattern in which the denti- male scutum homologous with the scutum cles of the hypostome are arranged; in the female. e.g., 3/3 indicates 3 files of denticles Scapulae: The anterior angles, or shoulders, on each side of the midline. of the ecutum of ixodids. Dorsal humps: Elevations of the dorsal sur- Scutum: The dorsal sclerotized plate cover- face of segments of the legs of some ing all or most of the idiosoma of argasids, but exclusive of the dorsal males, and the anterior portion in protuberance . females, nymphs, and larvae of ixodids. Dorsal plate: Sclerotized area of dorsum, Sensilla sagittifonnia: Four paired dermal usually centrally situated, on the larva sensory structures; one pair located of argasids (not to be confused with ventrally behind each pair of coxae and the scutum of ixodids). one pair dorsally on the posterolateral Dorsal protuberances: Dorsal subapical idiosoma of some ixodid larvae. elevations on the legs of same argasids. Spiracles (spiracular plates): Paired scle- Festoons: Uniform, cell-like divisions on rotic areas usually situated posterior the posterior edge of the body of cer- or lateral to coxae XV and containing tain ixodids. goblet cells (aeropyles). Foveae : Small, paired, subcircul ar s truc- Spur: Typically broad, subtriangular projec- tures in the cuticula of the dorsal tions on the coxae and trochanters of surface posterior to the scutum in the the legs or on the palpi, often reduced female and nymph and to the pseudoscutum to mild saliences. in males of certain ixodids. Sutural line: A definite peripheral line of Frame: The sclerotized periphery of the cleavage in Argas spp., dividing the spiracular plate. dorsal and ventral surfaces. Genital aperture: The sexual opening of adult ticks situated on the median line, ventrally between coxae 2-4. KEY TO TICK FAMILIES (ALL STAGES) Genital grooves: Paired, ventral, longitudi- AND GENERA OF ADULTS nal grooves extending from the genital OCCURRING IN CALIFORNIA^ aperture posteriorly, in some cases to the margin of the body. 1 Dorsal scutum absent; capitulum Goblets (Aeropyles): Larger cell-like pores of the spiracular plate. Smaller aeropyles are also often present near frame Genera of adults of ixodids, and nymphs or in dorsal prolongations of spiracular and adults of argasids, are included in plate. this key; for ixodid nymphs, see key under Haller's organ: A sensory structure on the Family Ixodidae. THE TICKS OF CALIFORNIA 5

ventral in nymphs and adults, Palpi short, 1/3 longer than partially or completely con- broad, and with acute trans- cealed from above; sexual verse ridges dorsally and lat- dimorphism of adults slight erally (Fig. 110) ; festoons (Figs. 33-36). Larvae with absent; coxae I shallowly segment 4 of palpi not greatly cleft; spiracular plate oval . reduced, and with palpal ...... Boophilus2 segments all of essentially Palpi short, 314 longer than the same shape. ... .Argasidae .7 broad, without transverse Dorsal scutum present; capitulum ridges dorsally and laterally at anterior margin, not con- (Fig. 353); festoons present; cealed from above; sexual coxae I deeply cleft; spiracu- dimorphism of adults well lar plate comma-shaped .... marked (Figs. 123-128). Larvae ...... Rhipicephalus3 with segment 4 of palpi greatly Margin of body with a sutural reduced and not similar in line separating dorsal and shape to other segments . ventral surfaces (Figs. 14, ...... Ixodidae .2 38); unengorged nymphs and Anal groove distinct, surrounding adults with body margin thin the anus anteriorly (Fig. 8); and acute ...... Argas festoons absent ...... Ixodes Margin of body without a sutural Anal groove distinct or indis- line separating dorsal and tinct but never surrounding ventral surfaces (Fig. 57); the anus anteriorly; festoons margin variably rounded...... 8 often present (Fig. 3)...... -3 Adults with integument granular Eyes absent; second palpal seg- (Fig. 95) and hypostome vesti- ment acutely produced lateral- gial (Fig. 99); nymphs with ly, extending beyond the basis integument spinose and hypo- capituli (Fig. 181). . Haemaphysalis stome well developed (Figs. 98, 100) Otobius Eyes present; second palpal seg- ...... ment not acutely produced lat- Adults and nymphs with integument erally (Fig. 1) ...... -4 alike, mamillated or tubercu- lated, and lacking spines; Capitulum long in relation to hypostome well developed in width; second palpal segment both adult and nymph. .... 3 X longer than wide (Fig. 103) ...... Ornithodoros ...... Amblyomma Capitulum short in relation to width; second palpal segment FAMILY ARGASIDAE CANESTRINI not appreciably longer than wide (Fig. 5) ...... 5 Dorsal scutum absent; little sexual Basis capituli rectangular dor- dimorphism. Adults and nymphs with capitu- sally (Fig. 5); usually ornate; lum partially or completely invisible from male without adanal and acces- above; integument leathery; spiracles usual- sory shields...... Dermacentor ly situated anterior to coxae IV; palpi of all instars free. Of the over 150 described Basis capituli hexagonal dorsally species from the world, 20 are known to be (Fig. 353); inornate; male established in California. As larvae cannot with adanal and accessory be keyed with the same characters as adults shields (Fig. 352)...... 6 and nymphs, a separate key is given here.

1 One species, A. americanum, is occasion- 2 A common New and Old World cattle para- ally encountered on cattle shipped to site, E. annulatus, not currently estab- California from states where the tick is lished in California. established. Only one species reported from North America: E, sanguineus. 6 BULLETIN OF THE CALIFORNIA INSECT SURVEY

KEY TO LARVAE OF SPECIES OF ARGASIDAE IN 7(6) 2 pairs of eyes; tarsus I with 4 CALIFORNIA^ paracapsular setae. ...-0. megnini Eyes absent; tarsus I with 3 Tarsus I with internal, elongate, paracapsular setae 0. lagophilus trumpet-shaped sensillum ex- . . tending posteriorly from cap- 8(6) Dorsum of body with 2 pairs of sule of Haller's organ. Para- eyes; palpal segment IV about sites of birds. . .Argas (Argas) .2 2 X length of any other segment. In deer and cattle bed- Tarsus I lacMng internal trum- ding areas...... pet-shaped sensillum extending .o. (Ornamentum) coriaceus posteriorly from capsule of ..... - Haller' s organ...... 3 Dorsum of body without eyes; palpal segment IV subequal or Dorsal setae: 2 pairs bordering shorter than any other segment . . 9 dorsal plate; typically 15 pairs laterally and 7 pairs on 9(8) Dorsum of body with 12-13 pairs posterior quadrant .A. (A.1 brevipes of setae (11 dorsolateral, 2 central); dorsal plate may be Dorsal setae: 1 pair bordering absent, never pyriform, elon- dorsal plate; typically 22 gated, or pointed if present; pairs laterally; 9 pairs on hypostome bluntly rounded ante- posterior quadrant. .A. (A.1 cooleyi riorly, with denticles 212 only With 3-4 central setae anterior on distal portion (Fig. 58) . . to the midlength of the dorsal ...... O. - (Pavlovskyella) - 10 plate. Parasites of birds. .. Dorsum of body with 14-30 pairs ...... Argas (Persicargas) .4 of setae (10-25 dorsolateral, With less than 3-4 central setae 3-4 central); dorsal plate anterior to the midlength of present, elongated or pyriform; the dorsal plate...... 6 hypostome usually pointed anteriorly, with denticles 313 to Engorged larvae large, about 4 X 515 throughout the length ... 2.5 mm...... A. (p.1 giganteus ...... -0. (Alectorobius) - 12 Engorged larvae small, not over Without either of the above 1 X 0.75 mm ...... 5 combinations of characters Dorsal posterolateral setae only (from woodrats and nests). .. slightly longer than dorsal ...... -0. sparnus anterolateral setae. Tarsus I lO(9) Dorsal plate present (Fig. 59); with 1 pair of lateral setae hypostome short, 0.062-0.076 and 2 pairs of ventral setae. mm (on rodents and their nests) ...... -A. (E.) persicus ...... -0. (E. hermsi Dorsal posterolateral setae about Dorsal plate absent; hypostome 2 X length of dorsal antero- long, over 0.086 mm (on many laterals. Tarsus I with 2 hosts)...... 11 pairs of lateral setae and 3 pairs of ventral setae. ... ll(10) Hypostome 0.086-0.101 mm long ...... -A. (2.1 sanchezi (on rodents and their nests)...... O. (2.)parkeri Venter of body with 5 pairs of setae; dorsum of body with 7-10 Hypostome 0.138-0.150 mm long pairs of setae...... Otobius .7 (on many hosts, including desert tortoises, burrow- Venter of body with 6-9 pairs of inhabiting birds, and rodents). setae; dorsum of body with more 0. (E.) turicata than 10 pairs of setae (except ...... - in 0. sparnus, which has 10 12(9) Capsule of Haller's organ with pairs)...... Ornithodoros .8 reticulations (from bats and bat retreats) ...... 13 Capsule of Haller's organ without Modified from Jones and Clifford, 1972. reticulations ...... 15 THE TICKS OF CALIFORNIA 7

13(12)Venter of body with 8 pairs of Genus Argas Latreille setae plus a posteromedian seta...... -0. (A.1 yumatensis Argas Latreille, 1796, Precis des caracteres gen6riques des insectes disposes Venter of body with 9 pairs of dans un ordre nature1 5:18; Nuttall et setae plus a posteromedian al., 1908, Ticks: monograph of the seta...... 14 Ixodoidea, part I. Argasidae: 4-5 14(13)Hypostome arises from a relative- ( synonymy 1 . ly short, subtriangular median extension of the basis capituli Type of genus: Argas reflexus (Fabricius, ...... O. (A.) dyeri 1794). Hypostome arises from a very Adults and nymphs of all California long, subtriangular median Argas species with body flattened extension of the basis capituli ...... O. (A.1 rossi dorsoventrally and peripheral margin distinctly flattened; sutural line present, 15(12)Hypostome dentition 5/5 distally separating dorsal and ventral surfaces. (from marine birds) ...... 1 Capitulum anteroventral, invisible from ...... o. (A.) capensis above. Integument leathery, minutely wrinkled. Discs present on dorsal and Hypostome dentition fewer than 5/5 distally. 16 ventral surfaces. Eyes absent...... Of the 56 described species of Argas 16(15)Hypostome dentition 4/4 distally in the world (Hoogstraal et al, 19791, 8 (from bats and bat retreats). . are known from the United States, including ...... o. (&.I stageri 5 from California. Hypostome dentition 313 distally. . 17 17(16)Posteromedian seta absent (from KEY TO ARGAS ADULTS AND LATE-STAGE NYMPHS marine birds) ...O. (A.1 denmarki IN CALIFORNIA2 Posteromedian seta present (not 1 Dorsal marginal integument com- from marine birds)...... 18 posed of striae with scattered 18(17)Junction of subtriangular median setae arising from pits or from extension of basis capituli linear interstices between with hypostome abrupt, dorsal ridges (Fig. 24). Postpalpal plate over 0.3 mm long (from setae absent (Fig. 23). ... bats and bat retreats)...... subgenus Argas - 2 .O. (A.1 kelleyi ...... Dorsal marginal integument com- Junction of subtriangular median posed of "cells" (Fig. 38). extension of basis capituli Postpalpal setae present (Pig. with hypostome not abrupt, 37) ....subgenus Persicargas - 3 dorsal plate usually less than 3 0.3 mm long 19 2(1) Legs short; tarsus I less than ...... X longer than broad: tarsus IV 19(18)Subtriangular median extension less than 4 X longer than of basis capituli absent or broad. Hypostome with teeth merely suggested (from rodents) bluntly pointed and short, 2/2 ...... -0. (&.I talaje anteriorly and 313-4/4 poste- Subtriangular median extension riorly (Fig. 23). Recorded from owls, woodpeckers, and of basis capituli present, but kestrel brevipes may be substantially reduced ...... (on bats and birds) ...... Legs moderately long; tarsus I ...... g. (4.1 concanensis more than 3 X longer than

A species close to both capensis and Argas (2.) giganteus is known only from denmarki occurs on marine birds on the the larvae. Farallon Islands of California. 8 BULLETIN OF THE CALIFORNIA INSECT SURVEY

broad; tarsus IV more than 4 X longer than broad. Hypostome with definite corona of 2-3 rows of small hooklets, and shaft with 3-4 rows of narrow pointed denticles and 5-7 rows of small hooklets, 2/2 anteriorly and 313-515 posteriorly (Fig. 28). From Cliff Swallows, California Condor, California Gull...... cooleyi 3(1) About 90 dorsal marginal cells, mostly rectangular to subrectangular, each with single large setiferous pit occupying much of surface area (Fig. 32). Mammillae of dorsal integument elevated above ridges (Fig. 32). Female palpi sparsely setose dorsally. Poultry parasite; common in Old World; in the United States rare west of the Mississippi River. . persicus Over 150 dorsal marginal cells, comparatively small, irregular, with 1-2 setiferous pits in MAP 1. Distribution in California of: Argas about 2 of the cells occupying brevipes Banks, half-circles; A. cooleyi 5 no more than half of surface Kohls and Hoogstraal, solid circles; A. area (Fig. 38). Mammillae and giganteus Kohls and Clifford, rectangles. ridges of dorsal integument of equal elevation. Female palpi densely setose dorsally. Com- Nymph. Similar to adult; large nymphs mon poultry parasite in Cali- may equal females in size and have depres- fornia, also on wild birds such sion in place of genital aperture. as quail, Mourning Dove . . sanchezi Larva. Two pairs of setae around dorsal plate; 15 pairs laterally; 7 pairs on posterior quadrant. Tarsus I with internal, Argas (Argas) brevipes Banks elongate, trumpet-shaped sensillum extending (Figs. 21-24; Map 1) posteriorly from capsule of Haller's organ.

Argas (k.) brevipes Banks, 1908, U.S. Dept. The short-legged North American bird Agric., Bur. Entomol. Tech. Ser. 15: argasid A. brevipes is known from Arizona, 15-16; Kohls, Hoogstraal, and Clifford, California, Baja California, and Sonora, 1961, Ann. Entomol. SOC. her. 54: Mexico, where it lives in close association 869-877 (all instars). with birds that build nests in hollow trees or in holes made in trees or Saguaro cactus. DIAGNOSIS. Adult. Pattern of discs Recorded hosts include Screech and Pygmy and ridges of dorsal integument as in Argas Owls, Mearns' Gilded Flicker, Sparrow Hawk, cooleyi. Closely resembles A. cooleyi, but Violet-Green Swallow, and the Cactus Wren. smaller: females 3.7-5.6 mm long by 2.5-3.6 The life cycle from egg to ovipositing mm wide, compared to females of A. cooleyi, female takes 42-82 days (avg. 561, and 4.3-8.2 m long by 3.1-5.7 mm wide. Hypo- adults may live for one year at room tem- stome (Fig. 23) with only a slight sugges- peratures without feeding (Clifford and tion of coronal hooklets, 3-4 rows of blunt- Kohls, 1963). ly pointed denticles in 212 arrangement, and several more proximal bluntly rounded CALIFORNIA RECORDS. The few California denticles. All legs short. Tarsus I about collections of this tick have been from the 2.8 X longer than broad; tarsus IV about central part of the state. ALAMEDA CO.: 3.5 X longer than broad. Berkeley (Skyline Blvd. and Redwood Rd. 1, THE TICKS OF CALIFORNIA 9 many larvae from Sparrow Hawk, Falco Wyoming, where its chief host is the Cliff sparverius, 111-30-1960 (P. Covel). NAPA Swallow (Petrochelidon pyrrhonota). Single CO.: Monticello Dam, 6 nymphs from owl, collections have been made from the Barred VI-4-1970 (E.E. Grissell). YOLO CO. : Owl (Strix varia helveola) in Texas and from Winters, 2 collections on XI-30-1965, 1 near or in the nests of a California Condor, from the California Woodpecker, Melanerpes Barn Owl, gull, mud dauber, and sparrow in formicivora, and 1 from the Plain Titmouse, California. Most collections have been made -Parus inornatus. from swallow nests from May through October, although records include March and December. According to Kohls and Hoogstraal (19601, Argas (Argas) cooleyi Kohls and Hoogstraal -A. cooleyi should be found in permanently (Figs. 25-28, Map 1) established colonies of Cliff Swallows primarily located on cliff sides as well as Argas (A,) cooleyi Kohls and Hoogstraal, from man-made structures over water sources. 1960, Ann. Entomol. SOC. her. 53:625- Since Cliff Swallows breed during spring 631, Figs. 1-15 (all instars). and early summer the nests remain vacant thereafter, and ticks may retreat into DIAGNOSIS. Adult. Body bluntly rounded earth, rock, wood, or concrete crevices, anteriorly and posteriorly (Fig. 25). thereby making them difficult to find. Lateral marginal integument (Fig. 28) simi- Johnson and Casals (1972) isolated the lar to that of A. brevipes, with distinct Mono Lake virus (Kemerovo group) from a pool suture encircling entire body and striated of adult 4. cooleyi collected from nests of margin bearing scattered setae. Dorsal -Larus californicus on an island in Mono integument with mammillae irregularly pat- Lake, California. terned; ridges smooth, forming a convergent pattern among discs anteriorly; discs large CALIFORNIA RECORDS. COLUSA CO. : 18 mi. and arranged in radiating rows. Hood ab- W Williams, 1 0 from Cliff Swallow nest sent. Basis capituli (Fig. 26) broadly under bridge over irrigation ditch, VI-5- rectangular centrally, 2.5 X wider than 1956; 17 mi. W Williams, 12 nymphs, 1 0 from long. Hypostome (Pig. 26) short and stout, Cliff Swallow nest on Cache Creek bridge on about 2 X as long as wide, apex bluntly Hwy 20, VI-10-1958 (both H.N. Johnson); Salt rounded, with corona of 2-3 rows of small Canyon, males and females from mud dauber hooklets; sharp-pointed denticles, 212 nest (Sphecidae), XI-7-1963. GLENN CO.: 4 apically, smaller and from 313 to 515 basal- mi. W Willows, 1 female, XII-5-1965. INYO ly. Genital aperture a transverse slit at CO.: 10 mi. S Independence, 1 nymph from level of bases of coxae I. Anus elliptical nesting hole of Otus asio, XI-20-1940 (K.E. at level of apex of trochanter IV. Legs Stager). KERN CO.: Coffee Canyon, 1 nymph moderately long and stout, with tarsi 11-IV from Passer domesticus nest, V-30-1946 (D.P. each with a slightly elevated subapical Furman, H.E. McClure). MODOC CO.: Newell, protuberance. Tarsus I (Fig. 27) about 3.3 from swallow nests, larvae on V-17-1959 and X longer than wide; tarsus IV about 4 X adults and larvae on V-26-1961. MONO CO.: longer than wide. Mono Lake, 10 adults from nest of Larus Nymph. Similar to adult, except that californicus, V-18-1966 (H.N. Johnson). basis capituli is more elongate and hypo- RIVERSIDE CO.: San Jacinto (Soboba Indian stome is slightly narrower and more elon- Reservation), adults and larvae from Tyto gate. -alba nest site, X-15-1961. SAN DIEGO CO.: Larva. One pair setae around dorsal Warner Hot Springs, 35 males, 31 females, plate; 22 pairs laterally; 9 pairs on pos- 27 nymphs, VIII-?-1954. (F.N. Gallup). terior quadrant. Tarsus I with internal, SISKIYOU CO.: Tule Lake Wildlife Refuge, 2 elongate, trumpet-shaped sensillum extending nymphs from Petrochelidon pyrrhonota nest, posteriorly from Haller's organ, as in A. IV-?-1963 (E. Hansen). VENTURA CO.: Santa brevipes. Paula Canyon, 7 males, 9 females, 13 nymphs near nest of condor (Gymnogyps californi- A. cooleyi, the Western North American -anus) in cave, IX-18-1939 and V-24-1940 bird-argasid, is recorded from California, (T.F. Kelley), and 58 adults near condor Colorado, Montana, Nevada, Oregon, South nest, 111-25-1946 (C. Koford). Dakota, Texas , Utah, W-ashington, and 10 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Argas (Persicargas) giganteus Kohls and Clifford (UP 1)

Argas (E.) giganteus Kohls and Clifford, 1968, Ann. Entomol. SOC. Amer. 61:1113- 1116 (larva).

DIAGNOSIS. Adult and Nymph unknown. Larva. Engorged larvae very large, up to 4 mm long by 2.57 11110. wide. Dorsal plate rounded. Dorsum with 14 dorsolateral setae and 9-10 central pairs. Venter with 7 pairs setae. Distance betwen 2nd pair of posthp postomal setae about 2 X that between first pair. Eypostome blunt; dentition 414 and 313 in distal third; 212 vroximallv. Engorged larvae of ?A. giganteus are about twice as large as those of other sicargas species.

This tick was described from larvae collected from several species of wild birds in Arizona, Utah, New Mexico, California, and Sonora, Mexico.

CALIFORNIA RECORDS. KERN CO.: 2.5 mi. MAP 2. Distribution in California of: Argas E Onyx, 75 larvae from the Brown Towhee, persicus (Oken), half-circles; A. sanchezi Pipilo fuscus, XII-19-1948. MONO CO.: Dug&, solid circles. White Mountains, 13,100? elev., 2 larvae from the Rock Wren, Salpinctes obsoletus, VI-30-197 4. row of about 90 large rectangular or subrectangular cells bordering body; each cell Argas (Persicargus) persicus (Oken) with large, setiferous pit (Fig. 32). Hood (Figs. 29-32; Map 2) absent. Basis capituli (Fig. 31) broadly rectangular; 1 pair postpalpal setae Rhynchoprion persicum Oken, 1818, Isis: present; 8 (rarely 10) pairs posterolateral 1567-1570 pl. 19, figs. 1-4. setae. Hypostome (Pig. 31) with broad base; Argas persicus, Fischer de Waldheim, 1823, lateral margins parallel and apex bluntly Mem. Soc. Imp. Nat. Moscou 6:269-283, rounded; sharply pointed denticles 212 in pl. 23, figs. 1-11; Cooley and Kohls, files of 3-4 on anterior half. Genital 1944, Amer. Midland Naturalist, monogr. aperture located as transverse slit between 1: 17-20 (synonymy, redescription, coxae I. Anus elliptical, situated medially figs.); Kohls et al., 1970, Ann. and posterior to coxae IV. Legs long and Entomol. Soc. her. 63: 591-595 (all narrow; tarsi moderately stout, with no instars). dorsal subapical humps. Nymph. Similar in late instar to DIAGNOSIS. Adult. Body usually oval, adults, but with fewer setae and lacking narrowly rounded anteriorly and broadly genital aperture. rounded posteriorly (Figs. 29 and 30). Larva. Body length unengorged less than Mammillae posterior of midlength regular 1 mm. Dorsal plate rounded, subquadrangu- and elevated; anterior of midlength fewer, lar, slightly longer than broad. Dorsum smaller, and less elevated. Ridges with 26-29 pairs setae; 1 or 2 pairs border- posteriorly narrow, short, irregular but ing dorsal plate; dorsal posterolateral numerous; centrally becoming longer and more setae short, subequal to, or slightly longer conspicuous ; anteriorly more numerous, than, anterolaterals. Capitulum with 2 straight or curving and in irregular rows. pairs posthypostomal setae; 2nd pair about Discs large and conspicuous in radiating 2 X as widely spaced as first pair. Tarsus rows, with those anterior slightly smaller I with 1 pair lateral setae and 2 pairs than those posterior. Dorsal peripheral ventrals. THE TICKS OF CALIFORNIA 11

A. persicus , the Persian poultry Argas , Nevada, New Mexico, Texas, and Utah, and in is more common in the eastern United States, Mexico it has been recorded in Baja with collections listed from Georgia, Mary- California, Durango, and Guanajuato. Hosts land, and Pennsylvania. Only one collection are chiefly chickens (chicken houses and is known for California - NEVADA CO.: Grass roosts) and turkeys (turkey houses), but a Valley, 1 male, 2 females, 12 nymphs from variety of wild birds have been found chicken house, VI-21-1951 (E.C. Loomis). infested. It has been incriminated in the Repeated attempts in 1966 and 1977 to col- natural transmission of avian spirochetosis lect this species from the Grass Valley area (Borrelia anserina) in California (da Massa were negative, although specimens of A. and Adler, 1979). sanchezi were taken from one of the few remaining backyard chicken coops found in In California 4. sanchezi has been col- this area. All A. persicus collections in lected during every month, but mostly from the eastern states are from chickens or May through September, the months of rapid chicken houses, from June through August. tick multiplication. The poultry tick is Routine inspections of such habitats would usually a problem only to small farm flocks undoubtedly show the presence of this of poultry where harborage for the ticks is species in other months. provided by loosely constructed wooden roosts and housing or under the bark of trees used as roosts (see Table 1). Eggs Argas (Persicargas) sanchezi Duges are laid in cracks or crevices, and in warm (Figs. 33-38; Map 2) weather hatch in about 2 weeks. Larvae attach to poultry and suck blood for about Argas sanchezi Dugss, 1887, La Naturaleza 7 days, then drop off at night and hide in (Mexico), ser. 2, 1:18-21, pl. 3, fig. available crevices. The 2 nymphal instars 2; Kohls et al., 1970, Ann. Entomol. and the adults attach and feed only a few SOC. Amer. 63:598-602, figs. 32-48 (all minutes at a time, usually at night, and instars). spend the rest of the time in hiding near the roosts or nests of their hosts. There DIAGNOSIS. Adult. Body as in A. may be 2-3 generations a year under favor- persicus, with similar pattern and formation able conditions, but developmental time is of mammillae, ridges, and discs (Figs. retarded by cold weather and lack of hosts. 33-36). Peripheral cells in a row of about Infestations may survive in empty poultry 150 or more, small, irregular outlines and housing for up to 2 years. each with 1-2 small setiferous pits occupy- The majority of collection records are ing no more than half of surface area (Fig. in drier areas of the state, particularly 38). Basis capituli, hypostome (Fig. 371, around the central valley from Shasta County genital aperture, anus, tarsi, and short, on the north to Kern County on the south, stout legs as in A. persicus. although A. sanchezi occurs also in the dry Nymph. Similar to adult. coastal and inland areas of southern Cali- Larva. Similar to A. persicus, but fornia. tarsus I with 2 pairs lateral setae and 3 pairs ventral setae. Differs from all other American species of the subgenus Persicargas Genus Ornithodoros Koch in having posterolateral body setae about 2 X as long as the anterolaterals. Ornithodoros Koch, 1844, Arch. Naturgesch. (Berlin) 10:219; Nuttall et el., 1908, Prior to the extensive study by Kohls Ticks: a monograph of the Ixodoidea et al. (1970) of representative specimens 1:39-41 (redescribed; synonymy). of the subgenus Persicargas from many countries, identification of species in this Type of genus : Ornithodoros savignyi subgenus was most difficult. These workers (Audouin, 1826). redescribed 4. (E. radiatus Railliet , E- chezi, and A. (E.) miniatus Koch and con- Adult and nymph similar, with peripheral sidered these to be valid species and not margins thick, not clearly defined (except synonyms of the Old World fowl tick, A. (E.) ;no. dyeri), and similar to rest of integu- persicus (Oken), as formerly believed. ment; sutural line separating dorsal and A. sanchezi is the common poultry tick ventral surfaces absent; body of unfed in California, often called the "blue bug" specimens flattened, engorged forms usually by farmers. It is also known in Arizona, very convex. Capitulum ventral and sub- 12 BULLETIN OF THE CALIFORNIA INSECT SURVEY

TABLE 1. California Records of Argas sanchezi Dugb

Host or Source Tick Collections

Adults Nymphs Larvae Chicken 10 5 2 Chicken house 14 8 4 Turkey house 1 1 - Aphelocoma californica - - 2 Passer domesticus, nest 1 1 - -Homo sapiens 1 1 - Bark of tree 1 - - Unrecorded 7 3 5 Shed 1 - -

terminal, with anterior parts often visible Legs unusually long (Fig. 86); dorsally. Integument with discs and mammil- tarsus I of female over 1 mm lae. Eyes present or absent. long, of male over 0.8 mm long ...... yumatensis Legs not unusually long; tarsus KEY TO ORNITHODOROS ADULTS AN4 LATE-STAGE I of nymphs and adults less NYMPHS IN CALIFORNIA than 0.7 mm long...... 6 2 pairs of eyes along antero- Body slightly wider posteriorly lateral margins (Fig. 44) .coriaceus than anteriorly; tarsus I without subapical dorsal pro- Without eyes along anterolateral 77) margins ...... 2 tuberance (Fig...... stageri Body width subequal posteriorly Cheeks present (Fig. 42) (except and anteriorly; tarsus with in nymphs and males of 0. I small subapical dorsal protu- stageri); dorsal humps on legs berance (Fig. 40) .7 absent (Fig. 87)...... 3 ...... Hypostome appreciably wider Cheeks absent (Fig. 67); dorsal distal to midlength (Fig. 41); humps on legs present or absent . 10 palpi hypertrichous, with From bats and bat retreats...... 4 accessory ventral setae on From birds or non-chiropteran segments 1-3...... concanensis mammals, from their nests, Hypostome with sides subparallel burrows, or ground surface) ... .8 and not appreciably wider distal to mid-length; palpi 2 pairs of dorsal parallel longitudinal submarginal ridges not hypertrichous and without (Fig. 47); body narrowly accessory ventral setae on elongate (Fig. 47) (about 2.3X segments 1-3...... kelleyi longer than wide) ...... dyeri From marine birds and their nests Without 2 pairs of dorsal longi- ...... -0. sp. near denmarki tudinal ridges; body not nar- Associated with rodents and ter- rowly elongate (not more than restrial birds...... 9 2X longer than wide). .5 ...... Hypostome appreciably wider distal to midlength (Fig. 41). .. Modified from Cooley and Kohls, 1944b...... concanensis THE TICKS OF CALIFORNIA 13

I- Hypostome not appreciably wider J distal to midlength .....talaje lO(2) Dorsal humps on tarsi I present (Fig. 68) ...... 11 i Dorsal humps on tarsi I absent. ..12 I ll(l0)Mammillae large, relatively few, and not crowded (Fig. 82) . turicata I Mammillae small, numerous, and crowded (Fig. 69) ...... parkeri i 12( 10)Corona of hypostome with 6-8 fine denticles; dorsoventral groove absent; a smaller tick, with females typically less than 4 mm long. From Neotoma and Peromyscus ...... sparnus ! Corona of hypostome with numerous fine denticles (Fig. 53); dorsoventral groove present but faint (Fig. 52); females about 5 mm long. From Eutamias . . hermsi

Ornithodoros ( Alectorobius) concanensis Cooley and Kohls MAP 3. Distribution in California of (Figs. 39-42; Map 4) Ornithodoros coriaceus Koch, solid circles.

Ornithodoros concanensis Cooley and Kohls, 1941, U.S. Pub. Health Serv. Rept. central pairs. Venter with posteromedian 56:910, figs. 1 and 2 (adult, nymph); seta (= postanal) plus 8 pairs setae; 2 Kohls, Sonenshine, and Clifford, 1965, pairs posthypostomal setae. Hypostome aris- Ann. Entomol. SOC. her. 58:340, figs. ing from small subtriangular extension of 16-18 (larva). basis capituli and tapering to point; denti- Ornithodoros aquilae Cooley, 1944b, J. tion 313 in distal half and 212 to base. Parasitol. 30:287. 0. concanensis closely resembles 0. DIAGNOSIS. Adult. Body (Fig. 39) sub kelleyi Cooley and Kohls and 0. talaje oval; sides nearly parallel; anterior margin (GuBrin-MBneville), from which it is separ- bluntly pointed and posterior rounded. able as indicated in the keys. It is known Mammillae large, close but not crowded; from bat retreats in Arizona and Texas, but usually with 1-2 circular or crescentic pits Kohls, Sonenshine, and Clifford (1965) on top; some pits with minute seta. Discs recorded it from a variety of avian hosts depressed on dorsum with elevated edges, on from Colorado, Wyoming, and Montana. venter lineally arranged in preanal and median postanal grooves. Hood short, ele- CALIFORNIA RECORDS. MODOC COUNTY: vated apically. Cheeks (Fig. 42) oval; Newell: 3 males and 6 nymphs from swallow anterior ends wider and free. Hypostome nest, V-17-1959; 1 female, V-26-1961 (both appreciably wider beyond middle, moderately J. Schuh). SAN LUIS OBISPO COUNTY: Cholame, large, notched apically, and with denticles 26 larvae from Tyto alba pratincola, IV-14- 212, with about 4 per file but limited to 1946 (M.S. Ray). distal third. Palpi hypertrichous (Fig. 41) with accessory ventral setae on segments 1-3. Eyes absent. Tarsus of leg I (Fig. 40) with mild subapical protuberance; dorsal Ornithodoros (Ornamentum) coriaceus Koch humps absent on all tarsi. Pajahuello Tick Nymph. Similar to adult. (Figs. 43-46; Map 3) Larva. Dorsal plate large, pyriform. 12-14 pairs dorsolateral setae; 4-5 dorsal Ornithodorus coriaceus Koch, 1844, Arch. 14 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Naturgesch. 10:217-219; Koch, 1847, Niirnberg 4, 136 pp. (redescription with figures); Cooley and Kohls, 1944, her. Midland Naturalist, monogr. 1:42-45. Argas coriaceus; Berlese, 1888, Boll. della SOC. Entomol. Italiana 20:193.

DIAGNOSIS. Adult. Body pointed anteriorly, with sides nearly parallel (Fig. 43). Mammillae numerous, irregular in shape and variable in size. Discs large, depressed, and less definite or absent on ventral surface. Hood separated from anterior dorsal projection of body wall. Cheeks absent (Fig. 45). Basis capituli broader than long, surface irregular with coarse transverse wrinkles. Hypostome with sides nearly parallel, denticles 2/2 with 4 large equal- sized ones in each file. Anterior pair eyes (Pig. 44) larger than posterior pair. Anus elliptical. Legs moderate in size, with femur, tibia, and metatarsus flared distally (Fig. 43). Subapical dorsal protuberances evident on all tarsi, with dorsal humps on tarsi I (Fig. 46) to 111 and one at base of tarsus IV. Nymph. Similar to adult. MAP 4. Distribution in California of: Larva. Detailed description given by Ornithodoros concanensis Cooley and Kohls, Kohls, Sonenshine, and Clifford (1965). 0. triangles; 0. hermsi Wheeler, Herms, and coriaceus larva differs from all other Meyer, solid circles; 0- sparnus Kohls and argasid larvae in California except that of Clifford, half-circles. Otobius megnini (Dug&) in possessing 2 pairs of eyes on dorsum, and differs from megnini in having segment 4 of palpi about range in coastal California, the tick was 2X as long as any other segment and venter designated "pajahuello" by Banks (1904) and of body with 7 pairs of setae plus a post- subsequently "pajaroello" by Herms (1916) anal median seta (PMS). and "pajahuello" by Herms and Wheeler (1935). We cannot find any interpretative -0. coriaceus is the only species of the meaning for the spelling of either "pajaro- genus in the Western Hemisphere with two nela" or "pajaroello," and prefer to use pairs of eyes located as in 0. savignyi the term "pajahuello" (paja = straw-colored, (Audouin), the type of the genus in Africa. huello = treading, as on lower parts of -0. coriaceus was originally collected in animals' hoof), as originally used by Banks Sonora, Mexico, and is distributed in the (1904) and subsequently by Hems and Wheeler other western states of Mexico southward to (1935). the Isthmus of Tehuantepec, where the 0. coriaceus is commonly found in the natives call this tick lltalajasll(tala =: beddTng grounds of deer and cattle under- destruction, havoc). Berlese (1888) states neath foothill chaparral, scrub oak, cotton- that he saw a specimen taken from cattle wood, manzanita, or mahogany brush areas at hides at Rio Apa, Paraguay. From 1904 185-1850 m elevations along the coast from through 1941 numerous collections of this San Diego County north to Humboldt County. species were made in the coastal regions of This tick can also be found at elevations California (Cooley and Kohls, 1944b), and up to 2450 m under juniper, yellow and pinon intensive surveys in subsequent years have pine, and bitterbrush on the western slope shown that this tick is widespread in Cali- of the Sierra Nevada and in valley areas fornia and occurs in Nevada and southern northeast of the Cascade Range and east of Oregon (Loomis and Furman, 1977). the Sierra Nevada (Map 3). It is recorded Originally called "pajaronela" by the from 50 of the 58 California counties. inhabitants of the Santa Lucia mountain Ticks may be collected the year round in THE TICKS OF CALIFORNIA 15

COa-pan traps (Hokama and Howarth, 1977) These symptoms usually disappear within 48 placed in the soil or tree-litter of animal hours, leaving a small, purplish nodule that bedding grounds below snow level; at higher disappears in 1-2 weeks. In other cases, elevations, ticks are more commonly trapped an umbilicated pustule surrounded by an from late spring to fall. The life cycle inflamed, painful, edematous area develops. from egg to adult normally takes more than More severe allergic reactions appear in one year, with larvae feeding on hosts for persons previously bitten and thus sensi- 9 days and nymphs passing through 7 instars, tized to a substance injected during the each requiring a blood meal that lasts from tick's blood-feeding process. 8 to 100 minutes; adults feed in 5-50 -0. coriaceus is not known to transmit minutes (Loomis, 1961). any disease of man, but on the basis of Fence lizards, Sceloporus occidentalis, circumstantial evidence it is capable of are reported to eat adult 0. coriaceus transmitting the agent of a cattle disease (Garcia, 1963) , and unidentifiG "red ants'' called epizootic bovine abortion ("foothill have been reported to carry off small abortion") (Schmidtmann et al. , 1976) , and nymphs. Ault and Elliott (1979) observed has been shown experimentally to transmit the jumping spider, Phidippus rimator, African Swine Fever virus to healthy pigs attacking nymphs of 0. coriaceus. (Groocock et al. , 1980). Cattle and deer are common hosts for this tick, but blood meals may be taken from almost any warm-blooded animal, including domestic poultry. Man is an accidental Ornithodoros (Alectorobius) dyeri host, and most tick bites on humans are Cooley and Kohls associated with outdoor, rural, and forest (Figs. 47-50) activities. California collection records are given in Table 2. Ornithodoros dyeri Cooley and Kohls, 1940, The pajahuello is best known because of U.S. Pub. Health Serv. Rept. 55:925-928 the severe reaction in humans following (nymph); Cooley and Kohls, 1944b, Amer. exposure to its bite. Initial bites usually Midland Naturalist, monogr. 1:95-98 result in a localized inflammatory reaction (adults); Kohls, Sonenshine, and accompanied by a burning sensation and a Clifford, 1965, Ann. Entomol. SOC. Amer. small nodule that forms around the wound. 58:342-343 (larva).

TABLE 2. California Records of Ornithodoros coriaceus Koch

Host or Source Tick Collections

Adults and/or Nymphs Larvae Unrecorded Instars Falco mexicanus - 1 - Nest of Petrochelidon albifrons 1 - - -Homo sapiens 29 1 16 Equus caballus 2 - - Odocoileus hemionus columbianus 2 - 14 "Deer" 2 4 - --Bos taurua 1 1 2 Deer bed 19 1 1 Cattle bedding area 9 - - Soil and litter 59 - 5 Vegetation 3 - - Not recorded 30 1 2 16 BULLETIN OF THE CALIFORNIA INSECT SURVEY

DIAGNOSIS. Adult and Nymph. Body Ornithodoros (Pavl ovskye 1 1a) he rms i yellow-brown, elongated, sides parallel, Wheeler, Hems, and Pleyer anterior end pointed, posterior end rounded, (Figs. 51-60; Map 4) flattened on top with distinct, parallel, marginal ridges (Fig. 47). Marmnillae Ornithodoros hermsi Wheeler, Hems, and indefinite or absent, although small, Meyer, 1935, Proc. SOC. Expmtl. Biol. irregular elevations present on body sur- and kd. 32:1290-1292 (laiva, nymph, face. Hood absent but with median ridge adult); Kohls, Sonenshine, and Clifford, present from mouth parts to the anterior 1965, Ann. Entomol. SOC. Amer. 58:352- region. Cheeks (Fig. 49) present, broader 353 (larva). anteriorly. Basis capituli broader than Ornithodoros hermsi; Wheeler, 1935, Amer. long, with micromamillae and transverse J. Trop. Med. 15:435-438 (male, female). wrinkles. Hypostome (Fig. 50) short and broad, denticles 5/5, blunt and U-shaped; DIAGNOSIS. Adult and nymph. Body oval, anterior posthypostomal setae long, faintly wider behind, pointed anteriorly (Fig. 51); barbed, reaching to tip of hypostome, and mamillae numerous, moderate in size, elon- posterior second pair shorter in length. gated, not conical , with sinuous radiating Coxal and supracoxal folds present, plus a ridges on sides (Fig. 57); equal in size fold or ridge above the supracoxal fold throughout dorsal surface but smaller in opposite legs 111 and IV (Fig. 48). Spira- median areas than along sides of ventral cle located posterior to coxa IV. Genital surface. Discs present but shallow (Fig. opening of female located in a V-shaped 51). Hood well developed. Cheeks absent. depression (also present in nymphs but Basis capituli as long as wide, surface absent in males). Eyes absent. Anus small, irregular with interrupted transverse ovoid, distant from transverse postanal wrinkles. Hypostome (Fig. 53) apically groove. Legs small, micromamnillated; tarsi notched, corona with nuuerous fine denti- without subapical dorsal protuberances or cles, denticles 2/2 limited to distal third dorsal humps. with 5 files of about equal size. Eyes Larva. Differentiated from those of absent. Anus in a long oval pattern. most other bat-infesting Ornithodoros Dorsoventral groove faint (Fig. 52). Pre- species by reticulations on the capsule of anal and transverse postanal grooves dis- Haller's organ. Distinguished from larvae tinct. Legs smooth, not micramammillated, of 0. yumatensis by 9 pairs of ventral setae with few hairs and all tarsi without dorsal and smaller dorsal plate, compared to 8 hunps but with mild subapical dorsal pro- pairs with large dorsal plate on 0. yumaten- tuberances (Fig. 55). -sis. The hypostome of 0. dyeri arises from Larva. Dorsa (Fig. 59) with dorsal subtriangular median extension and tapers plate, 11 pairs dorsolateral setae and 2 to a point; large denticles 3/3 on distal pairs central setae. Venter with 7 pairs three-fourths and 2/2 posteriorly to base. setae but lacking posteromedian (postanal) The hypostome of 0. rossi arises from a very seta (Fig. 60). Palpi with 5 setae on seg- long extension of the basis capituli, with ment 3. Hypostome (Fig. 58) not arising distance from base of palpi to basal denti- from median extension of basis capituli, cle of hypostane subequal to length of hypo- short, truncate or notched apical 1 y; dent i- stome, as opposed to about half length of tion only on apical half, 2/2. First pair hypostane in 0. dyeri. posthypostomal setae about 2X length of 2nd pair. 0. dyeri is known from bats and their The absence of cheeks and dorsal humps retreats in Arizona, California, Mexico, on all tarsi distinguish adults and nymphs and El Salvador. of this species from all other Ornithodoros species with the exception of 0. sparnus, CALIFORNIA RECORDS. IMPERIAL CO.: 15 which is usually smaller than 0. hermsi and mi. NE Yuma, 4 nymphs, and 1 adult from rock lacks the dorsoventral groove; the corona crevices in mine tunnel, V-20-1940. RIVER- of its hypostane has 6-8 fine denticles, SIDE 0.:Alice Mine (Riverside Mts.) 1 and larva has smaller dorsal plate than that female frcm spider web in mine shaft, V-16- of 0. hermsi: Larvae of 0. hermsi are much 1954 (W. MacDonald). SAB BERNARDINO CO.: simpler to distinguish from 0. sparnus than 16 mi. N Needles, several nymphal cast skins are nymphs or adults. from rock crevices in bat caves, VI-14-1940. 0. hermsi is commonly associated with chipmunk nests, Eutamias spp., in snags and THE TICKS OF CALIFORNIA 17 decayed stmps of trees and in cabins at IX-23-1934; 1 female, VIII-17-1934 (all C. elevations above 1500 m in California; it Wheeler). occurs also in northern Arizona, Colorado, Idaho, Nevada, Oregon, and Washington. Unusual records are from the toad, Bufo Ornithodoros (Alectorobius) kelleyi boreus halophilus, and from the nest of an Cooley and Kohls English sparrow in the southern Central (Figs. 61-63) Valley of California. This species vectors the spirochete of Ornithodoros kelleyi Cooley and Kohls, 1941, relapsing fever, Borrelia hermsi, and Philip U.S. Pub. Health Serv. Rept. 56:912-914; and Davis (1940) have records of this tick Cooley and Kohls, 1944, Amer. Midland biting humans. Human cases of tick-borne Naturalist, monog. 1:117 (distrib. 1; relapsing fever in California are recorded Sonenshine, 1962, J. Parasitol. 48:470- mainly from inhabitants of mountain cabins 485 (all instars described); Kohls, in which ticks were collected from nests of Sonenshine, and Clifford, 1965, Ann. chipmunks, rats, and squirrels. Porter et Entomol. SOC. her. 58:344-346 (dis- al. (1932) identified spirochetes from the trib. 1. blood and organs of chipmunks and squirrels collected in the immediate vicinity of these DIAGNOSIS. Adult and nymph. Closely cabins. resemble those of 0. talaie and 0. concanen- Adults and nymphs of 0. hermsi have been -sis except for the following: body propor- collected in California from February tionally longer (Pig. 611, with smaller through October. The majority of these cheeks (Fig. 63) and a hypostome with sides records (77%), however, represent col lec almost parallel (Fig. 63), compared to adult tions made from May through September, dur- and nymph of 0. concanensis; absence of a ing the period in which mountain cabins are distinct notch between the hood and anterior usually occupied. extension of the dorsal body wall when viewed in lateral profile (Fig. 62) sepa- CALIFORNIA RECORDS. EL DORADO CO.: Lake rates 0. kelleyi from adults and nymphs of Tahoe: 1 male and 1 female from unknown --0. tala je. host, X-18-1947; 2 females from drags (=tick Larva. Much longer (0.95-1.04 mm long flags) , VI-8-1953. INYO CO. : Independence, unengorged) than larva of 0. talaje , vi th 6 mi. W Gray's Meadow campground, 1 nymph hypostome arising from large, conical median from Peromyscus maniculatus, IX-10-1965 extension, compared to small, subtriangular (R.B. Loomis). KERN CO.: Shafter, 4 nymphs median extension found on larvae of 0. con- and 5 adults from English sparrow nest, canensis and 0. talaje; denticles 3f3 on IV-2-1947 (W. Doetschman); Hart Park, 1 distal third of hypostome, compared to 3/3 nymph from ground squirrel burrow, VIII-24- on distal half of hypostome in concanensis 1956 (H. Johnson); Iron Canyon near Garlock, and talaje. 4 nymphs from unknown host, IV-12-1960 (C.A. In addition to the 4 California bat Toschi); Maricopa, 1 female from Bufo boreus records, 0. kelleyi is known from a variety halophilus, 111-2-1968 (J.M. Sheppard). of bats and bat retreats, including bat- MARIPOSA CO.: Yosemite National Park, 1 infested houses in Alabama, Arizona, Arkan- female, IX-10-1948 (Park Service staff). sas, Colorado, Illinois, Iowa, Maryland, MODOC CO: Unrecorded locality, 3 nymphs from Minnesota, Montana, Nevada, New York, Ohio, Spermophilus beldingi, V-13-1949 (Welsh). Pennsylvania, South Dakota, Texas, West MONO CO.: Mammoth Lake, 2380 m elev., 3 Virginia, Utah, and Wisconsin. It has been males, 1 female, 5 nymphs from cabin, recorded also from Alberta, Canada, and VIII-23-1980; island on Mono Lake, 2 females Sonora , Mexico . from nest of California gull, Larus californicus, 1980 (D. Winkler). PLACER CO.: CALIFORNIA RECORDS. ALAMEDA CO.: SW Unrecorded locality, 10 specimens from Livermore, 3 larvae from the bat Antrozous Douglas fir log, IX-13-1941 (R.H.L.); Lake pallidus, 18-5-1945. IMPERIAL CO. : Kane Tahoe, 1 nymph and 2 adults from animal Springs, 8 larvae from A. pallidus, III-29- burrow, tree, and cabin, IV-?-1947; Lake 1945 (D. Constantine). SAN LUIS OBISPO CO.: Tahoe (Kings Beach) , 2 females, 11-1-1949 4.5 mi. NE Shandon, 91 larvae from A- pal- (W. Graef); Forest Hill, 4 nymphs from bed- lidus, IX-20-1947 (R.T. Orr). SANTA CLARA ding in mineshaft, X-10-1973 (K.H. Hansgen). CO., San Jose, 11 larvae from A. pallidus, SAN BERNARDINO CO.: Big Bear Lake: 1 female, V-6-1942 (D. Constantine). VIII-14-1933; 1 male, VIII-17-1934; 1 male, 18 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Ornithodoros (Pavlovskyella) parkeri Cooley (Figs. 64-69; Map 5)

Ornithodoros parkeri Cooley, 1936, U.S. Pub. Health Service Rept. 51:431-433, 1 pl. (adults). Ornithodoros wheeleri McIvor 1937, J. Para- sitol. 23: 365-367.

DIAGNOSIS. Adult and nymph. Both ends of body rounded, but broader posteriorly, similar in appearance to 0. turicata (Figs. 64, 65). Mammillae (Pig. 69) more numerous and smaller than in 0. turicata, larger at sides and largest posteriorly. Hood large and evident from above. Cheeks absent (Fig. 67). Basis capituli (Fig. 66) as wide as long; surface irregular, with transverse wrinkles. Hypostome (Fig. 67) long and rounded apically, with denticles in lateral and median files equal in size 212. Geni- tal aperture of adult at interval between coxae I and I1 (Fig. 66). Eyes absent. Anus in an oval pattern. Legs long, moder- - I_c_- .- -- --....-J ate in size but appear heavier than in 0. I I--- s,I--.*-_-- -__ - - - ~* I turicata, with segments flared distally; mild subapical dorsal protuberances present MAP 5. Distribution in California of on all tarsi except IV; tarsus I with 3-4 Ornithodoros parkeri Cooley, solid circles. dorsal humps (Fig. 68). Larva. Oval, wider anteriorly, without dorsal plate. Hypostome moderately long, Ornithodoros (Alectorobius) rossi rounded apically, with denticles 212. Palp Kohls, Sonenshine, and Clifford segment 3 extending distal to tip of hypostome. Dorsum with 10-12 dorsolateral pairs Ornithodoros rossi Kohls, Sonenshine, and setae and 2 central pairs. Venter with 7 Clifford, 1965, Ann. Entomol. SOC. her. pairs setae and single posterior median 58:347 (larva). seta. DIAGNOSIS. Adult and nymph unknown. 0. parkeri lives in close association Larva. Most structures of the larva with- various rodents, lagomorphs and the similar to those of 0. yumatensis. Nine Burrowing Owl (Speotyto cunicularia) in the pairs of ventral setae and absence of a pair western and northwestern U.S. Davis (1939) of anterolateral setae on tarsus I distin- reported this tick feeding on man. Davis guish 0. rossi from 0. yumatensis. Larvae (1941) also reported that specimens col- also resemble closely those of 0. dyeri, lected from Idaho and Utah were found to from which they are distinguished as des- transmit the spirochetes of relapsing fever. cribed under that species. Feldman-Muhsam (1973) demonstrated autogeny in 0. parkeri. 0. rossi is known only from larvae coll&ted from various species of bats in In California most collections of this Arizona, California, Utah, Mexico, Colombia, species have been made in the Central and Venezuela. In California, Poch6 and Valley, with a few from central and southern Keirans (1975) reported a collection of coastal localities (Map 5). 0. parkeri has larvae from the leaf-nosed bat, Macrotus been collected chiefly from rodents and californicus, in the Turtle Mountains, San their burrows or nests (Table 3) from April Bernardino County. We have not seen any through October. California specimens. THE TICKS OF CALIFORNIA 19

Ornithodoros sparnus Kohls and Clifford not only morphologically distinct from those (Figs. 70-73; Map 4) of 0. hermsi but required from 9 to 15 days to engorge on the guinea pig, whereas larval Ornithodoros sparnus Kohls and Clifford, --0. hermsi commonly feed in less than 1 hour 1963, J. Parasitol. 49:857. on adult mice, rabbits, guinea pigs, or man.

DIAGNOSIS. Adult and nymph. Similar CALIFORNIA RECORDS. SAN BBNITO CO.: to 0. hermsi, except for the following: 2.7 mi. S Panoche, 1 female, 3 nymphs from smaller females to 3.90 X 2.15 nun and males nest and juniper litter in tree, VII-31-1965 to 3.17 X 1.84 mm, compared to 0. hermsi (K. Hom and V.F. Lee). SAN BKRNARDINO CO.: females 5.0 X 3.1 mm and males 3.8 X 2.4 Kramer Hills, 5 mi. S Kramer Junction, 5 mm; tip of corona of hypostome (Pig. 72) females, 8 males from nest of Neotoma with only 6-8 very fine denticles, whereas lepida, IV-15-1962 (D.P. Furman and F.J. corona of 0. hermsi has numerous fine den- Radovsky). ticles; dorsoventral groove absent in 0. sparnus (Fig. 701, present in 0. hermsi. Larva. Markedly different from 0. Ornithodoros (Alectorobius) stageri hermsi: dorsal plate very large, dorsal body Cooley and Kohls surface with 10 pairs setae and venter with (Figs. 74-77; Map 6) 6 pairs; hypostome and palpi much longer than in 0. hermsi, with 212 dentition along Ornithodoros stageri Cooley and Kohls, 1941, entire length. U.S. Pub. Health Serv. Rept. 56:589, figs. 1-2 (male, female, and larva). 0. sparnus has been found only on woodrats- (Neotoma) and deer mice (Peromyscus) DIAGNOSIS. Adult. Body oval, wider in Arizona, California, levada, and Utah. posteriorly (Figs. 74, 751, differentiating Laboratory studies by Davis and Mavros this sDecies from adults and nvmDhs.. of 0. (1956) on specimens collected from woodrat coacantkis, 0. kelleyi, and 0. yumatensi;, nests in 1954 and 1955 showed that these which have sides parallel. Mammillae ticks were biologically distinct from 0. large, few, not crowded (Fig. 74). Discs hermsi: Utah ticks failed to transmit depressed, large and circular. Hood indis- Borrelia hermsi from California, a strain tinct to absent. Cheeks small, variable in which was readily transmitted by 0. hermsi shape (absent in male). Basis capituli from Oregon and other states. Larvae were (Fig. 76) as wide as long, surface irregular

TABLE 3. California Records of Ornithodoros parkeri Cooley

~~ ~ Host or Source Tick Collections

Adults and Nymphs Larvae Unrecorded Instars Gopherus agassizi 1 - - Homo sapiens 1 - - Spermophilus beecheyi 1 - - Spermophilus beecheyi burrow 13 - - Spermophilus burrow 6 1 - Dipodomys sp. - 1 - Deer 1 1 - Animal burrow 6 - 1 Barnyard 1 - - Unknown source 4 1 - Under rocks 2 1 - 20 BUUETIN OF THE CALIFORNIA INSECT SURVEY

Adults of 2. stageri were taken from bats, Myotis velifer and Tadarida mexicana, in Arizona and Texas respectively. Cooley and Kohls (1944b) and Eads et al. (1956) report this species to feed readily on man.

CALIFORNIA RECORDS. IMPERIAL CO.: Senator mine, 21 mi. NE Yuma, adults and nymphs on bat guano, V-21-1940 (G.M. Kohls). MADERA CO. : San Joaquin Experimental Range, 54 larvae from Antrozous pallidus (103 bats), VI1 and VIII-1951 (Augustson and Wood). SAU LUIS OBISPO CO.: Simmler. 6 adulis and 8 nymphs from A. pallidus, IV-14-1951 (R.T. Orr) .

Ornithodoros (Alectorobius) talaje (Gutkin-MBneville) (Figs. 78-81; Map 6)

Argas talaje GuBrin-Mdneville, 1849, Rev. Mago ZOol. 1~342-344, pl. 90 Ornithodoros tala e Neuraann, 1896, MLm. SOC. 2001. France-4- 34-37; Cooley and Kohls, 1944, Amer. Midland Naturalist monog. MAP 6. Distribution in California of: 1:82-88, figs. 38-39, pl. 8 (adults, Ornithodoros stageri. Cooley and Kohls, larva); Kohls, Sonenshine, and Clifford, rectangles; 0. talale (GuBrin-Mbneville) , 1965, Ann. Entomol. SOC. Amer. 58:349- solid circles; 0. turicata (DugBs), half- 350, figs. 34, 35 (larva). circ lee. Alectorobius talaje, Pocock, 1907, in A System of Medicine 2:187-189. Ornithodoros dugesi Mazzotti, 1943, Rev. and wrinkled. Hypostome notched in both Inst. Salub. Enfenn. Trop. 4:371-374. sexes; principal denticles 212 and large in female; 2 short files of small denticles on DIAGNOSIS. Adult and nymph. Body each side of median line in males. Genital pointed anteriorly, with curve of posterior aperture at interval between coxae I and border slightly flattened, sides nearly 11. Eyes absent. Anus elliptical. Legs parallel (Figs. 78, 79). Mammillae larger moderate in size and length, surface smooth at sides, largest on posterior border; small and shining, neither subapical dorsal pro- on venter. Discs large, distinct, in large tuberances nor dorsal humps present (Fig. depressed areas over median area of dorsum; 77). linearly arranged on venter. Intersection Nymph. Cheeks lacking in early stages; of hood and dorsal body wall forming a notch in later stages, cheeks smaller than in (Fig. 81). Hood small, bent ventrally in adult females. contact with cheeks. Cheeks (Fig. 80) oval, Larva. Short, oval, with dorsal plate largest of any California species. Basis large, pyriform, wider posteriorly. Hypo- capituli (Fig. 80) wider than long, with stome lacks long conical base, bluntly irregular transverse wrinkles and micro- pointed apically with denticles 4/4 apical- mammillae. Hypostome short and small, ly, 3f3 medially, 2/2 at base; lateral files notched apically; denticles 2/2 with 4 large large, median files small. Larvae of this ones in each file. Genital aperture of species are distinguished from those of 0. adult between coxae I. Eyes absent. Anus dyeri and 2. yumatensis by absence of reti- in an oval frame. Legs small and surfaces culations on capsule of Haller?s organ. micromammillated; Tarsus I with mild subapical dorsal protuberance, absent on other 0. stageri is found on bats and in bat tarsi; dorsal humps on tarsi absent. retreats in Arizona, California, Oklahoma, Larva. Similar to 0. kelley i., a bat- and Texas. It is also reported from Mexico, infesting species. Larvae of talale have a Nicaragua, Bolivia, Brazil, and Venezuela. short-based but broad hypostome with rounded THE TICKS OF CALIJNRNIA 21 sides and pointed apically, while in kelleyi Kohls, 1944, Amer. Midland Naturalist the hypostome is long and narrow, pointed monog. 1:56-61, figs. (adults, larva); apically, with sides nearly straight distal Kohls, Sonenshine, and Clifford, 1965, to abrupt constriction marking junction of Ann. Ent. SOC. Amer. 58:356-357 (larva). median extension of basis capituli with hypos tome. DIAGNOSIS. Adult and nymph. Body with both ends evenly rounded (Figs. 82, 83). 0. talaje is found on rodents and in Mammillae (Fig. 82) numerous and large, close association with their burrows or smaller in the midventral area, larger at nests in Arizona, California, Florida, posterior margin; mammillae larger and less Kansas, Nevada, and Texas. It is also known crowded than in 2. parkeri. Hood large, from Mexico and Central and South America evident from above. Cheeks absent. Basis and found in close association with wild capituli (Fig. 84) as wide as long, surface rodents. In Panama, however, Dunn (1933) irregular with few transverse wrinkles. collected larvae chiefly from rats, monkeys, Hypostome long, rounded apically, with den- dogs, cats, chickens, a snake (Epicrates ticles 212 and large teeth limited to distal cenchris), and man. But as noted by Fair- hhs of length. Sex pore at interval child et al. (19661, Dunn's specimens may ktwen coxae I and 11. Eyes absent. Anus have been 0. puertoricensis Fox. in an oval pattern. Legs moderate in size The spirochetes of relapsing fever were and length, with segments slightly flared recovered from specimens of 0. talaje col- distally; mild subapical dorsal protuber lected in Arizona and Texas (Davis, 1940; ances present on all tarsi except IV; tarsus Hackie et al., 1946). I (Fig. 85) with 3-4 dorsal humps. Larva. Oval, slightly wider posteri- CALIFORNIA RECORDS. CONTJlA COSTA CO.: orly, without dorsal plate. Hypostome long, W slope Mt. Diablo, 1 nymph by a rodent rounded apically with denticles 212; extends burrow, VIII-6-1938 (T. fitken). FRESNO to or beyond apex of palpal segment 111. CO.: 7 mi. E Hinkler, 1 nymph under granite Dorsum with 10-11 pairs thick, dorsolateral chip, XI-27-1965 (A. Jung). LOS ANGELES setae and 2 central pairs. Venter with 7 CO.: San Cleeaente Island, 1908 (Banks); pairs setae and a posteromedian seta. Palmdale, 2 adults from rotted base of Joshua tree, LV-14-1938 (T. Aitken); Long 0. turicata is found in Arizona, Cali- Beach, female from unspecified host, IV-20- fornia, Colorado, Kansas, New Mexico, Okla- 1957 (0. Roberts); Lovejoy Buttes, adults homa, Texas, and Utah, on various rodents and nymphs from woodrat nest, X-31-1965 and lagomorphs and in their burrows or (S.F. Wood). MADERA CO.: O'Neals (U.C. San habitats, and on man and livestock. In Joaquin Experimental Range), 1 nymph from Florida and Mexico, this species has been Spermophilus beecheyi, IX-7-1939 (T. found in close association with pigs. 0- Kellev). RIVERSIDE CO.: Desert Center, 15 turicata is rarely collected in California. adults- and 2 nymphs from burrow of Spe-mo- -0. turicata can transmit the spirochetes of philus spa, IX-26-1939 (C.B. Philip). SAN relapsing fever, particularly in the hot, BERNARDINO CO.: 13.9 mi. S Baker, 3 nymphs, dry regions of the southwestern states. It 1 larva under rocks, VII-2-1978 (S. Wernan). is quite similar in appearance to 0. TIJLARE CO.: near Terminus Reservoir on parkeri . State Hwy 198, 1 female under rock, XI-26- 1965 (V. Lee); 13.7 mi. S Badger, 2 females CALIFORNIA RECORDS. ALAMEDA CO.: under granite chip, XII-27-1965 (D.O. Yong). Calaveras Dam: 1 adult and 9 nymphs from unknown host, V-31-1940 (R. Holdenried); 1 male from Spermophilus beecheyi, VIII-28- 1942. MS ANGELES 0.: Hi Vista, 65 nymphs from desert tortoise (Gopherus agassizi), (Figs. 82-85; Map 6) VIII-27-1961. MADERA CO.: 20 mi. NE Madera, 1 nymph from 5. beecheyi, VIII-30-1939 (T. Argas turicata Dug&, 1876, El Repertorio Kelley). RIVERSIDE CO.: Thermal, 80 speci- de Guanajuato, Mexico (newspaper), mens from 2. agassizi, X-15-1935 (Harbison) . April 25. SAN BERNARDINO CO.: Joshua Tree, 1 nymph Ornithodoros americanus Marx, 1895, Proc. from 2. agassizi, IV-26-1959 (R.E. Ryclanan). Entomol. SOC. Wash. 3:195-201, 1 pl. TULARE CO.: 8 mi. N Orange Cove on Drake Ornithodoros turicata; Neumann, 1896, Mem. ranch, 1 female and 2 nymphs from 2. Soc. 2001. France 9:31-34; Cooley and beecheyi, V-14-1942 (R.V. Wings). 22 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Ornithodoros (Alectorobius) yumatensis mine, 21 mi. NE Yuma, 1 larva from bat Cooley and Kohls guano, V-21-1940 (G.M. Kohls). RIVERSIDE (Figs. 86-88) CO.: Alice Mine (Riverside Mts.), nymphal skin from spider web in mine shaft, V-16- Ornithodoros yumatensis Cooley and Kohls, 1954 (W. MacDonald). 1941, U.S. Pub. Health Serv. Rept. 56: 592-594, figs. (larva, nymph, male, and female); Cooley and Kohls, 1944, Amer. Genus Otobius N. Banks Midland Naturalist, monog . 1 :7 4-7 9 ; figs. (adults, larva); Kohls, Sonen- Otobius Banks, 1912, Proc. Entomol. SOC. shine, and Clifford, 1965, Ann. Entomol. Wash. 14:99. Soc. Amer. 58:350-351 (larva). Type of genus: Argas megnini DugBs, DIAGNOSIS. Adult and nymph. Body oval, 1884. sides almost parallel, bluntly pointed anteriorly (Fig. 86). Mammillae irregular Adult and nymph dissimilar, sexes in shape, moderate in size and number, but similar. Adult with integument granulated, equal in size in median and peripheral areas no mammillae, discs indefinite; nymph on all surfaces, close together but not striated, set with spines. No change in crowded. Discs moderate in size, mildly pattern of integument at sides. Capitulum depressed; lineal arrangement on venter in distant from anterior margin. Hood preanal, transverse postanal, and median absent. Hypostome of nymph well developed; postanal grooves. Hood indefinite. Cheeks vestigial in adult. Eyes absent in nymph (Fig. 88) large, twice as long as wide, and adult, present or absent in larva. attached along one side, with few short hairs. Capitulum (Fig. 881, when extended, reaching beyond anterior end of body; basis KEY TO OTOBIUS IN CALIFORNIA capituli with surface granulated and faintly ADULTS AND LATE-STAGE NYMPHS wrinkled. Hypostome (Pig. 88) with sides subparallel, apex mildly notched. Denticles Nymphal integument vi th heavy small or faint, lateral files large but spines anteriorly and lighter smaller toward median line. Genital aper- spines posteriorly (Fig. 98); ture of adult at interval between coxae I adult integument with dorsal and 11. Eyes absent. Anus large and ellip- pits separated by distance 2X tical. Legs long and slender, with surfaces pit diameter (Pig. 97). . . . .megnini made irregular by numerous granulations; tarsi (Fig. 87) without subapical dorsal Nymphal integument with spines all protuberances or dorsal humps. Hypostome one size (Fig. 93); adult integu- smaller, with smaller denticles, in male. ment with dorsal pits separated Larva. Body sub-oval. Dorsal plate by distance diameter of 1 pit . . pyriform, widest posteriorly. Capitulum . (...... - . lagophilus large, as long as half length of body. Hypostome long and slender, arising from a conical base nearly as long as hypostome, Otobius la o hilus Cooley and Kohls sointed apically; dentition 313 in apical -94, Hap 7) -ths, 212 near base of toothed portion. kegs long and slender. 14-16 pairs of Otobius lagophilus Cooley and Kohls, 1940, dorsal setae; 8 pairs of ventral setae, plus U.S. Pub. Health Serv. Rept. 55:928, posteromedian seta. figs. 3-5 (male, female, nymph); Kohls, Sonenshine, and Clifford, 1965, Ann. 0. yumatensis is known from Arizona, Entmol. SOC. her. 58:361-362 (larva Caliiornia, Georgia, New Mexico, and Texas, fig.). where infrequent collections have been made from bats and bat retreats. It is also DIAGNOSIS. Adult. Body rounded on both reported from Mexico, Uicaragua, Colmbia, ends (Pigs. 89-90), but less constricted at and Venezuela. the sides behind legs IV than ing. megnini. Integument granular, with dorsal pits sepa- CALIFORNIA RECORDS. IMPERIAL CO.: 15 rated by l pit diameter (Fig. 92). Hood mi. NE Yuma, 2 nymphs, 1 male from rock and cmerostome not apparent. Basis capitu- crevices in mine tunnel, V-20-1940; Senator li (Pig. 91) broad, short as ;no. megnhi. THE TICKS OF CALIFORNIA

Hypostome vestigial. Spiracle ovate, con- females from burrows of Spermophilus vex; eyes absent; anus subcircular. Legs beecheyi, VIII-15-1956 (D. Roberts and E. short, heavy, with moderate subapical pro- Smith); Cantil, 4 nymphs from Lepus califor- tuberances definite on all tarsi, more pro- -9nicus V-9-1954 (B. Prunty); Hart Park, Kern nounced on IV. River, 1 nymph from ground squirrel burrow, Nymph. Body shape and size as in adult, VIII-24-1956 (D. Roberts); 3 mi. W Wasco, generally smaller than nymph of 0. megnini. 29 males, 2 females from ground squirrel Slender spines over entire body surface, burrow, VIII-17-1956 (H. Johnson); Lerdo more abundant and longer anteriorly (Fig. airport, 1 nymph from 5. californicus, 93), shorter posteriorly, in contrast to 111-13-1942; Bakersfield, 1 nymph from L. spines on nymphs of 0. me nini Hypostome californicus, V-1962. LOS ANGELES COT: (Fig. 94) well developed *with 3 3 dentition. Sierra Pelona Valley, 2 nymphs from jack- Spiracle slightly convex instead of conical in shape as in megnini. Legs short and heavy, with subapical protuberances absent or very small on first 3 tarsi but distinct on IV. Larva. Similar in most aspects to larva -L. calif ornicus deserticoia, of 0. megnini but with 7-9 pairs of dorsal SAN JOAQUIN CO.: Lonetree C body setae, 5 ventral pairs, and without Tracy, from rodent burrows: 5 females and eyes. over 200 larvae, VIII-18-1951; 13 males, 11 0. lagophilus is the spinose tick asso- females, and 2 nymphal skins, VI-1952 (both ciatzd typically with Lepus species (jack- E.C. Loomis). rabbits), on which the nymphs are usually attached to the face. Specimens collected from burrows in Mexico were found to be Otobius megnini (DugSs) naturally infected with rickettsiae of Rocky Spinose Ear Tick Mountain spotted fever (Silva-Goytia and (Figs. 95-100, Map 7) Elizondo, 1952). Philip et al. (1955) and Eklund et al. (1955) reported that 0. lago- Argas megnini Dugls, 1884, La Naturaleza philus collected in Nevada and Utah were 6:197, figs. infested with spotted fever rickettsiae and Otobius megnini; Banks, 1912, Proc. Entomol. Colorado tick fever virus respectively. SOC. Wash. 14:99; Cooley and Kohls, Laboratory experiments have demonstrated 1944b, her. Midland Naturalist monog. that larvae and nymphs of 0. lagophilus 1:21 (synonymy, redescription adults, retained tularemia organisms up to 676 days, nymphs, larva); Kohls, Sonenshine, and but transovarian transmission did not occur. Clifford, 1965, Ann. Entomol. SOC. Bacha (1957) reared this tick on Lepus Amer. 58( 3) :362-363 (larva). californicus in the laboratory. The life cycle from egg hatch to ovipositing females DIAGNOSIS. Adult. Body rounded behind, required from 55 to 134 days. Other experi- slightly attenuated anteriorly; broadest at mental hosts such as cottontail and domestic legs I1 and 111, constricted behind leg IV white rabbits, guinea pigs, an adult kanga- (Figs. 95-96). Integument granular; dorsum roo rat (Dipodomys), suckling deer mice with numerous small, slightly depressed (Peromyscus) , and harvest mice (Rei throdon- pits, each separated by 2X pit diameter tomys) were unsatisfactory. (Fig. 97); not distinct on ventral side. 0. lagophilus is known from Alberta, Hood short and broad. Camerostome lined Canaza, to Chihuahua and Coahuila, Mexico. with numerous fine, long hairs. Basis of In the United States it is recorded from capitulum (Fig. 99) very broad and short, California, Colorado, Idaho, Montana, curved to approach reniform shape. Hypo- Nevada, Oregon, Utah, and Wyoming. stome (Fig. 99) vestigial, short, broad, Of 16 collections of 0. lagophilus in concave dorsally and without denticles. California 5 are from jackrabbits, 4 from Genital aperture at level of posterior ends pikas, and 7 from dirt at entrances to of coxae I (Fig. 96). Spiracle circular, ground squirrel and other burrows. These mildly convex. Eyes absent. Anus very collections were made from March through small and circular. Legs short and heavy; June and in August. except for tarsi I, all tarsi with subapical dorsal protuberance moderate. CALIFORNIA RECORDS. KERN CO.: 17 mi. W Nymph. Second instar nymphs, easily Bakersfield, 3 collections of 7 males and 4 distinguished from other ticks, are the 24 BULLETIN OF THE CALIFORNIA INSECT SURVEY

surface. 2 pairs of ocellus-like eyes on dorsum over legs I and 11. Moderately large elongated dorsal plate with 2 pairs of lateral sensilla. Hypostome dentition 212.

0. megnini, the spinose ear tick, is an important pest of livestock and horses in the western United States. It is also known as far north as British Columbia and south through Mexico, with reports of specimens from Argentina, Brazil, and Chile. Intro- ductions of this species have been found in South Africa, India, and Australia, as a result of long-term feeding in host's ears and the rapid, long-distance travel by livestock transport. 0. megnini is known to transmit the Q-Gver agent, Coxiella burnetti, in sheep and dairy stock. Dense ear infestations on livestock will cause intense head-rubbing and loss of hair similar to that caused by lice. Tick-bite areas are subject to secondary infection, and serious auditory and ear problems may result. In California, this species is more common in the warm, arid, livestock ranching MAP 7. Distribution in California of: areas of the Central Valley and the clsmon- Otobius lagophilus Cooley and Kohls, half- tane southern region. It has been collected circles; 0. megnini (Dugb), solid circles. in 38 counties, but recorded distribution within counties is spotty because of failure to report most infestations. There are 228 collections recorded from 11 hosts in Cali- forms commonly seen in ears of domestic fornia (Table 4). Most common hosts are animals. Body shape of fed tick similar to cattle and horses and, in descending order, adult, but lateral constriction often less dogs, man, sheep, cat, elk, and deer. The pronounced. Unfed nymph much narrowed pos- pronghorn antelope, Anti locapra americana, teriorly (Fig. 98). Integument with shiny should not be overlooked as a host, since surface and fine striae continuous over Shad (1958) found this tick in the ears of depressed areas, which are counterpart of antelopes in New Mexico. Non-host collec discs. Spines over entire body surface tions are recorded from the environs fre- except around capitulum (Fig. 98); heavy, quented by livestock. These include barns, rshape spines on dorsum extending in front salt troughs, and dirt in cattle pens. One of spiracles anteriorly, those on ventral collection by the junior author included surface extending from anus anteriorly; adults, nymphs, and larvae taken from under- slender spines on posterior dorsal and neath eucalyptus bark as high as 12 feet lateral areas and on those ventral areas above ground. An unusual collection of 1 not described above. Hood and camerostome nynph was recorded from a swallow nest in absent. Capitulum subrectangular, as broad Aliso Canyon, Orange County, by J. Belkin as long, with few setae at sides in front (UCLA) . and with short, heavy spines on each side Collection data for 0- megnini shows behind. Hypostome (Pig. 100) large, taper- that this species may be collected as nymphs ing, denticles 414 with about 8 per file. throughout the year. Adults are not common- Spiracle conical. Eyes absent. Legs short ly found unless inspections are made under- and heavy, tarsi with subapical dorsal neath wooden salt troughs, between cracks protuberance absent or very small. First in board sidings of barns or in wooden nymphal stage similar to second, but corral posts, and under bark of trees. smaller, more slender, legs and hypostane Collections from these sites suggest nega- half as large. tive geotropism, positive thigmotropism, Larva. Body oval, striated with 9-10 and possible avoidance of light. Larvae pairs of setae on dorsum, 5 pairs on ventral are usually not found unless an inspector THE TICKS OF CALIFORNIA 25

TABLE 4. California Records of Otobius megnini (DugSs)

Host or Source Tick Collections

Adults Nymphs Larvae Unrecorded Instars “Swallow nest” - 1 - - -Homo sapiens 8 1 “Rabbi t” - Marmota monax - Canis familiaris 14 Felis domestica 4 - Equus caballus 21 1 Cervus canadensis 2 - Cervus nannodes 1 Odocoileus sp. 2 - --Bos taurus 127 22 --Ovis aries 4 1 Wooden feed or salt troughs 2 3 Dairy barn 1 1 - - Soil and vegetation 4 1 - Tree bark 1 1 1 Miscellaneous or unknown 7 8 -

makes a thorough skin scraping deep within KEY TO NYMPHS OF IXODIDAE GENERA the ears of hosts. OF CALIFORNIA^ This species was successfully reared in the laboratory on the closely shaved stomach 1 Anal groove distinct, surrounding area of rabbits, and required from 62 to anus anteriorly and extending 118 days for the complete life cycle from to posterior body margin (Fig. egg hatch to ovipositing females (Loomis, 265); no foveae, eyes, or 1961). festoons. . - . . . - - . . . Ixodes Anal groove, if present, never FAMILY IXODIDAE HURRAY surrounding anus anteriorly; foveae present, with or without Dorsal scutum present; marked sexual eyes or festoons...... 2 dimorphism; male scutum almost covering 2(1) Anal groove absent; legs poorly body; female scutum small, covering about developed, short; no marginal half or less of body. Capitulum conspicu- festoons...... Boophilus ous, terminal; fourth palpal segment reduced and typically inserted subterninally in Anal groove distinct or indis- segment 111; porose areas present only on tinct, surrounding anus pos- basis capituli of female. Spiracles posteriorly; legs normal. marginal terolateral to coxae IV. Of sone 650 des cribed species (Hoogstraal, 1973), 29 species in 4 genera are established in ~~ California. ’Adapted from Serdyukova (1955). 26 BULLETIN OF THE CALIFORNIA INSECT SURVEY

festoons1 present (Fig. 165) scutum (Fig. 16); 1 pair of ...... 3 po s thypo s tomal se tae bordering 3(2) Eyes absent; coxa I with internal hyopostome (Fig. 17); sensilla sagittiformia present (Fig. spur, no external spur (Fig. 16) .2 183)...... Haemaphysalis ...... 2( 1) Marginal festoons absent; palpi Eyes present; coxa I with inter- short and thick. Boophilus nal and external spurs (Fig...... 165)...... -4 Marginal festoons present; palpi 4(3) Palpi with segment I not visible variable...... 3 dorsally (Fig. 354), small and 3(2) Eyes absent ...... Haemaphysalis angular ventrally; apices of palpi sharply triangular in Eyes present...... 4 dorsal view (Fig. 354); post- 4(3) Two pairs marginal dorsal setae anal groove distinct (Fig. 355) anterior to sensilla sagitti- ...... Rhipicephalus formia; 11 festoons present . . .Amblyomma Palpi with segment I visible ...... dorsally (Fig. 149), well 3-4 pairs of marginal dorsal developed and of varied shape setae anterior to sensilla vent rally; palpi bluntly sagittiformia; 9 festoons tipped in dorsal view (Fig. present ...... 5 149); postanal groove may be 5(4) Auriculae absent; palpi appear 2- indistinct. .5 ...... segmented, triangular, with a 5(4) Postanal groove distinct; basis sharp tip; 4 pairs of marginal capituli in dorsal view short, dorsal setae anterior to sen- rectangular, with posterior silla sagittiformia. . Rhipicephalus margin nearly straight .Amblyoma 2 . Auriculae present, palpi appear Postanal groove indistinct; basis 3-segmented, with blunt tips; capituli in dorsal view sub- 3 pairs of marginal dorsal triangular or, if subrectangu- setae anterior to sensilla lar, posterior margin concave. sagittiformia (Fig. 16) .Dermacentor ...... Dermacentor Genus Amblyomma Koch KEY TO LARVAE OF IXODIDAE GENERA OF CALIFORNIA^ Amblyomma Koch, 1844, Arch. Naturgesch. (Berlin) 10:223; Robinson, 1926, Ticks: 1 Anal groove present, interrupted a monograph of the Ixodoidea 4:9; Cooley anterior to anus but extending and Kohls, 1944, J. Parasitol. 30:82. posteriorly beside anus; eyes absent; 4-5 pairs of setae on Type of genus. Amblyomma cajennense scutum; 2 pairs of posthyposto- (Fabricius, 1787). mal setae spaced well posterior to hypostomek; sensilla sagit- Amblyomma species have very long mouth- tiformia absent ...... Ixodes parts with segment 2 of the palpi notably elongate. Usually ornate. Eyes and fes- Anal groove absent; eyes present toons present. Postanal groove present. or absent; 3 pairs of setae on Males without adanal plates. Of the approximately 100 described species none is established in California, but 5. ameri- 1Marginal festoons may not be apparent canum, is occasionally found on cattle on fully engorged nymphs. imported from other states. 2A.- americanum occasionally Introduced on imported cattle. Adapted from Serdyukova (1955). 4 1 pair of posthypostomal setae in Ixodes -uriae. THE TICKS OF CALIFORNIA 27

Amblyomma americanum (Linnaeus) Margaropus Neumann 1907 (nec Karsch 1879), Lone Star Tick Arch. Parasitologie 11:223 (Figs. 101-104) Boophilus Salmon and Stiles, 1901, 17th Rept. U.S. Bur. Anim. Ind., Washington: Acarus americanus Linnaeus, 1758, Regnum 419 (description, synonymy); Cooley, animale , ed. 10 :615. 1946, U.S. Pub. Health Serv., Natl. Amblyomma americanum; Koch, 1844, Arch. Inst. Health Bull. 187:9; Arthur, 1960, Naturgesch. (Berlin) 10:229; Cooley and Ticks: a monograph of the Ixodoidea Kohls, 1944, J. Parasitol, 30:87 5 :205 (description, synonymy). (synonymy listed). Type of genus: Boophilus annulatus (Say, DIAGNOSIS. Female (Fig. 102). Palpi 1821). slim, elongate; segment 2 about 2X length of segment 3. Scutum typically with white Boophilus species are inornate, without spot near posterior end (Fig. 102) on other- festoons but with eyes. Palpi short, com- wise inornate surface. Coxa I (Fig. 104) pressed; segments I1 and 111 with transverse with external spur much longer than internal ridges on dorsal and lateral surfaces. spur. Hypostome dentition 414 in California Male (Fig. 101). Palpi (Fig. 103) as species. Female scutum small, but in male in female. Scutum typically with 2 pairs extending length of body. Coxa I (Fig- 111) of semicircular white lines in anterolateral with 1-2 spurs, and in male with long and posterolateral areas; represented in anterior process. All tarsi with terminal some only by interrupted pale lines (Fig. ventral spurs. 101). Coxa I similar to that of female. Nymph. Palpi similar to those of adults. Inornate. Coxae with external Boophilus annulatus (Say) spurs. Basis capituli with lateral margins Cattle Tick parallel. (Figs. 105-114) Larva. Separable from other ixodid genera by presence of eyes in combination Ixodes annulatus Say, 1821, J. Acad. Nat. with 2 pairs of marginal dorsal setae in Sci. Phila. 2:75, front of sensilla sagittifonnia, and --Ixodes bovis Riley, 1869, Reports on the presence of 11 festoons. diseases of cattle in the U.S., p. 168. Boophilus bovis, Curtice, 1891, J. Comp. The lone star tick, which is not known Med. Vet. Arch. 12: 313. to be established in California, is an Boophilus annulatus; Salmon and Stiles, important pest of man, domestic stock, and 1902. 17th Rept. Bur. Anim. Ind.. deer in other areas. It can transmit the Washington, p. 621; Cooley, 1946, U.S; causal agents of Rocky Mountain spotted Pub. Bealth Serv., Natl. Inst. Health fever, tularemia, and Q fever, and can cause Bull. 187:ll (synonymy, literature, tick paralysis in man and dogs. descriptions female, male, nymph, A. americanum is a 3-host tick, all larva); Arthur, 1960, Ticks: a monograph instars of which will attack man and live- of the Ixodoidea 5:219 (synonymy, stock as well as a variety of other mammals. literature, descriptions, biology) - Immature instars also occur cormonly on birds. In the United States the tick is DIAGNOSIS. Female. Unfed specimens established from Missouri to Texas and east- small, 2.75 mm long from tips of palpi to ward to the Atlantic coast. The northern posterior margin, 1.4 mm wide; engorged limit of its breeding range in the eastern specimens up to 12 X 8 mm (Cooley, 1946a). United States is Pennsylvania and southern Basis capituli (Fig. 110) much broader than New Jersey (Bequaert, 1946). In California, long, with lateral margins forming short adults of A. americanum have been collected points; cornua essentially absent. Scutum mainly from cattle shipped to feed lots in wider anteriorly (Fig. 107); anterolateral Imperial County from Texas. margins almost parallel and straight. Coxae without spurs except short rounded external spur on coxa I. Tarsi 11-IV each with sub Genus Boophilus Curtice apical ventral spur (Fig. 112). Male. Length 2.16-2.28 mm from tips of Boophilus Curtice, 1891, J. Comp. Med. Vet. palpi to posterior margin; width 1.26-1.32 Arch. 12: 317. mm (Cooley, 1946a). Body without caudal 28 BULLETIN OF THE CALIFORNIA INSECT SURVEY appendages. Basis capi tuli (Fig. 109) much B. annulatus is of great veterinary broader than long, with definite cornua and importance as a biological vector of Babesia straight posterior margin. Scutum with bigemina, the protozoan parasite causing slightly constricted, sinuous margins in redwater, or Texas cattle fever. posterolateral areas. Adanal and accessory shields (Fig. 106) each with blunt caudal point. Coxa I (Fig. 111) with long anterior Genus Dermacentor Koch process, free terminally. Coxa I with external and internal spurs of subequal Dermacentor Koch, 1844, Arch. Naturgesch. length and variable width. Coxae I1 to IV (Berlin) 10:217-239. without spurs. Tarsal spurs as in female. Dermacentor; Nuttall and Warburton, 1911, Nymph. Basis capituli much wider than Ticks: a monograph of the Ixodoidea long, with short lateral points as in adult 2 :120. (Fig. 114). Cornua absent. Palpi similar Dermacentor; Cooley, 1938, U.S. Pub. Health to those of adults. Hypostome with denti- Serv., Natl. Inst. Health Bull. 171: cles 313. Scutum subpentagonal (Fig. 113), 15-19 length subequal width. Short rounded spur on each of Coxae 1-111; coxa IV without Type of genus: Ixodes reticulatus (Fabri- spur. cius, 1794). Larva. Capitular length from tips of palpi equal to width of basis capituli; Dermacentor species possess eyes and without cornua. Palpi short, broad. Hypo- festoons,. typically._ ornate. Basis capituli stome short, with denticles 2/2. Scutum of adults rectangular, of nymphs triangular subpentagonal, covering about half length or rectangular. Palpi short and relatively of body of unfed specimen. Eyes indistinct. broad. Anal groove indistinct, situated Coxae without spurs. posterior to anus. Coxa I bifid. Coxae I-IV increasing in size progressively. In At one time 2. annulatus was widely engorged females and nymphs, neither fes- distributed in the southern part of the toons nor anal groove may be apparent. United States, including California. Eradi- There are 6 species of Dermacentor which cation programs during the early part of occur in California. Intraspecific varia- the twentieth century eliminated the tick tion in morphology is common, but specimens from the United States, but it continues to may be identified in most cases with the exist in Mexico and occasionally may be aid of a key and illustrations supplemented introduced. Of the 5 recognized Boophilus by text diagnoses. species in the world, E. annulatus is the only one known to have been established in the United States. In California cattle KEY TO ADULTS OF DERMACENTOR IN CALIFORNIA ticks are easily recognized by the characters given for the genus, most of which are 1 Spurs on coxa I with facing edges shared by the nymphs. widely divergent (Fig. 155). In common with other Boophilus species, Scutum with large, deep puncta- the life cycle from larvae to engorged and tions bearing setae; with long mated adults is completed on a single host. rugo-punctate cervical grooves Hooker et al. (1912) reported that at in female parumapertus Dallas, Texas, the minimum period from ...... attachment of the larvae to dropping of the Spurs on coxa I with facing edges first engorged female was 20 days and the parallel or nearly so (Fig. maximum period 59 days. The fertilized 147). Scutum with shallow female drops from the host and deposits punctations bearing setae, about 2,000 to more than 4,000 eggs which without elongate or rugo- hatch in 19 to over 200 days, depending on punctate cervical grooves in the prevailing temperature. Larvae climb female...... 2 to the tips of vegetation and attach to 2( 1) Spiracular plate lacking dorsal passing hosts. The usual hosts are cattle, prolongation (Fig. 118) or but records of other hosts include deer, with broad truncate prolonga- equines, sheep, goats, and buffalo. The tion; goblet cells very large, biology of the cattle tick was described in of uniform size (75-85) albipictus detail by Hunter and Hooker (1907) and by . Hooker, et al. (1912), who included other Spiracular plate oval with dis- pertinent references to the 1i terature a tinct dorsal prolongation (Pig. THE TICKS OF CALIFORNIA 29

130), with goblet cells minute KEY TO "PIIS OF DERMACENTOR IN CALJFORIJIA1 to large, 50 or fewer to more than 100. (Goblet cells large 1 Lateral points of basis capituli in E. hunteri, but spiracular indistinct, short (Pig. 121); plate with narrow dorsal pro- idiosoma . slender (Fig. 1191, longation containing smaller with lateral margins weakly punctations.) (Pig. 137)...... 3 curved...... albipictus 3(2) Spiracular plate with numeroua Lateral points of basis capituli goblet cells, minute, granular distinct, extending beyond in appearance (Pig. 163). ... apices of scapulae (Fig. 139); ...... variabilis idiosoma broad posteriorly, Spiracular plate with medium to with lateral margins strongly curved (Fig. 140) large goblet cells (Pig. 148) ...4 ...... 2 4(3) Male with attenuated dorsal pro- 2(1) Internal spur of coxa I reduced, longation of spiracular plate; not apparent (Fig. 157); ven- goblet cells fewer than 50, tral basal spurs of basis capi- larger than in andersoni or tuli absent or not apparent (Fig. 158). .3 occidentalis ; smaller puncta- ...... tions lacking or sparse toward Internal spur of coxa I small frame but present in dorsal but apparent (Fig. 152) ventral prolongation; female spiracular basal spurs of basis capituli plate similar to male's but distinct (Pig. 151) ...... 4 with broader dorsal prolong& 3(2) Anterolateral edges basis tion (Fig. 137). .hunteri of ..... capituli straight in dorsal Spiracular plate with moderately view; posterior border even. broad to slender dorsal pro- Spiracular plates subcircular longation and goblet cells (Fig. 159) with ring of about distinct but moderate in size, 7 relatively large goblet cells 50 to more than 100, tending around eccentric macula .... to be smaller toward frame and ...... parumapertus dorsal prolongation (Fig. 148) .5 Anterolateral edges of basis ...... capituli slightly convex over 5( 4) Scuta1 punctations relatively anterior margins of palpal uniform in size; gray pattern segment I (Fig. 139); posterior color of scutum, legs, and border irregular. Spiracular palpi dorsally, with promi- plates broader mesally, with nent, reddish-brown, uniformly ring of relatively small goblet "measled" appearance from base cells around macula in broad color around punctations (Fig. end of plate (Fig. 138). ..hunteri 141): cornua of male about as long as broad basally, of 4(2) Spiracular plate small, with female broader than long (Figs. anteroposterior diameter usual- 143, 144) occidentalis ly about 90-111 pm, typically ...... irregularly ovoid (Fig. 150) , Scuta1 punctations of markedly with broader end toward ventral different sizes, larger ones side, but aome subcircular; deeper and up to 4X larger than ring of 6-7 larger goblet cells the more numerous small ones; around eccentric macula. gray pattern color of scutum, External spur of coxa I about legs, and palpi dorsally, with as long as broad; internal spur only a faintly "measled" not deflected outward (Fig. appearance due to minute size 152) ...... occidentalis of the majority of punctations (Fig. 123). Cornua of both Spiracular plate large, with sexes broader than long (Figs. anteroposterior diameter usual- 127, 128) ...... andersoni ly over 125 DIU, typically ovoid

'Modified from Brinton et al. (1965). 30 BULLETIN OF THE CALLFORNIA INSECT SURVEY

to subcircular (Fig. 167); 5(4) Spur on coxa I acute; spurs on group of 6-20 or more larger coxae I1 and I11 not extending goblet cells around eccentric beyond posterior margin of macula. External spur of coxa plate; margins of cervical I longer than broad; internal grooves parallel .....variabilis spur deflected outward (Fig. 166)...... 5 Spur on coxa I sub-acute; spurs on coxae I1 and 111 extending 5(4) Anterolateral edges of basis beyond posterior margin of capituli slightly convex in plate; margins of cervical dorsal view (Fig. 133). Spira- grooves divergent...... hunteri cular plate with about 6-10 moderately large goblet cells around macula and 1 or more Dermacentor albipictus (Packard) rows of small punctatations Winter Tick bordering frame (Fig. 134). . (Figs. 115-122; Map 9) ...... andersoni Anterolateral edges of basis Ixodes albipictus Packard, 1869, Ann. Rept. capituli slightly concave (Fig. Peabody Acad. Sci. 1:65-66 (female). 164). Spiracular plate as Ixodes nigrolineatus Packard, 1869, Ann. viewed with SEM with about 20 Rept. Peabody Acad. Sci. 1:66. relatively small goblet cells Dermacentor variegatus Marx, in Neumann, around macula and several rows 1897, Mem. SOC. Zool. Fr. 10:367-370. of minute punctations bordering Dermacentor albipictus; Banks, 1908, U.S. frame (Fig. 167). ....variabilis Dept. Agric., Bur. Entomol. Tech. Ser. 15:44-45; Cooley, 1938, U.S. Pub. Health Serv., Natl. Inst. Health Bull. 171: 59-64 (male, female, nymph, larva); KEY TO LARVAE OF DERMACENTOR IN CALIFORNIA' Gregson, 1956, Canada Dept. Agric. Sci. Serv., Entomol. Publ. 930:31-33. 1 Dermal sensilla present (Fig. 16) . .2 Dermal sensilla absent...... 4 2(1) Spur on coxa I broadly tapered and sub-acute .....parumapertus Spur on coxa I moderately tapered or acute (Fig. 17)...... 3 3(2) Posterodorsal margin of basis capituli straight to slightly concave in unmounted specimens...... occidentalie Posterodorsal margin of basis capituli appears slightly convex on unmounted apechens...... andersoni 4(1) Basis capituli with lateral points much reduced, not .extending beyond the scapulae; spurs on coxae I1 and 111 inconspicuous...... albipictus Basis capituli with lateral points extending beyond scapulae (Pig. 16); spurs on coxae 11 and 111 distinct (Fig. 17) .. .5

MAP 8. Distribution in California of 'Adapted from Brinton, et al. (1965). Dermacentor andersoni Stiles, solid circles. THE TICKS OF CALIFORNIA 31

Dermacentor nigrolineatus; Banks 1908, U.S. with broad internal spur. Coxae I1 and 111 Dept. Agric., Bur. Entomol. Tech. Ser. with indistinct marginal spurs, or lacking 15 :48-49. spurs. Dermacentor salmoni Stiles, 1910, U.S. Pub.

Health Mar. ksp. Serv., Hyg. Lab. Bull. &I). albipictus adults differ from other 62:55-60. species in California in the shape of the spiracular plate and the larger goblet cells DIAGNOSIS. Female (Fig. 116). A of the plate. Nymphs differ from other medium to large Dermacentor, 2.8-5.5 rrrm long Dermacentor species in lacking sharp lateral from tip of scapula to caudal end, 1.6-3.5 points on the basis capituli, which is thus mm wide; engorged specimens up to 17 X 12 roughly rectangular rather than triangular. mm, with olive green body. Silvery-gray Larvae differ from other species by reduced pattern color of dorsal capitulum, scutum, lateral points on the basis capituli and and legs predominant or absent. Scutum with lack of dermal sensillae on the dorsal reddish-brown base color in some populations aspect of palpal segment 11. In California, limited to spots around eyes, cervical larvae and nymphs of Dermacentor occurring grooves and their posteriorly diverging on large herbivores in the winter are almost extensions, and pair of lines parallel and always E. albipictus. Adult Dermacentor lateral to these extensions. Scutum (Pig. species commonly seen on large herbivores 116) longer than wide, with section pos- in the winter season include both albipictus terior to eyes narrower than in other Cali- and occidentalis. fornia species; subangulate laterally. Variation in ornamentation and degree Punctations not markedly different in size. of sclerotization is pronounced in different Coxa I with spurs of variable length, but populations of E. albipictus. Cooley (1938) proximal edges parallel to slightly diver- noted the predominance of the unornamented gent. Coxae I1 and 111 with internal spur form ("nigrolineatus") in the south and east small, broad; external spurs large, acute and the ornamented form in the western (Fig. 117). Coxa IV lacking internal spur, United States. Both types occur in Califor- but external spur large, acute. Spiracular nia. Similarly, there is considerable plates (see Fig. 118) typically rounded, variation in size and shape of the spiracu- subcircular to ovate, only occasionally lar plate. with rudimentary broad, truncate, dorsal The winter tick is a widely distributed projection; with goblet cells much larger species in North America. It is a one-host than in other Dermacentor species and with- tick vhich occurs primarily on large herbi- out smaller perforations near frame such as vorous mammals during the fall, winter, and seen in E. occidentalis. early spring months. In parts of California Male (Fig. 115). A medium-sized Derma- the tick is a severe pest of horses during -9centor 2.8-6.1 X 1.7-3.9 mm. Silvery-gray the winter. Heavily infested horses may pattern color variable as in female. In become emaciated and anemic and develop an well-ornamented males, pseudoscutum evident. abdominal swelling known as "water belly." Punctations of scutum of moderate difference This tick can transmit anaplasmosis of in size, but not obvious. Coxal spurs and bovines, but is not considered as important spiracular plates as in female. a vector as other Dermacentor species. Nymph. Slender, elongate, with sides The larvae of the winter tick become almost parallel (Fig. 119). Basis capituli active in September or later and typically (Fig. 121) roughly rectangular dorsally, attach to large herbivores such as horses with at most rudimentary lateral points. or deer. They feed and molt on the same Scutum length and width subequal, with pos- host, and the resultant nymphs remain on terior tip narrowly rounded. Spiracular the host to feed and molt to the adult males plates (Fig. 122) large, subcircular, with and females. Nymphs and larvae typically 15-20 large, distinct goblet cells. Coxa I may be found on their hosts from September with external spur of moderate size, inter- to April, although they may each engorge nal spur small. Coxae I1 and 111 with only and molt in as few as 9 days. Female ticks small external spurs. Coxa IV with external drop from their hosts after engorging for spur small or absent (Fig. 120). 8-30 days to oviposit on the ground. Adult Larva. Subcircular body shape. Basis ticks are found most frequently on their capituli with reduced lateral points not hosts during the winter but occur in lesser extending laterally beyond apices of scapu- numbers as late as June and as early as lae. Dermal sensillae absent. Sensillae September. porosa and sagittifomia present. Coxa I Eggs hatch in spring and typically pro- 32 BULLETIN OF THE CALIFORNIA INSECT SURVEY duce larvae which diapause until fall. tions of adults in California through this Wright (1969) indicated that activation of method. the diapausing larvae may be a function of photoperiodicity. The unfed larvae may Dermacentor andersoni Stiles survive for up to 346 days, according to Rocky Mountain Wood Tick Bishopp and Wood (1913). (Figs. 123-134; Map 8)

California collections of 2. albipictus Dermacentor venustus Marx, in Neumann, 1897, are most concentrated around the coastal Mem. SOC. 2001. Fr. 10:365; Intl. Com, range in the north-central part of the State Zool. Nomencl. 1978, Bull. Zool. and on the western foothills of the Sierra Nomencl. 35:88 (Opinion 1107, Suppres- Nevada, but this tick has been found in sion of Dermacentor venustus Marx). widely scattered areas from the extreme Dermacentor andersoni Stiles, 1908, U.S. north to the southeast. Pub. Health Mar. Hosp. Serv. Rept. As seen in Table 5, California collec- 23: 949. tions of E. albipictus- are most numerous Dewacentor modestus Banks, 1909, Proc. from horses and deer. The relatively few Entomol. SOC. Wash. 10:170-171. collections from cattle indicate that cattle Dermacentor andersoni; Cooley, 1938, U.S. are not common hosts of the winter tick in Pub. Health Serv., Natl. Inst. Health California. The single collection of 2 Bull. 171:31 (male, female, nymph, males from a jackrabbit reported by Lane et larva); Gregson, 1956, Canada Dept. al. (1981) represents the only record of Agric. Sci. Serv., Ent. Publ. 930:28; this tick from a lagomorph. One would not Intl. Com. 2001. Nomencl. 1978, Bull. expect to collect adults of E. albipictus Zool. Nomencl. 35:88 (Opinion 1107, by dragging, or flagging, vegetation. This Conservation of Dermacentor andersoni is reflected by a total of only 3 collec- Stiles, 1908).

TABLE 5. California Records of Dermacentor albipictus (Packard)

Host or Source Tick Collections

Adults Nymphs Larvae Unrecorded Instars

Circus cyaneus 1 I - - Lepus californicus 1 - - - Spermophilus sp, - 1 - - -Felis concolor 1 - - Equus caballus 41 14 2 3 Odocoileus hemionus columbianus 14 18 10 Odocoileus hemionus 6 4 1 1 Odocoileus sp. 15 3 1 1 Cervus nannodes 1 - - - Cervus roosevelti 1 - - - --Bos taurus 9 3 - 1 Antilocapra americana 1 - - - -Ovis canadensis 1 I - - Vegetation (tick drag) 3 - 3 - Under rock 1 - - - Unrecorded host 5 - - 2 THE TICKS OF CALIFORNIA 33

Male (Figs. 123, 124). 2.5-6.1 X 1.6-4.1 mm, broadly ovoid. Ornamentation variable as described for female. Cornua (Fig. 127) as in female but slightly longer and with apices subacute. Scutum with cervical grooves deep and up to 3X longer than wide; punctations as in female. Spira- cular plates as described for female, with fewer goblet cells (Fig. 129). Coxae as in female, usually progressively enlarged from anterior to posterior (Pig. 124). Nymph. Basis capituli dorsally sub triangular, with lateral sharp points extending at almost right angle to longitudinal axis of body past apices of scapulae and with anterolateral margins slightly convex (Fig. 133); distance between apices of lateral points 340-360 pm; ventrally with short basal spurs (Fig. 132), in some reduced to small rounded projections. Scutum slightly longer than wide to wider than long, strongly convex posteriorly, covering less than half of idiosoraa. Spira- cular plate (Fig. 134) subcircular, rela- ______-- -. .- - ___ - ...l...,h tively large (representative diameter 129 um); macula eccentric; goblet cells numer- MAP 9. Distribution in California of: ous, small, with ring of larger goblet cells Dermacentor albipictus (Packard), solid around macula. Coxa I (Fig. 132) with circles; 2. hunteri Bishopp, half-circles. external spur large and much longer than wide (representative measurement: 73 X 54 pm); internal spur distinct, small, with DIAGNOSIS. Female (Figs. 125, 126). A broadly pointed apex deflected outward. medium to large Dermacentor measuring about Coxae I1 and I11 with small external spurs 2.8-5.4 X 1.9-3.7 nun; engorged specimens up (Fig. 132). Coxa IV with external spur very to 16 X 10 mm. Silvery-gray ornamentation small or absent (Pig. 134). pattern superimposed on a reddish-brown base Larva. Ovoid, tapering anteriorly. color typically on scutum, dorsal aspects Basis capituli with subacute, broadly of legs, and to lesser extent on basis tapered lateral points extending laterally capituli. Degree of ornamentation variable. beyond apices of scapulae; distance between Cornua (Fig. 128) typically much broader apices of lateral points 188-193 um; antero- than long, rounded apically. Scutum with lateral margins irregularly curved and cervical grooves 3-4 X longer than wide; posterodorsal margin slightly convex as seen punctations of 2 distinctly different sizes: in unmounted specimens. Palpal apex sub- relatively few large deep ones, and often acute. Dermal sensillae on dorsal surface bearing a seta, and more numerous small of palpal segment 11. Internal spur on punctations without setae; small punctations coxa I subacute. Coxae I1 and 111 each with inconspicuous and usually seen only on orna- small broad spur. mented parts of scutum as minute reddish dots. Spiracular plates (Fig. 130) oval Adults of E. andersoni resemble E. &- with moderately broad to slender dorsal dentalis, from which they may be distin- prolongation; typically over 100 goblet guished by the larger size of the small cells of moderate size subtended by numerous scuta1 punctations on occidentalis, which smaller punctations toward frame and dorsal in well-ornamented males give the latter a prolongation. Coxa I (Fig. 131) bifid with distinctly measled appearance. In both inner spur broader than external spur, with sexes the cornua of occidentalis are longer, facing margins subparallel, apices rounded. and in the female more pointed, than in External spurs on coxae 11-IV about as broad andersoni. Adults of the similar-appearing at base as long; internal spur on coxae 11 -D. variabilis differ in having spiracular and 111, short, broad, rounded; no internal plates with much smaller and more numerous spur on coxa IV. goblet cells. Nymphs of E. andersoni may 34 BULLETIN OF THE CALIFORNIA INSECT SURVEY be difficult to separate from those of e- ern Inyo County in the Bishop, Big Pine dentalis or variabilis by any one character, Creek, and Round Valley areas, a popular but the combination of characters described recreational region. above will usually serve to identify them. While 25 hosts of the Rocky Mountain Larvae of occidentalis usually have the wood tick are recorded from California posterodorsal margin of the basis capituli (Table 61, the majority of collections have straight to slightly concave as seen in been made by flagging vegetation. Adults unmounted specimens, rather than slightly of this tick were identified by us from 5 convex as in andersoni. In the generally of 167 bears (Ursus americanus) examined by larger larvae of andersoni, compared to D. Graeber from 1974 to 1978 for ectopara- occidentalis, there is a greater distance sites in Yosemite National Park. Infested between lateral points of the basis cavi- bears were found at Tuolumne Meadows at an tuli. Larvae of E. parumapertus have a much elevation of 2,630 m and on the floor of broader and more rounded internal spur on Yosemite Valley at 1,225 m elevation. In a coxa I than do andersoni. similar study made in Sequoia National Park. -D. andersoni is notorious as a vector no E. andersoni were found on 73 bears of Rocky Mountain spotted fever in the examined from May to December 1980. northwestern United States and western Canada. It is also the only proven vector to man of Colorado tick fever virus, and a Dermacentor hunteri Bishopp vector of the tularemia agent Francisella (Figs. 135-140; Map 9) tularensis, and the Q fever agent Coxiella burneti. The virus of Powassan fever has Dermacentor hunteri Bishopp, 1912, Proc. been isolated from this tick and it has been Biol. SOC. Wash., 25:33 (male, female); incriminated as a vector of anaplasmosis in Cooley, 1938, U.S. Pub. Health Serv., cattle. It is the most important cause of Natl. Inst. Health Bull. 171:45 (all tick paralysis in North America in humans, instars). livestock, and wild mammals. D. andersoni occuss in western North DIAGNOSIS. Female (Fig. 136). Large AmerTca from British Columbia to Saskatche- ticks, 3.5-4.3 X 1.8-2.5 mm. Idiosoma wan southward through North Dakota to New broadest posteriorly at level of spiracular Mexico, Arizona, and California. It is a plates. Silvery-gray ornamentation similar 3-host tick, in vhich larvae and nymphs to that of 2. andersoni. Basis capituli typically feed on small mammals such as with cornua broader than long, with ends ground squirrels, and adults usually feed rounded to subacute. Scutum tapered pos- on large mammals such as deer, sheep, teriorly, almost subangulate; cervical horses, cattle, coyotes, and bear; man is grooves 2X as long as broad; large puncta- attacked readily by adult ticks. The life tions relatively sparse; small punctations cycle is completed in 1-3 years. more numerous but inconspicuous even on D. andersoni is encountered most fre- ornamented areas of scutum. Typically quenily in the spring in California, begin scutum of both sexes with 2 pairs of diago- ning in March and reaching a peak abundance nal marks of base color located postero- of adults in Nay. Adults as well as nymphs median to eyes (Figs. 135, 136). Spiracular may be encountered in lesser numbers, how- plates diagnostic: oval with slender dorsal ever, throughout the summer until October. prolongation slightly broader than in male (Fig. 137); goblet cells large, fewer than In California E. andersoni is largely 50; lacking smaller punctations toward frame restricted to the northeastern and central but with 10-20 small punctations in dorsal eastern portions of the State, in the south- prolongation. Coxae I of both sexes with eastern Cascade Mountains, the Modoc opposing margins of internal and external Plateau, and the eastern slopes of the spurs parallel; coxae 11, 111, and IV with Sierra Nevada. Some collections are external spurs distinct, semi-acute; inter- recorded also from western slopes of the nal spurs of coxae I1 and I11 reduced, Sierra Nevada, where the tick appears to be rounded; coxa IV lacking internal spur. established, but the isolated collections Hale (Fig. 135). Large ticks, 3.9-4.6 recorded from southern California and the X 2.6-3.1 mm. Silvery-gray ornamentation San Francisco Bay area seem to represent similar to that of E. andersoni, but more only specimens introduced from other areas. boldly marked. Basis capituli with cornua The southernmost established populations of about as long as width at base and with the tick in California seem to be in north- subacute tips. Scutum with cervical grooves THE TICKS OF CALIFORNIA 35

TABLE 6. California Records of Dermacentor andersoni Stiles

Host or Source Tick Collections

Adults Nymphs -Homo sapiens Ochotona princeps Sylvilagus sp. Lepus townsendii Lepus sp. Eutamias amoenus Eutamias minimus Eutamias quadrimaculatus Eutamias speciosus Eutamias townsendi Eutamias 9. Marmota flaviventris Spermophilus beecheyi Spermophi lus be Id ing i Spermophilus lateralis Spermophilus sp. Tamiasciurus douglasii Glaucomys sabrinus Peromy scus boy lii Peromyscus maniculatus Neotoma fuscipes Erethieon dorsatum Canis familiaris Ursus americanus Equus caballus Odocoileus sp. --Bos taurus --his aries Vegetation (tick drag) C02 trap House Unrecorded 36 BULLETIN OF THE CALIFORNIA INSECT SURVEY

2X as long as broad; punctations disparate The above diagnoses of 2. hunteri nymph in size: relatively few large punctations, and larva are modified from those of Brinton less conspicuous than in E. amlersoni; small et al. (19651, based on specimens reared in punctations inconspicuous, even on orna- our laboratory. Adults feed readily on the mented areas of scutum. Spiracular plates laboratory rabbit. During an initial and coxal spurs as described for female, attachment period of several days there is except for narrower dorsal prolongation of little feeding, but females undergo a rapid male plates. engorgement period in the last 2-3 days of Nymph. Basis capituli (Pig. 139) sub- attachment, when they reach a body length triangular dorsally with lateral points of 12-1s 11111. prominent, extending beyond apices of scapu- Adults of this rather large and well- lae; ventral basal spurs absent or merely ornamented tick typically parasitize bighorn rounded saliences (Fig. 140). Scutum (Fig. sheep in Arizona, California, Nevada, Utah, 139) short and weakly curved posteriorly, New Mexico, and Mexico, including Baja Cali- about 1 idiosomal length in unfed specimen. fornia. Little is known of this tick's 3 Idiosoma 1.35-1.47 mm long. Spiracular biology nor has it been implicated as a plates (Fig. 138) ovate with broader end vector of disease agents. anteromedial; with eccentric macula in broader end; 110-116 X 85-98 pm wide; goblet In California, 2. hunteri has been col- cells smaller than in adult; ring of larger lected only in the southeastern arid regions cells around macula. Coxa I (Fig. 140) with of the State within the range of Desert external spur as broad basally as long; Bighorn Sheep. With one exception, only internal spur rounded, indistinct. External adult ticks have been collected (Table 7). spurs on coxae I1 and I11 small, marginal, Of the 31 recorded collections from Califor- broad, rounded; coxa IV with spur absent or nia, 27 have been identified by us; 17 of rudimentary. these were taken from 1963 to 1980 at Deep Larva. Basis capituli with lateral Canyon, 3.5 miles south of Palm Desert, points extending beyond apices of scapulae; Riverside County. Several of these collec- dermal sensilla and sensilla porosa absent; tions were made by entomologists beating anterolateral margins irregularly curved. vegetation to collect insects. The majority Scutum about 1 length of idiosoma, with pos- of California collections were made in March .3 terior margin slightly curved convexly; and April, but adults have been taken in cervical grooves each with posteriorly all seasons. diverging margins, but paired grooves con- The single California record of a wild- verging posteriorly. Coxa I with broadly caught nymph of 2. hunteri is for a specimen subacute internal spur; coxae I1 and I11 taken from Neotoma lepida, V-30-80, in the with small, broad, postmarginal spurs. Kelso Mountains, San Bernardino County (S.

TABLE 7. California Records of Dermacentor hunteri Bishapp

Host or Source Tick Collections

Adults -Homo sapiens 5 Neotoma lepida - -Ovis canadensis 3

Vegetation (including tick drag) 10 - Malaise trap baited with COP 1 - Unrecorded 11 THE TICKS OF CALIFORNIA 37 i Dike). Identification of the nymph was confirmed by comparison with nymphs reared in our laboratory from wild-caught females of 2. hunteri. Records from outside of California indicate that deer and more rarely rabbits may serve as at least occasional hosts of 2. hunteri adults.

Dermacentor occidentalis Harx Pacific Coast Tick (Figs. 141-152; Map 10)

Dermacentor occidentalis Marx, in Curtice, 1892, J. Comp. Med. Vet. Arch. 13:226; Cooley, 1938; U.S. Pub. Health Serv., Natl. Inst. Health Bull. 171:39 (all instars). DIAGNOSIS. Female (Fig. 142). A medium-sized tick measuring about 2.9-3.8 X 1.8-2.5 mm. Typical specimens with silvery- gray pattern color prominent on scutum and dorsal surface of legs, with less on basis capituli and dorsal side of palpi. Extent of ornamentation highly variable. Basis capituli much wider than long. Cornua MAP 10. Distribution in California of (Fig. 144) variable, typically slightly Dermacentor occidentalis Marx, solid broader than long, terminating in subacute circles. to rounded point. Scutum about half length of idiosoma; punctations in both sexes (Fig. 146) numerous, with smaller punctations plate (Fig. 148) as in female, but with more (Fig. 141) more conspicuous and larger than pronounced posterolateral projection. Coxal in 2. andersoni and each surrounded by wide spurs as in female. areola of base color producing red "measled" Nymph. fdiosoma maximum length to tips appearance on ornamented areas of scutum of scapulae 1.21 mm (maximum length of 2. (not visible on SEM photos). Cervical andersoni 1.45 mm) (Brinton et al., 1965); grooves (Fig. 145) short, broad, with pos- distance between lateral points of basis terolateral margins undefined. Spiracular capituli about 306-317 pm; ventral spurs of plate broadly oval with short, broad pos- basis capituli distinct, postmarginal (Figs. terolateral prolongation; goblet cells 151, 152); external spur of coxa I moderate- moderate sized, typically fewer than 100, sized, 48 X 40 pm wide (73 X 54 pm in with smaller punctations bordering frame andersoni); internal spur small, distinct and extending into posterolateral prolonga- (Fig. 1521, not deflected outward; spiracu- tion. Coxa I (Fig. 147) bifid, with inner lar plate subcircular to oval (Fig. 1501, spur broader than external spur and inner with longest axis at right angle to long margins subparallel; apices rounded. Exter- axis of tick body; anteroposterior length nal spurs on coxae 11-IV subacute to acute. about 94-113 pm (about 127 pm in andersoni) Internal spurs on coxae I1 and 111 marginal, and with moderate-sized goblet cells (up to rounded. 17 pm diameter). Male (Fig. 141). Length from tips of Larva. Broadly ovoid, tapering anteri- scapulae to posterior margin 2.6-3.9 mm; orly. As described for 2. andersoni, but width 1.6-2.6 mm. Ornamentation as des- palpi rounded apically, posterodorsal margin cribed for female. Basis capituli width of basis capituli straight to slightly con- slightly greater than length. Cornua (Fig. cave, distance between lateral points of 143) large, prominent; length variable, up basis capituli 167-182 pm, and internal spur to as long as wide, terminating in round to on coxa I more acutely pointed. pointed apex. Scutum with cervical grooves deep, rounded, pit-like; punctations as in Immature instars of E. occidentalis are female, but more prominent. Spiracular particularly difficult to distinguish from 38 BULLETIN OF THE CALIFORNIA INSECT SURVEY

-D. andersoni in areas where both species tsia of Q fever, and the causal agent of occur. Adults are usually distinguishable tularemia. Lane et al. (1981) isolated as desc.ribed under E. andersoni. Although spotted fever group rickettsiae from E. hybridization between male occidentalis and occidentalis collected in Mendocino County. female andersoni has been described (Oliver et al., 19721, the authors concluded that The Pacific Coast tick occurs throughout this very rarely happens in nature. most of California, except for the south- The Pacific Coast tick is known only eastern desert region and parts of the San from California, Oregon, and northern Baja Joaquin and Sacramento valleys. Collec- California, Mexico. Adult ticks occur tions, primarily of adults, are recorded commonly on cattle, horses, deer, and man. from 54 of 58 counties. As indicated in Both sexes of E. occidentalis remain Table 8, man and deer are the most frequent- attached to their hosts for extended ly recorded hosts of the adult ticks. In periods. Oliver and Brinton (1972) deter- part this reflects the results of a sys- mined that the male usually requires 5 days tematic survey of the ectoparasites of deer attachment before becoming sexually potent. conducted by Westrom (1975). Over a two- Larvae and nymphs normally feed on various year period 59 of 72 (82%) black-tailed deer rodents and other small mammals. It is our examined at the Hopland Field Station in impression that nymphs of E. occidentalis Mendocino County were found to be infested attack humans more frequently than indicated with this tick. It was the most frequently by the record; the irritation produced by encountered tick on the deer of that area. their feeding is easily mistaken for that Of 2,112 E. occidentalis collected from produced by other arthropods such as deer, all but 2 nymphs and 2 larvae were spiders. The total impact of this tick on adults. California records of adult ticks the health of man and livestock is incom- from man, horses, and cattle are common, pletely known, but it is associated with a but verified collections from dogs are formidable list of diseases. E. occiden- infrequent. This is surprising, because talis causes tick paralysis in cattle, adults of this tick are so commonly col- horses, and deer in several different areas lected from vegetation in areas frequented of California and is associated with the by dogs. Adults of E. occidentalis were transmission and spread of bovine anaplas- identified by us from 2 of 167 bears mosis. It has been found naturally infected examined for ectoparasites by D. Graeber with Colorado tick fever virus, the ricket- from 1974 to 1978 in Yosemite National Park,

TABLE 8. California Records of Dermacentor occidentalis Marx

__ ~- __ - ~~ Host or Source Tick Collections

Adults Nymphs Larvae Unrecorded Instars -Homo sapiens 108 7 2 4 Ochotona princeps - 1 - - Sylvilagus audubonii 3 1 - Sylvilagus bachmani 1 1 Sylvilagus sp. 5 1

Lepus californicua 4 L Lepus sp. 1 Eutamias merriami 3 Eutamias speciosus 1 Eutamias townsendi 1

Eutamias spb 1 1 1 Spermophilue beecheyi 8 55 12 Spermophilus beldingi 1 THE TICKS OF CALIFORNIA 39 i TBBLE 8 continued

Host or Source Tick Collections

Adults Unrecorded Instars "Ground squirrel" 1 - Tamiasciurus douglasi 1 1 Perognathus californicus 8 - Perognathus sp. - - Dipodomys venustus 2 - Reithrodontomys megalotis - Reithrodontomys sp. 1 - Peromyscus maniculatus 4 Peromyscus truei - Peromyscus californicus - "Deer mouse" - Neotoma fuscipes 1 Neotoma lepida - Neotoma sp. 2 Microtus californicus 3 Microtus sp. - "House mouse" 1 -Canis familiaris 7 Urocyon cineroargenteus 1 Ur sus ameri canus 2 Mustela frenata - Taxidea taxus 1 Mephitis mephitis - Equus caballus 82 Shetland pony - "Mu le" 5 --Sus scrofa 1 Odocoileus hemionus columbianus 59 "Deer" 23 "Fallow deer" 2 --Boa taurus 63 --Ovis ariee 1 Vegetation (tick drag) 418 C02 trap 6 Miscellaneous 3 House 4 Unrecorded 245 40 BULLETIN OF THE CALIFORNIA INSECT SURVEY

,"" .1 but none were found on 73 bears examined ______v- -3%- Y " :.* from May to December 1980 at Sequoia National Park. Incidence of adult E. occidentalis in California is greatest during April and May, although adult ticks are recorded for every month. Nymphs and larvae are most conrmon during spring and summer.

Dermacentor parumapertus Neumann (Figs. 153-159; Map 11)

Dermacentor parumapertus Neumann, 1901, Tc.Zool. Fr. 14:267-268 (female); Banks, 1908, U.S. Dept. Agric., Bur. Entomol. Tech. Ser. 15:45-46 (male, female); Cooley, 1938, U.S. Publ. Health Serv., Natl. Inst. Health Bull. 171:52 (nymph, larva). Dermacentor parumapertus marginatus Banks, 1908, U.S. Dept. Agric., Bur. Ent. Tech. Ser. 15:46.

DIAGNOSIS. Female (Fig. 154). A relatively small Dermacentor, 2.2-3.4 mm long from tips of scapulae to posterior margin. MAP 11. Distribution in California of Typically brown with gray pattern restricted Dermacentor parumapertus Neumann, solid to spot on posterior margin of scutum, but circles. variable to extensive pattern. Basis capituli about 2X as broad as long. Cornua short, rounded. Scutum about half length tions large, prominent in lateral grooves. of idiosoma in unfed specimen; posterior Small punctations minute, inconspicuous. scuta1 margins irregularly subangulate; Spiracular plate similar to that of female, cervical grooves deep, elongate, diverging but posterolateral prolongation typically posteriorly, with posterolateral margins narrower. Coxal spurs (Fig. 155) as des- merging imperceptibly with adjacent surface cribed for female. (Fig. 156); median anterior area and areas Nymph. Basis capituli (Fig. 158) sub- anterolateral to cervical grooves elevated; triangular, with lateral points extending punctations large, most numerous along pos- past apices of scapulae; ventral spurs terior cervical grooves; small punctations indistinct. Scutum over length of idio- so minute as to be easily overlooked. soma; cervical grooves elongate, deep Spiracular plate broadly oval with distinct, anteriorly with parallel sides; shallow broad posterolateral prolongation; goblet posteriorly. Spiracular plate (Fig. 159) cells larger in ring around macula, with subcircular to slightly ovoid, with narrower fine punctations in remainder of plate end laterodorsal; macula eccentric, sur- including posterolateral prolongation. Coxa rounded by ring of 6-7 larger goblet cells I bifid, with opposing margins of internal and 1 to several rings of irregular perfora- and external spurs divergent (Fig. 155). tions. Coxa I (Fig. 157) with moderate- Coxae I1 and 111 with external and internal sized external spur but internal spur an spurs; coxa IV with external spur. indistinct swelling. Coxae 11, 111, IV Male (Fig. 153). Length 2.4-3.1 mm from (Fig. 157) with small external spurs, some- tips of scapulae to posterior margin. times reduced on IV. Typically dark reddish-brown with gray Larva. Basis capituli subtriangular, pattern restricted to posterolateral mar- moderately pointed laterally. Palpi rounded ginal spots of scutum, but variable to apically; dermal sensillae present. Cervi- extensive pattern. Basis capituli wider cal grooves distinct, deep anteriorly. Coxa than long. Cornua wider than long, rounded. I with internal spur small, broadly tapered, Scutum with cervical grooves parallel, long, rounded to subacute apically; coxae I1 and deep anteriorly, open posteriorly; puncta- 111 with only small, broad, external spurs. * t I THE TICKS OF CALIFORNIA 41 s’ Adults of E. parumapertus differ from Sci. Phila. 2:77 (male; female, des t all other California species of Dermacentor cribed as Ixodes punctulatus; synonymy). in that the opposing margins of the internal Dermacentor variabilis; Banks, 1908, U.S. and external spurs of coxa I obviously Dept. Agric., Bur. Entomol. Tech. Ser. b- diverge. Nymphs resemble those of E. 15:49; Cooley, 1938, U.S. Pub. Health andersoni and E. occidentalis, but andersoni Sen., Natl. Inst. Health Bull. 171:23 has a much larger external spur on coxa I, ab (all instars described); Arthur, 1960, and both andersoni and occidentalis have a Ticks: a monograph of the Ixodoidea 5:40 well-defined internal sDur on Coxa I. From (synonymy) the very similar nymphs of 2. hunteri, parumapertus differs in having less acutely DIAGNOSIS. Female (Fig. 161). Medium pointed lateral points on the basis capi- to large Dermacentor; unfed specimens tuli, more rounded spiracular plates, and a 2.7-5.2 X 2.2-3.0 mm (Cooley, 1938); smaller external spur on coxa I. Larvae engorged female up to 15 X 10 mu. Basis differ from other species of the genus by a capituli wider than long, with cornua broad, combination of dermal sensilla and a broadly short, and apices narrowly to bluntly tapered, rounded to subacute spur on coxa I. rounded. Scutum with silvery-gray pattern -D. parumapertus does not normally feed of variable extent, typically more extensive on man, but it is considered of great poten- than red-brown base color. Cervical grooves tial significance in the maintenance of (Fig. 161) shallow and constricted anterior- zoonoses. It has been found naturally ly; deep, prominent, long in midsection, infected with Colorado tick fever virus, shallow posteriorly; combined appearance of with Francisella tularensis, and with a apposed cervical grooves hourglass-shaped. rickettsia of the spotted fever group. Large scuta1 punctations prominent; small Experimentally it has been shown to transmit punctations more numerous, but obvious only the rickettsia of Rocky Mountain spotted in patterned areas. Diameter large puncta- fever and the bacillus of tularemia. tions up to 4X small punctations. Scutum t D. parumapertus is widely distributed less than half length of idiosoma; posterior in t?;e southwestern United States and adja- margin angulate. Spiracular plate shape cent regions of Mexico. The adults occur most abundantly on the Black-tailed Jack- f rabbit. Nymphs occur in lesser numbers on jackrabbits, and both nymphs and larvae commonly infest a variety of small rodents (Gastfriend, 1955; Allred and Roscoe, 1956).

In California, 2. parumapertus is widely distributed, particularly in lowland areas of the Central Valley, the mountains, and F Great Basin region in the east, the southeast corner of Lassen County, and the south- 6r eastern arid part of the State. It is also known along the coast from Mendocino County f south to the Mexican border. California 1 records of the tick reflect the recorded E host distribution of Lepus californicus, to which adult ticks primarily are restricted. Immatures occur on a variety of rodents as well as on Sylvilagus and Lepus (Table 9). c California collections of adult E. paruma- f pertus have been limited to March through August. Nymphs and larvae have been taken I in every season except late fall.

Dermacentor variabilis (Say) American Dog Tick (Figs. 160-167; Hap 12) MAP 12. Distribution in California of Ixodes variabilis Say, 1821, J. Acad. Nat. Dermacentor variabilis (Say), solid circles. 42 BULLETIN OF THE CALIFORNIA INSECT SURVEY variable, typically broad with blunt pos- Male (Fig. 160). Medium to large, terodorsal prolongation; goblet cells very 2.3-4.4 X 1.6-2.4 mm (Cooley, 1938). small and numerous, with smaller punctations Silvery-gray pattern of variable extent but near f rarne and in posterodorsal prolongation frequently less extensive than red-brown on (Fig. 163). Coxa I with large inner spur, scutum. Basis capituli as described for roughly triangular; external spur of sub- female. Scutum with cervical grooves deep, equal length, but slimmer; opposing margins short. Punctations and spiracular plate of spurs subparallel. Coxae I1 and I11 with (Fig. 162) as in female. Coxal spurs as in external spurs and smaller, rounded internal female, but no vestige of internal spur on spurs. Coxa IV with small external spur; Coxa IV. internal spur absent or at most represented Nymph. Basis capituli (Fig. 164) sub- by small marginal swelling. triangular with anterolateral margins

TABLE 9. California Records of Dermacentor parumapertus Neumann

Host or Source Tick Collections

Adults Nymphs Larvae Unrecorded Instars -Homo sapiens - Sylvi lagus audubonii - S ylvilagus bachmani 1 Sylvi lagus SQ . 5

" Rabbi t" 1 Lepus californicus 9 Lepus dese rticola - Lepus sp. 37 Eutamias minimus 1 Eutamias sp. - Spe rmophi lus be ldingi 1 Spermophilus lateralis 1 Spermophilus sp. - Perognathus longimembri s 3 Perognathus parvus 2 Dipodomys agilis 1 Dipodomys heermanni 1 Dipodomys merriami 12 Dipodomys ordii 2 Dipodomys pananti ntinus 9 Dipodomys sp . 1 Peromyscus calif ornicus 1 Cani s f ami liaris - Canis latrans - Vegetation (tick drag) - Unrecorded host - THE TICKS OF CAL.IFORNIA 43 straight or slightly concave; lateral points lagomorphs, particularly meadow mice long, slender; posterodorsal margin almost (Microtus spp.). Adult ticks mate on the straight; ventral basal spurs (Fig. 166) host while females feed. Following an variable but distinct. Palpi long, slender. engorgement period of up to 13 days the Scutum (Fig. 164) with posterior margin female drops to the ground, where she may arcuate, extending over one-third to half lay up to 6,500 eggs. The life cycle of of idiosoma. Spiracular plate (Fig. 167) this 3-host tick may be completed in less large, 134 X 162 pm, suboval, with longer than a year, but in the absence of a suit- apex parallel to that of body. Approximate- able host all stages are able to withstand ly 20 small goblet cells, 6-9 pm diameter, relatively long periods of starvation; in surrounding eccentric macula; frame bordered such cases completion of the life cycle may by band of many minute punctations. Coxa I take 2 or more years. The remarkable (Fig. 165) with external spur large, semi- ability of the American dog tick to survive acute; internal spur small, acute to sub- adverse environmental conditions is exempli- acute, distinct. Coxae 11 and 111 with fied by our collection of 2 female ticks small external spurs, those on IV indistinct from an area recently flooded for more than to absent (Fig. 165). 6 weeks to a depth of approximately 12 feet. Larva. Palpi relatively slender. The ticks were caked with fine sediment, Dermal sensillae absent. Basis capituli indicating prolonged immersion, but it is with lateral points prominent; anterolateral possible that they had crawled out of the margins irregularly curved; posterodorsal water on the emergent trunks of trees in margin straight. Scutum extending one-third the vicinity during the course of the flood. to half length of idiosoma; posterior margin shallowly arcuate, irregular. Coxa I with In California, adults of the American subacute internal spur; 11 and 111 each with dog tick have been recorded for every month, broad, indistinct, external spur. with the majority from Uarch through July. Few records of immature instars have been Adults of D. variabilis differ from all made for the State, but larvae and nymphs other species 7f Dermacentor in the finely appear to be active from late winter until granular appearance of the spiracular plate. summer. Collection records indicate that Nymphs closely resemble 2. andersoni, but the American dog tick is a long-time resi- the spiracular plate has smaller goblet dent of California: numerous collections cells, the palpi are slenderer, and the are recorded from a variety of locations anterolateral margins of the basis capituli early in the twentieth century. are not elevated nor is the Dosterior margin- Recorded hosts of D. variabilis in composed of 3 distinct arcs as in andersoni. California are given in Table 10. Dogs and Larvae lack the dermal sensilla seen in humans are commonly attacked by the adult andersoni; they are distinguished from the tick, with a variety of carnivores also very similar larvae of 2. hunteri by a more serving as hosts. D. variabilis occurs most acute spur on coxa I and by a more arcuate abundantly in the coastal ranges from posterior margin of the scutum. northern to southern California; it also D. variabilis is the most important appears to be well established on the floor vector of Rickettsia rickettsii, the causal of the interior valleys, particularly in organism of Rocky Mountain spotted fever, the Sacramento Valley. Several collections in the eastern United States. It is also have been recorded from mountainous areas capable of transmitting to man Francisella of east-central California, including 4 tularensis. It causes tick paralysis in females collected from a black bear at an the eastern United States as well as in elevation of 6,400 ft at Sequoia National California (Robinow and Carroll, 1938; Park, Tulare County, VI-27-1980 (D. Graber). Jessup, 1979). The American dog tick is widespread in the eastern and central United States as well as in California, Oregon, eastern Genus Eaemaphysalis Koch Washington, and northern Idaho. In Canada it occurs in southeastern and south-central Haemaphysalis Koch, 1844, Arch. Naturgesch. regions, and Bishopp and Trembley (1945) (Berlin) 10:237; Nuttall and Warburton, record it from Mexico. 1915, Ticks: a monograph of the Ixodoi- The dog is a preferred host of the adult dea 3:349 (synonymy, literature, diag- ticks, but man and many wild and domesti- nosis); Cooley, 1946, U.S. Pub. Health cated large mammals are attacked. Immature Serv., Natl. Inst. Health Bull. 187: instars feed on a variety of rodents and 30-51. 44 BULLETIN OF THE CALIFORNIA INSECT SURVEY

TABLE 10. California Records of Dermacentor variabilis (Say)

Host or Source Tick Collections

Adults Gerrhonotus sp. - Sturnus vulgaris - Didelphis marsupialis virginiana 1 -Homo sapiens 16 Sylvilagus audubonii 1 Lepus californicus 1 Peromyscus sp. - Microtus californicus 1 Canis familiaris 91 Canis latrans 12 Urocyon cinereoargenteus 7 "Fox" 2 Ursus americanus 1 Procyon lotor 8 Mustela frenata 1 Taxidea taxus 7 'IS kunk" 1 Felis domestica 2 --Felis rufus 10 "Wild cat" 1 Equus caballus 1 "Deer" -1 -~Bos taurus 5 Vegetation (tick drag) 16 Unrecorded source 59

Type of genus: Haemaphysalis concinna Koch. KEY TO HAEMAPHYSALIS SPECIES IN CALIFORNIA

DIAGNOSIS. Small, inconspicuous ticks. Ventral cornua present (Fig. All instars with lateral expansions of 179); hypostome dentition 3/3 palpal segment I1 extending beyond margins in adults (Fig. 179); male with of basis capituli (Fig. 18l>, characteristic short internal spur on coxa IV of no other genus found in California. All (Fig. 180); nymph with basis instars inornate; eyes absent; festoons capituli quadrangular dorsally present. Anal groove posterior to anus in (Fig. 182); larva with lateral adults and nymphs. Male without shields or margins of basis rounded. . . . plates ventrally. Larvae with 2 marginal ...... leporispalustris dorsal pairs of setae anterior to sensillae sagittiformia and 1 pair of posthypostomal Ventral cornua absent (Fig. 170); hypostome dentition in setae. 515 THE TICKS OF CALIFORNIA

adults (Pig. 171); male with long internal spur on coxa IV (Fig. 174); nymph with basis capituli hexagonal dorsally (Pig. 175); larva with lateral margins of basis acute. . chordeilis

Ixodes chordeilis Packard, 1869, Ann. Rept. Peabody Acad. Sci. 1:67 (female). Haemaphysalis punctata var. cinnabarina, Neumann, 1905, Arch. Parasitologie 9:237 (nee Koch, 1844). Haemaphyslis chordeilis; Banks, 1908, U.S. Dept. Agric., Bur. Entomol. Tech. Ser. 15~34,pl. iv, fig. 11. Haemaphysalis cinnabarina; Nuttall and Warburton, 1- a monograph of the Ixodoidea 3:372, figs. 316-320 (nec Koch) (all instars).

DIAGNOSIS. Female. Size and shape similar to H. leporispalustris. Unfed MAP 13. Distribution in California of female 2.8-2.9-X 1.4-1.8 mm (Hooker et al., Haemaphysalis chordeilis (Packard), half- 1912). Basis capituli (Fig. 173) with circles; E. leporispalustris (Packard), porose areas large, oval, depressed, well solid circles. separated; dorsal cornua rudimentary to absent; basal spurs (ventral cornua) absent (Fig. 170). Palpal segment I1 with lateral circular; cervical grooves distinct. Legs angle short, truncated; segment I11 with with only trochanter I bearing dorsal spur. short, retrograde ventral spur (Pig. 17 1). Trochanters without ventral spurs. Coxae Hypostome (Fig. 171) clavate, with small essentially as in female. corona; dentition 5/5. Scutum (Fig. 172) Larva. Basis capituli hexagonal doe oval, widest anteriorly, with large, rather sally; lacking either dorsal or ventral blunt scapulae. Cervical grooves conspicu- cornua. Hypostome rounded apically; denti- ous. Legs lacking ventral trochanteral tion 2/2. Scutum broader than long. Coxae spurs; trochanter I with large, pointed with distinct spur only on 11. dorsal spur; all tarsi with terminal ventral spurs. Coxa I with moderately long subacute H. chordeilis adults are easily distin- internal spur. Coxa IV with spur as large guisKed from those of g. leporispalustris or larger than spur of Coxa 11 or 111. by the lack of a ventral cornua on the basis Male. Essentially same size as unfed capituli and a denticular hypostomal formula female. Shape (Pig. 168) elongate ovoid; of 5/5 rather than 313. A long, prominent widest behind middle. Basis capituli shape internal spur on coxa IV of chordeilis males similar to female, but dorsal cornua dis separates them from leporispalustris. tinct and longer. Palpi and hypostome as Nymphs and larvae of chordeilis are separa- in female (Fig. 170). Scutum (Pig. 168) ble by the hexagonal shape of the basis ovoid, convex, widest behind middle. Legs capituli in dorsal view as well as by the and coxae similar to those of female, except absence of a ventral cornua. that coxa IV with very long acuminate spur €I.chordeilis is primarily a bird tick, (Pig. 174). althGqh it has been recorded from a variety Nymph. Basis capituli (Fig. 175) he? of mammals, including man. It is generally agonal dorsally; cornua absent both dorsally considered to be widespread in the eastern and ventrally. Palpi as in adults. Hypo- United States and throughout Canada except stome clavate; dentition 2/2. Scutum sub- for the Pacific coast wet belt (Gregson, 46 BULLETIN OF THE CALIFORNIA INSECT SURVEY

1956). It has been implicated in the death behind midlevel. Cervical grooves conspicu- of turkeys in the United States. ous. Legs and coxal spurs as in female, with spur of coxa IV short, acute (Fig. CALIFORNIA ReCORDS. H. chordeilis has 180). been recorded rarely in Caiifornia, and from Nymph. Basis capituli and palpi (Fig. only two locations, both near the coast in 182) similar to male, segment 2 of palpi northern California. HARIN 0.:Fort Barry: with 4-8, only slightly barbed hairs on 1 male and 1 female, 1-3-1952, and 1 male, inner margin. Hypostome rounded distally; 1-9-1952, both collections taken from vege- dentition 212 to base. Scutum subcircular; tation by a tick drag (Rocky Mountain Labo- cervical grooves distinct. Trochanter of ratory collection); 1 nymph, 11-4-1952, no leg I with spurs as in adults. Coxae (Fig. other information, but identified by G.M. 183) as in adults. Kohls. SONOMA CO.: Bodega Bay Head, 1 male, Larva. Basis capituli rectangular dor- 1-27-1980, collected from vegetation by tick sally, similar to nymph, including ventral drag (A. McGowan) . cornua. Hypostome similar to nymph, fewer teeth per file. Coxa I with raised lobe on inner side; coxae I1 and 111 each with Haemaphysalis leporispalustris (Packard) definite spur. Rabbit Tick (Figs. 176-183, Map 13) All instars of fl. leporispalustris are separable from 11. chordeilis as described Ixodes leporis-palustris Packard, 1869, Ann. under the latter. Rept. Peabody Acad. Sci. 1:67 (female). The rabbit tick is widely distributed Haemaphysalis leporis-palustris; Nuttall throughout the Bestern Hemisphere. It is a and Warburton. 1915. Ticks: a monograph-- 3-host tick that commonly parasitizes of the Ixodoidea 31387 (synonymy; des- lagomorphs during all stages of its life cription of all instars). history. Larvae and nymphs frequently parasitize a variety of rodents and ground- DIAGNOSIS. Female. Small, broadly frequenting birds, but adults occur almost ovoid tick, widest posteriorly (Fig. 177). exclusively on rabbits. Sonenshine and Length of unfed female, tips of palpi to Stout (1970) reported 40 species of birds posterior margin, 1.71-2.5 mm; engorged as hosts for immature instars of the rabbit female up to 10 X 7 mm (Cooley, 1946a). tick. As demonstrated by George (1971), Basis capituli quadrate, with porose areas the drop-off rhythm of engorged rabbit ticks mildly depressed, oval, well separated (Fig. is adapted to the resting habits of the 181); dorsal cornua present or absent; basal rabbit host. Camin and Drenner (1978) found spurs (ventral cornua) present on posterior host specificity of larval ticks linked to ventral margin as in male (Fig. 179). strato-orientation of the larvae on vegeta- Palpal segment I1 with lateral angle more tion during the host-finding stage. reflexed and longer than in g. chordeilis The rabbit tick is involved in the (Fig. 181); segment 111 with short retro- transmission of the causal rickettsia of grade ventral spur as in male (Fig. 179). Rocky Mountain spotted fever and the bacil- Hypostome clavate, with small corona; denti- lus of tularemia. It is important in main- tion 313. Scutum oval, widest anteriorly taining reservoirs of these infections in (Fig. 177); scapulae rounded, large; cervi- nature, but it is not responsible for direct cal grooves elongate, conspicuous. Legs transmission of these agents to man, since with small ventral spurs on trochanters; it does not bite humans. Lane (personal trochanter I with large, acute, dorsal spur communication, 1980) reports that 2.75% of (Fig. 181); tarsi lacking spurs. Coxa I -H. leporispalustris pools taken in Hendocino with large, rounded internal spur and small County produced antibodies to spotted-f ever- external spur; coxae 11-IV each with short group rickettsiae when inoculated into mice. spur. Tully et al. (1977) reported that a spiro- Male. Body shape and size similar to plasma from E. leporispalus tris experimen- unfed female or slightly smaller (Fig. 176). tally is associated with ocular and other Basis capituli (Fig. 178) shape similar to disease symptoms in laboratory mice and female, dorsal cornua always distinct and rats. longer than in female. Ventral cornua, palpi (Fig. 179) as in female. Hypostome In California, adult and nymphal rabbit as in female except for larger corona (Fig. ticks are found on hosts throughout the year 179). Scutum (Fig. 176) ovoid, wldest but have been collected most abundantly in THE TICKS OF CALIFORNIA 47

the spring. Larvae have been recorded dur- internal and external spurs ing every season except winter. The tick (Fig. 333) ...... has been collected in 35 counties of the ....subgenus Ceratixodes - uriae .State, throughout the coastal range from Without this combination Hendocino County in the north to San Diego of characters. .2 County in the south. It occurs in inland ...... areas around and within the Central Valley, 2(1) Porose areas large, occupying the northeastern Hodoc plateau, and in rela- most of dorsal surface of basis tively arid parts of southern counties. As capituli, almost contiguous or seen in Table 11, almost all collections of fused (Fig. 306). Cornua adult 11. leporispalustris in California have absent. Coxae I-IV with exter- been from jackrabbits or brush rabbits. nal spurs, lacking internal Although the majority of collections of spurs (Fig. 225). Trochanters immature instars are also from lagomorphs, I and 11 with small spurs (Fig. a variety of rodents and birds have been 225) ....subgenus Scaphixodes - 3 found infested with larvae and/or nymphs of the rabbit tick. The 2 collections recorded Without this combination of Erom man and the single collections each characters...... 4 from the dog, wildcat, deer, and cow may 3(2) Hypostome arising from a small not represent specimens feeding on these median extension of basis capi- hosts. tuli; auriculae as bulbous lateral extensions (Fig. 224). Genus Ixodes Latreille ...... howelli

Ixodes Latreille, 1795, Mag. Encycl. J. Sci. Hypostome not arising from a Lett. Arts, 4:18. median extension of basis capi- Ixodes Salmon and Stiles, 1901, 17th Rept. tuli; auriculae as evenly curved, shelf-like lateral U.S. Bur. him. Ind., p. 459. Ixodes Nuttall and Warburton, 1911, Ticks: extensions (Fig. 307) ...signatus a monograph of the Ixodoidea, 2:116. 4(2) Cornua present (Fig. 194). Auri- Ixodes Cooley and Kohls, 1945, U.S. Pub. culae large, posteriorly Health Serv., Natl. Inst. Health Bull. directed spurs or bulbous pro- 184: 7. jections. Palpal article I Ixodes Keirans and Clifford, 1978, J. Med. with large, anteriorly directed Entomol. Suppl. 2:3. spur (Fig. 195). Trochanters with spurs. Hypostome denti- Type of genus: Acarus ricinus Linnaeus, tion 5/5 anteriorly...... 1758. . subgenus Multidentatus - auritulus Ixodes species lack ornamentation, eyes, Without this combination of and festoons. Anal groove surrounding anus characters...... -5 anteriorly. Palpi variable. Coxae of sub- 5(4) Palpi either thick, club-like, equal size, with or without spurs. Males 1ength:width ratio 3:l or less with 7 nonsalient ventral plates. (Fig. 2821, or if greater than Of 36 species of Ixodes recorded from 3:l (angustus, holdenriedi, the United States, 20 are known from Cali- ochotonae, woodi), hypostome fornia. pointed to very narrowly rounded apically (Pigs. 189, 347). Auriculae absent or faint (except for small, defi- KEY TO FEMALES OF IXODES IN CALIFORNIA' nite triangular saliences in some hear lei) ...... 1 Porose areas large but not con- .subgenus Pholeoixodes 6 tiguous. Cornua absent. Pal- ..... - pal article IV visible dorsally Palpi much longer than wide, (Fig. 336). Auriculae absent lengthwidth ratio greater than (Fig. 337). Coxae lacking 3:l (Fig. 248). Auriculae either prominent as angular shelf-like extensions or pos- 'Adapted from Keirans and Clifford (1978). teriorly directed spurs or 48 BULLETIN OF THE CALIFORNIA INSECT SURVEY

TABLE 11. California Records of Haemaphysalis leporispalustris (Packard)

Host or Source Tick Collections

Larvae Unrecorded Instars Lophortyx californica - Melanerpes formicivorus - Chamaea fasciata - - Thry omane s bewi cki 1 - Toxos toma bendirei - - Toxostoma redivivum 1 - Oreoscop tes montanus - Pipi lo e ry throphthalmus 2 Pipilo fuscus 1 Zonotrichia atricapilla 1 Zono trichia leucophrys 1 1 Melospiza melodia 1 2 -Homo sapiens 2 - Sylvilagus audubonii 13 5 S ylvilagu s bachmani 23 14 Sylvi lagus cinerescens - - Sylvilagus nuttallii - - Sylvilagus sp . 17 10

" Rabbit" 10 4 Ory ctolag us cuniculus - -Lepus californicus 17 19 Lepus sp. 17 10 Eutamias maculatus 1 Eutamias sp. Spermophilus bee cheyi Spermophi lus belding i Spermophilus lateralis S permophi lus sp . Neotoma sp. Canis f amiliaris --Felis rufus Odocoileus hemionus --Bos taurus Ground litter - Unrecorded 1 THE TICKS OF CALIFORNIA 49

curved horns (Fig. 2491, or if lO(9) Very small tick (ca. 1.5 mm weakly developed as lateral long), with furrows on poste- saliences (jellisoni, pacif i- rior margin of scutum (Fig. -cus), hypostome rounded api- 310). Barbed setae on all tally .....subgenus Ixodes .15 coxae (Fig. 311)...... soricis 6(5) Basis capituli ventrally with Without this combination of angular hump on each side of characters...... 11 hypostome (Fig. 327). Scutum ll(10)Hypostome 3/3 throughout length; about as long as wide, with lateral denticles flaring (Fig. rugose lateral areas (Fig. 324) 189)...... angustus ...... texanus Hypostome 3/3 only at apical end; Basis capituli without angular 2/2 at base (Fig. 215); lateral hump on each side of hypostome denticles not flaring ...... 12 (Pig. 347) (rounded shoulders on each side of hypostome in lZ(l1)Scutum narrowly elongate (Fig. -I. kingi). Scutum as long or 2501, over 1.2 mm long; lateral longer than wide; lateral areas carinae near lateral margin . . smooth, punctate, and/or rugose . .7 ...... ochotonae 7(6) Space between small porose areas Scutum broadly elongate (Fig. 1.5-2 X greater than width of 2161, about 1.05 mm long; one porose area (Fig. 210). lateral carinae not close to Auriculae, if apparent, lateral lateral margin. .... holdenriedi bluntly triangular protuber- 13(9) Scutum usually distinctly rugose, ances (Fig. 211). Hypostome especially in anterolateral rounded. Coxa I with short, area (Fig. 287). Cornua absent weak internal spur .....hearlei ...... rugosus Space between medium sized to Scutum punctate but not distinct- large porose areas half to ly wrinkled (Fig. 235). Cornua subequal width of one porose present, distinct 14 area (porose areas weak in 2...... soricis and small with deep 14(13)Posterior margin of basis capitu- pores in i. kingi). Auriculae li sinuous between cornua (Fig. absent or small and weakly 298)...... sculptus defined (Fig. 347). Hypostome Posterior margin of basis capitu- pointed, or if rounded, coxa I li straight or evenly curved with strong internal spur .... .8 (Fig. 238) ...... kiwi 8(7) Ventral surface of basis capituli 15(5) Auriculae laterally extended, very elongate, smooth, impunc- sharp-edged protuberances (Fig. tate, not constricted laterally 203). Porose areas diffuse (Fig. 347). Hypostome dental (Fig. 202). Trochanters with formula 2f2 throughout length. spurs (Fig. 201). brunneus Cornua absent. Prominent ..... lateral ridges bordering porose Without this combination of areas (Fig. 346). Scutum (Fig. characters...... 16 345) much longer than wide, 16(15)Auriculae reduced or not apparent with definite lateral carinae (Fig. 263). 17 and large anteriorly projecting ...... scapulae...... woodi Auriculae as definite curved horns (Fig. 317) or posteriorly Without this combination of directed protuberances (Fig. characters. .9 ...... 273)...... 18 9(8) Coxa I with internal and external spurs subequal (Fig. 311) 10 17(16)Genital aperture between coxae .... IV. Transverse sutural line Coxa I with internal spur defi- on venter of basis capituli nitely longer than external usually visible (Fig. 263). (Fig. 297)...... 13 Apex of hypostome bluntly 50 BULLETIN OF THE CALIFORNIA INSECT SURVEY

rounded (Fig. 263). Hypostome without spurs (Pig. 331). denticle formula 4/4 apically. Ventral plates terminally with File 3 of hypostome denticles a fringe of long setae (Fig. extends over half of length. 334). . subgenus Ceratixodes - uriae Tarsus I approximately 0.84 mm long exclusive of ambulacrum. . Without this combination of ...... pacificus characters...... 2 2 Genital aperture at level of 2(1) Scutum smooth, with pit-like coxae 111. Transverse sutural depressions dorsolaterally line not visible. Apex of (Fig. 196)...... auritulus hypostome very narrowly rounded Scutum smooth or rough, but lack- but not acuminate. Hypostome ing 2 pit-like depressions dental formula 3/3 apically. dorsolaterally...... 3 File 3 of hypostome denticles 3(2) Palpal articles 1-3 fused, cornua extends not more than half of absent (Fig. 304). Hypostome length (Fig. 231). Tarsus I rudimentary, denticles weakly approximately 0.54 mm long exclusive of ambulacrum .jellison< developed (Fig. 305). Genital . aperture situated between 18(16)Auriculae as posteriorly directed coxae 11. Coxa I-IV with short spurs (Fig. 273)...... peromysci external spurs; internal spurs Auriculae as curved, retrograde absent (Fig. 222) ...... 4 horns (Fig. 249)...... 19 Without this combination of 19(18)Genital pore at level of poste- characters...... 5 rior margins of coxae IV. 4(3) Hypostome rounded apically (Fig. Hypostome arises from smoothly 305). Terminal end of palpal acuminate medial projection of segment 3 rounded (Fig. 305). . basis capituli (Fig. 317)...... signatus Spur on palpal article I usually broadly pointed, hook-like, Hypostome pointed apically. Ter- with sides not equilateral minal end of palpal segment 3 (Fig. 317)...... spinipalpis pointed (Pig. 223)...... howelli Genital pore at level of coxae 5(3) Hypostome with lateral denticles 111 or 111-IV. Hypostome different in shape and size arises from round-shouldered from median denticles (Fig. medial projection of basis 261)...... 6 capituli (Fig. 249). Spur on Hypostome with lateral and median palpal article I sharp, forming denticles of subequal shape and an isosceles triangle with size (Fig. 205) ...... 9 palpal margin (Fig. 249) .neotomae . 6(5) Without definite cornua (Fig. 261)...... pacificus KEY TO MALES OF IXODES SPECIES With definite cornua (Fig. 246) ...7 IN CALIFORNIA^ 7(6) Median ventral plate about 21 X 2 1 Fourth palpal segment not termi- as long as anal plate. Coxae nal, protruding from ventral I with moderately long, pen- surface of 3rd segment (Pig. ciliform, bluntly pointed 335). Hypos tome deeply internal spur (Fig. 274). indented apically, denticles Body (idiosoma) not over 1.1 obsolete (Fig. 335). Coxae mm long ...... peromysci Median ventral plate about 3X as long as anal plate. Coxa I with long, tapering, pointed Adapted from Keirans and Clifford (1978). internal spur (Fig. 243). Key excludes the male of 1. holdenriedi, Body (idiosoma) 1-1.5 mm long .. .8 which is unknown, and of jellisoni and 8(7) Pseudoscutum demarked by zone of ochotonae, which are poorly known. punctations (Fig. 314). Basal THE TICKS OF CALIFORNIA 51

teeth of hypostome as large as Larger tick (idiosoma over 1.3 largest lateral teeth (Fig. mm long). Palpi elongate (Fig. 316). Body 1.38-1.5 mm long. 343). Hypostome not reduced, ...... spinipalpis with regular crenulations (Pig. 343). 16 Pseudoscutum not demarked by zone ...... of punctations. Basal teeth 16(15)Hypostome with large, regular of hypostome not as large as crenulations overlapping base largest lateral teeth (Fig. of preceding row (Fig. 187) . . 249). Body 1.02 mm long. . neotomae ...... angustus 9(5) Trochanters with spurs (Fig. Hypostome with smaller crenula- 207) ...... brunneus tions not overlapping base of preceding row (Fig. 343). .woodi Trochanters without spurs .....10 .. lO(9) Cornua distinct (Fig. 236). Lateral carinae present on KEY TO NYMPHS OF IXODES SPECIES scutum ...... kingi IN CALIFORNIA^ Cornua indistinct or absent. 1 External spurs vestigial or Scutum without lateral carinae. 11 . absent on all coxae ...... 2 11(1O)Small projection ventrally on External spur present on coxa I palpal segment I. Coxa I with very short, blunt internal (Fig. 349)...... 4 spur. Hypostome with mild 2(1) With only 1 pair of posthyposto- transverse crenulations. Lack- mal setae. Hosts are marine ing well-defined shoulders at birds ...... uriae base of hypostome .hearlei ..... With 2 pairs of posthypostomal Without this combination of setae (Fig. 309). Varied hosts characters...... 12 ...... 3 12(11)Palpi very rugose dorsally. Pos- 3(2) Scutum widest anteriorly. Shoul- terior margin of basis capituli ders of hypostome humped or at sinuous (Fig. 322). Ventrally, right angle to hypostome ven- basis capituli with angular trally (Fig. 329). Distinct hump on each side of hypostome rugose areas laterally on scu- (Fig. 323)...... texanus tum and dorsally on palpi. Without this combination of Hosts primarily mustelids . .texanus characters...... 13 Scutum widest near middle. 13(12)Internal spur of coxa I relative- Shoulders of hypostome not ly long, thin, acuminate (Fig. humped, but gradually tapered. 293). Hypostome with definite Palpi and scutum lacking dis- teeth, not crenulations (Figs. tinct rugose areas. Hosts 283, 295)...... -14 primarily squirrels .....hearlei Internal spur of coxa I relative- 4(1) Internal spur of coxa I absent ly short, not acuminate (Fig. (Pig. 309). Hosts usually 341). Hypostome with denticles marine birds ...... signatus as crenulations (Fig. 343). ...15 Internal spur of coxa I present 14(13)Hypostome with numerous rows of (Fig. 349). Various hosts. ... .5 long, pointed, overlapping 5(4) Hypostome dentition 414 with 4 teeth (Fig. 283)...... rugosus or more denticles in the inner- Hypostome with fewer rows of most file. Segment 1 of palpi short, rounded, widely spaced teeth (Fig. 295)...... sculptus 15(13)Extremely small tick (idiosoma ca. 1.0 mm long). Palpi bul- Modified from Gregson (1956). Nymphs of bous. Hypostome reduced, with I. holdenriedi, jellisoni, and neotomae irregular crenulations. ...soricis are unknown. 52 BULLETIN OF THE CALIFORNIA INSECT SURVEY

with anterior spur but without short, triangular. Spiracular posterior spur (Fig. 199). plate nearly circular: 0.11 X Hosts marine birds ....auritulus 0.10 mm. Hypostome length ca. 0.2 mm. Hosts rodents, lago- Without this combination of morphs, birds .spinipalpis characters...... 6 ..... lO(6) Palpal segment I (Fig. 190) with 6(5) Palpi long and slender (Fig. anterior and posterior spurs 266); 1ength:width ratio or with only a posterior spur. usually exceeds 3.5:l. Hypo- 11 stome apex pointed or narrowly ...... rounded (Fig. 267) (rarely Palpal segment I without spurs; blunt). Auriculae present. ....7 projections, if any, of segment I rounded and lobe-like (Fig. Palpi (Fig. 190) short to moder- 290). ately long, thickset, length: ...... 16 width ratio usually less than ll(1O)Palpal segment I with posterior 3.5:l. Hypostome apex blunt spur but without anterior spur (Fig. 190). Auriculae present (Fig. 257) (ochotonae may have or absent ...... 10 a rudimentary anterior spur). . . 12 7(6) Basis capituli with 2 auriculae Palpal segment I with both ante- on each side (Fig. 209). Hypo- rior (usually, but see kingi) stome pointed apically. Hosts and posterior spurs (Fig. 190). . 13 birds brunneus ...... 12(11)Auriculae faint lateral protuber- Basis capituli with single auri- ances (Fig. 257). Internal cula on each side (Fig. 319). spur of coxa I short, barely Hypostome narrowly rounded larger than external spur. apically (Fig. 319). Hosts Hosts pikas, woodrats ...ochotonae various -8 ...... Auriculae short rounded teeth 8(7) Basis capituli appears subtrian- (Fig. 348). Internal spur of gular in dorsal view (Fig. coxa I much larger than exter- 264). Internal spur of coxa I nal spur (Fig. 349). Hosts about 2X as long as external woodrats...... woodi spur (Fig. 267). Scutum 13(11)Scutum longer than wide. Very slightly wider than long and small tick; scutum 0.40-0.50 X with scapulae short and rounded 0.36 hypostome 0.07-0.08 (Fig. 264). Unfed body length mm; m long. Hosts shrews, moles. .soricis 1.0-1.2 mm. Hosts lizards, small and large mammals, and Scutum as wide or wider than birds ...... pacificus long. Medium-size ticks; Without this combination of scutum 0.45-0.54 X 0.48-0.54 characters...... 9 mm; hypostome 0.15-0.18 mm long . 14 9(8) A very small tick; body ca. 0.65 14(13)Coxa I with internal and external mm long in unfed state; capitu- spurs subequal (Fig. 191). lum length 0.25 mm; width 0.16 Hosts rodents...... angustus mm. Scutum (Fig. 278) length Coxa I with internal spur appre- subequal to width. Scapulae ciably larger than external short and rounded. Spiracular spur (Fig. 241). Hosts pri- plate oval, 0.11 X 0.07 mm. marily rodents...... 15 Hypostome length ca. 0.14 mm. Hosts Peromyscus maniculatus, 15(14)Posterior margin of basis capitu- Rattus rattus, Gerrhonotus li sinuous in dorsal view (Fig. multicarinatus; Channel Islands, 300). Internal spur on coxa I California. .peromysci large (Fig. 297). Spiracular ...... plate with numerous goblet A small tick; body ca. 0.77 mm cells. Hosts burrowing mammals long in unfed state; capitulum ...... sculptus length 0.39 mm; width 0.23 mm. Scutum longer than wide, ca. Posterior margin of basis capitu- 0.60 X 0.51 mm. Scapulae li straight in dorsal view THE TICKS OF CALIFORNIA 53

(Fig. 240). Internal spur on coxa I of moderate size (Fig. 241). Spiracular plate with few small goblet cells (anterior spur of palpal segment I may be present or absent). Hosts rodents ...... kingi 16(10)Scutum (Pig. 289) with lateral carinae, length subequal to width; scapulae short, angular. Auriculae (Fig. 290) curved saliences. Hosts skunk, fox, dog, weasel ...... rugosus Scutum without lateral carinae, broader than long; scapulae lacking. Auriculae (Fig. 226) triangular lateral projections. Hosts birds...... howelli

Ixodes (Pholeoixodes) angustus Neumann (Figs. 184-191; Map 14)

Ixodes angustus Neumann, 1899, Mem. SOC. Zool. Fr. 12:136 (female); Nuttall and Warburton, 1911, Ticks: a monograph of MAP 14. Distribution in California of the Ixodoidea 2:195, 197, figs. 187, Ixodes angustus Neumann, solid circles. 189, 190 (male, nymph, larva).

DIAGNOSIS. Female. Unfed female about 1.74 nm~ long, slightly smaller than Ixodes Nymph and larva. Characterized by pacif icus. Capitulum (Fig. 188) lacks presence of both anterior and posterior cornua. Porose areas rather large, sub- spur on first palpal segment, giving the triangular or suboval. Palpi slightly more ventral aspect a diamond-shaped appearance than 3X longer than wide. Auriculae absent (Fig. 190) typically larger and more (Fig. 189). Hypostome acuminate, pointed, conspicuous than in I. soricis, I. kingi, and with strong sharp denticles 3/3 to base or 1. sculptus. Cooley and Kohis (1945) (Fig. 189). Scutum (Fig. 185) slightly note, however, that these projections are longer than wide, widest at midlevel, broad- variable in angustus nymphs, and that the ly rounded behind. Coxa I with internal anterior horn may be moderately long, and external spurs subequal, short to moder- short, or absent. Coxa I (Fig. 191) with ate in length as in nymph (Fig. 191). Coxae spurs subequal as in adults. 11-IV lacking internal spur but with external spur. Females of similar -I. ochotonae -I. angustus in all instars is a conanon distinguishable by denticles 313 only api- parasite of rodents. It is of no proven cally. Superficially similar to 2. pacifi- economic or public health significance, E, which has hypostome rounded apically although reported occasionally biting man and inner spur of coxa I much longer than or dogs. It is widely distributed in North external spur. America, occurring from Alaska to California Male. Capitulum (Fig. 186) narrower on the Pacific Coast, throughout Canada, behind, with sides and posterior border and from the Pacific to Atlantic coasts of straight, salient; cornua absent; auriculae the United States. (Fig. 187) represented by short lateral Easton and Goulding (1974) note that 5. ridges. Hypostome (Fig. 187) rounded api- angustus is probably a nest-specific tick. cally, with bluntly rounded median and Males typically are found in rodent or lateral tooth-like denticles of subequal insectivore nests, but only rarely on hosts. size and shape. Coxa I with a relatively Apparently the tick does not quest for hosts short internal spur. All coxae with exter- on vegetation, hence it is rarely recovered nal spurs. Scutum (Fig. 186) with large by flagging. punctations on lateral margins. 54 BULLETIN OF THE CAJiIFORNIA INSECT SURVEY

In California 2. angustus has been col- Ixodes (Multidentatus) auritulus Neumann lected in every month, with immature stages (Figs. 192-199, Map 15) recorded in each of the four major seasons. Distribution is primarily in the northern Ixodes thoracicus Neumann, 1899 (nec Koch, to central parts of the State, with no 18441, M6m. SOC. 2001. Fr. 12:149, records south of the Greenhorn mountains. figs. 22, 23 (female) (homonym). In California, 67 collections of 1 to Ixodes auritulus Neumann, 1904, Arch. numerous specimens are recorded from 16 Parasitologie 8:450 (nomen novem for I. identified and 4 unidentified species of thoracicus Neumann, 1899) ; Nuttall, hosts. I. angustus has been taken most 1916, Parasitology 8:314, figs. 15, 16 commonly lrom meadow mice, Microtus species, (nymph, larva). Cooley and Kohls, but it also readily infests deer mice, 1945, U.S. Pub. Health Serv., Natl. Peromyscus species, and other rodents (Table Inst. Health Bull. 184:195. Kohls and 12). Only two nonrodent hosts have been Clifford, 1966, J. Parasitol. 52:815, recorded as infested with I. angustus: figs. 20-24 (male). Scapanus latimanus, the CaliTornia mole, and Bassariscus astutus, the ringtailed cat.

TABLE 12. California Records of Ixodes angustus Neumann

Host or Source Tick Collections

Adults Nymphs Larvae Unrecorded Instars Scapanus latimanus - 2 2 1 Aplodontia rufa 2 1 - - Eutamias speciosus 1 - - - Spermophilus beecheyi - 1 - - Sciurus griseus 1 - - - Sciurus species 1 - - 1 Peromyscus californicus 2 2 1 - Peromyscus boylii 4 - 1 - Peromyscus maniculatus 8 2 - - Peromyscus truei 2 - - - - Peromyscus sp. 2 1 1 Neotoma fuscipes 2 1 - - Microtus californicus 9 1 - - Microtus longicaudus 1 1 - - Microtus montanus 3 2 - - '%louse" 1 - - - Rattus norvegicus 1 1 - - --Rattus rattus 1 - - - "Rodent" 1 - - 2 Bassariscus astutus Vegetation (tick drag) Unrecorded 2 2 - - THE TICKS OF CALIFORNIA 55

internal margins salient; each coxa with a small, pointed external spur. Nymph. Yellowish brown. Strong anteri- or projection on first palpal segment (Fig. 1991, as in female. Cornua present and ventrally basis capituli with large, triangular auriculae (Fig. 199). Coxal spurs essentially as in female (Fig. 198). Larva. Palpal article 1 with a large, acute, tooth-like anterior projection. Auriculae well developed.

-I. auritulus is of no known economic importance. It is primarily parasitic on birds, particularly ground-frequenting passerine and galliform hosts. Its recorded distribution extends from Alaska to the most southerly tip of South America. The 13 collections of A. auritulus from California were taken from birds collected from February to November, indicating year- round activity. The recorded distribution probably reflects areas of ornithological activity rather than the actual distribution of the tick.

MAP 15. Distribution in California of: CALIFORNIA RECORDS. ALAMEDA CO.: Ixodes auri tulus Neumann, half-circ les ; Berkeley, 1 nymph from Zonotrichia e- Ixodes brunneus Koch, solid circles. capilla, XI-26-1954. MARIN CO.: Fort Barry, 1 female from Circus hudsonicus, XI-11-1944; Point Reyes Bird Observatory: 3 females from 5. atricapilla, X-27-1976 (B. Sorrie); DIAGNOSIS. Female. Readily recognized 1 nymph each from Hylocychla ustulata by characteristic combination of features. swainsoni, VIII-24-1976 and IX-8-1976 (B. Dorsally basis capituli with strong cornua Sorrie); 1 female and 2 nymphs from --H. u2 (Fig. 1941, and ventrally with even stronger swainsoni, VII-9-1979; 1 female from B. u. horn-like to bulbous auriculae directed swainsoni, VIII-1-1979; 4 larvae from posteriorly (Fig. 195). First palpal seg- Passerella ileaca, XI-9-1978 (P. Paton). ment with a prominent anterior spurlike MENDOCINO CO.: Ten Mile River, 1 nymph from projection visible both dorsally and ven- Zonotrichia leucophrys nuttalli, V-10-1959 trally, but no corresponding posterior pro- (R.C. Banks). NAPA CO.: Napa Mead Ranch, 1 jection (Fig. 195). Hypostome broadly nymph from Pheucticus melanocephalus, rounded apically, with denticles 5/5 dis- VIII-25-1970. SAN HATE0 CO.: Pescadero, 6 tally, subtended by 414 and 313 consecutive- females from a "blackbird," 11-28-1964 (W. ly to base (Fig. 195). Scutum (Fig. 193) Plummer). SAN FRANCISCO CO.: Farallon widest at middle, broadly rounded behind; Islands, 2 larvae from z. atricapilla, anterolateral margins flattened; scapulae X-27-82, and 2 nymphs from z. atricapilla, reduced or absent; surface smooth; cervical X-30-82 (R. P. Henderson). grooves shallow, forming hourglass pattern. Coxae all with external spurs. Distinct internal spur of coxa I shorter than exter- Ixodes (Ixodes) brunneus Koch nal spur. Internal spur absent on coxa IV, (Figsx209; Map 15) and reduced or not apparent on coxae I1 and 111. Ixodes brunneus Koch, 1844, Arch. Natur- Male. Known from a single specimen gesch. (Berlin) 10:232 (female). collected in Chile and described by Kohls Anastos and Smith, 1957, J. Parasitol. and Clifford (1966). Scutum smooth, with 43:535-537, pls. I, 111, Iv (male, two pit-like depressions dorsolaterally nymph, larva). (Fig. 196). No cornua nor auriculae on Ixodes californicus Banks, 1904, Proc. basis capituli. Coxae (Fig. 197) lacking Calif. Acad. Sci. 3:369, pl. 41, fig. well-defined internal spurs, but postero- 57. 56 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Ixodes kelloggi Nuttall and Warburton, 1908, ground-frequenting birds. It is widespread Proc. Camb. Phil. SOC. 14:396, figs. in the United States and may be expected to 6-8 occur in Canada and Mexico. Males are rarely collected from birds, but they as DIAGNOSIS. Female. Mouthparts rela- well as nymphs and larvae were described tively long, slim (Pig. 202), with palpi 4X and illustrated from colony-reared specimens or more longer than wide and segment 2 much by Anastos and Smith (1957). longer than 3. Hypostome (Fig. 203) narrow -I. brunneus has been reported to affect and pointed, with dentition 414 becoming its avian hosts adversely (Bishopp and 313 and 212 near the base. Cornua short Trembley, 1945). Rickettsia rickettsii, and bluntly rounded. Auriculae, distinctive has been recorded from the ticks in nature for this species, large, laterally extended, by Clifford et al. (1969). This raises the sharp-edged protuberances of somewhat vari- issue of possible transfer of pathogens for able shape (Pig. 203). Dorose areas dif- long distances by ticks attached to migrat- fuse, large (Fig. 202). Coxa I with 2 sub ing birds. equal spurs (Pig, 201). Coxae 11-IV with only external spur obvious, but I1 and 111 In California, the recorded collection with suggestion of internal spur (Pig. 201). sites represent in part locations where Male. Palpi (Pig. 204) short and wide; ornithological activities are most inten- article 2 slightly wider than 3. Hypostome sively pursued. (Fig, 205) 313 with lateral and median den- Hosts from which the tick has been ticles small, blunt, subequal. Basis capi- recorded in California are given in Table tuli (Pig. 205) without cornua, and auricu- 13. lae inapparent. Scutum with prominent marginal grooves, deep and continuous posteriorly. Coxa I with 2 short, blunt, subequal Ixodes (Pholeoixodes) hearlei Gregson spurs. Coxae 11 and I11 with external spur (Figs. 210-213; Map 16) larger than internal spur; coxa IV with external spur only. Trochanters with spurs Ixodes hearlei Gregson, 1941, Caned. Ento- (Fig. 207). mol. 73:220, pl. XV, figs. 1-11 (male, Nymph. Outer palpal margin straight to female, nymph). convex; first segment narrowed in basal third (Fig. 208). Hypostome narrow, DIAGNOSIS. Female. Unfed specimens pointed; denticles 313, then 2/2 to base; oval, wider behind midbody; 2.5 X 1.6 mm. lateral denticles large, pointed. Cornua Yellowish brown. Hypostome with almost large, directed laterally. Auriculae (Fig. parallel sides and a rounded apex; dentition 209) appearing double on each side, each 212 except for coronal area. Hypostome triangular and bluntly pointed laterally. arising from anterior projection of basis Marginal groove of dorsum distinct. Coxa I capituli which originates at inner bases of with internal spur slightly longer than palpi, tapering anteriorly, with mildly external spur; coxae 11-IV with external rounded shoulders (Fig. 211). Basis capi- spur only. tuli with small, widely separated porose Larva. Basis capituli, including hypo- areas (Fig, 210). Cornua weakly developed, stome, similar to that of nymph, palpi Auriculae absent or small, bluntly triangu- relatively shorter and wider. Tvo pairs of lar protuberances (Fig. 211). Palpi thick posthypostomal setae; 3 pairs of marginal and club-like, almost straight on lateral ventral setae; 5 pairs of central dorsal margins but rounded medially; segment 1 with setae; 5 pairs of scuta1 setae. Coxal spurs small, ventral, pointed plate (Fig. 211). similar to those of nymph. Scutum (Pig. 212) about as wide as long, widest anterior to midlevel; mildly rugose The disputed validity of brunneus anterolaterally; punctations moderate; and 11. frontalis Panzer, 1795 was clarified cervical grooves distinct. Coxae appearing bv Homsher and Sonenshine (1975).-- who des- superficially devoid of spurs; coxa I with cribed discontinuous morphological distinc- a very short, rounded internal spur. Geni- tions in the structure of Haller's organ in tal opening between coxae 111. females of the two forms. L. frontalis is Male. Oval, broader behind midlevel, an avian parasite of birds in Europe, but about 2.5 X 1.7 mm. Capitulum lacking occurs on migrating birds in Asia and cornua, auriculae represented by mildly Africa. triangular lateral . projections. Palpi -I. brunneus is a common parasite of short, oval dorsally, lacking a suture THE TICKS OF CALIFORNIA 57

TABLE 13. California Records of Ixodes brunneus Koch

flost or Source Tick Collections

Adults Lophortyx californica - Tyto alba pratincola 1 Stellula calliope 1 Cyanocitta stelleri 1 Thryomanes bewicki 1 Parus inornatus - Toxostoma redivivum - Turdus migratorius 1 "Thrush" 1 Lanius ludovicianus anthonyi 1 Dendroica townsendii 1 Agelaius phaeniceus 1 Piranga ludoviciana 1 Pheucticus melanocephalus 1 Pipilo erythrophthalmus - Carpodacus pupurius 2 Zonotrichia leucophrys 2 Zonotrichia atricapilla 1 Vegetation (-1 1 Unrecorded 2

between segments 2 and 3; segment 1 with a pointed ventral plate easily confused with small, pointed, ventral plate. Hypostome larger posterior spur of other Ixodes such broad at base, tapering anteriorly, mildly as I. sculptus and L. ochotonae. Scutum bilobed apically, denticles weak and appear- widest near middle. Coxae lacking external ing as mild, transverse crenulations. Scu- spurs, but coxa I (Fig. 213) with a short, tum with conspicuous, large, shallow puncta- rounded internal spur, as in female. tions and a lesser number of small punctations. Scapulae short, rounded. Lateral The larva of I. hearlei is undescribed. carinae absent. Coxa I with a short, -I. hearlei is known as a squirrel tick, and rounded internal spur; external spurs very it closely resembles I. marxi, a species short on all coxae. Trochanters without found on squirrels in eastern North America. spurs. -I. hearlei has been recorded from British Nymph. Cornua lacking; dorsal posterior Columbia, Oklahoma, Washington, Montana, margin of basis capituli straight, salient. Texas, and Oregon, in addition to Cali- Hypostome rounded apically and dentition fornia. 212 (Fig. 213). Two pairs of posthypostomal setae (Fig. 213). Auriculae as rounded CALIFORNIA RECORDS. MONTEREY CO.: protuberances anteriorly positioned, almost Hastings Reservation, 3 females from Sciurus adjoining palpi. Palpi (Fig. 213) short griseus, VIII-14-1948 (J.M. Linsdale). and broad; segment 1 with a transverse, PLACER CO.: near Brockway, 3 males and 4 58 BULLETIN OF THE CALIFORNIA INSECT SURVEY

215). Transverse sutural line distinct on ventral side of basis capituli. Auriculae absent. Hypostome (Fig. 215) tapering in distal half to a point; dentition 3/3 in apical Zths, then 2/2 to base. Scutum (Fig. 5 216) oval, widest anterior to middle. Coxa I with internal and external spurs subequal in length, but internal spur narrower (Fig. 217). External spurs present on coxae I1 and 111, but absent on IV. Genital aperture at level of interval between coxae I1 and 111.

Morphologically, L. holdenriedi is very similar to 2. ochotonae, but the sides of the hypostome taper in the distal half instead of remaining essentially parallel until reaching the distal fifth, as in ochotonae.

The only recorded California collection is of 2 females from Thomomys bottae, the Valley Pocket Gopher, at Fitch Mountain, 3 mi. E Healdsburg, Sonoma County, 111-25-1945 (M. Hobmaier). A gopher collected from the type locality in June 1981 by Furman and MAP 16. Distribution in California of: Lane harbored only larvae of I. pacificus. Ixodes hearlei Gregson, solid circles; I. holdenriedi Cooley , rectangles; I. howelii Cooley and Kohls, half-circles. Ixodes (Scaphixodes) howelli Coolev and Kohls (Figs. 218-227; Map 16) females from nest in decayed tree, X-1946 (D. Longanecker); Sawtooth Springs Rd. S Ixodes howelli Cooley and Kohls, 1938, U.S. Truckee, 4 females from Spermophilus Pub. Health Serv. Rept. 53:1616, fig. 1 lateralis, VII-11-1978 (Hansgen and Clover). (female); Kohls, 1947, J. Parasitol. PLUMAS CO.: 1 female from Tamiasciurus 33:57, figs. 1, 2 (male, nymph, larva). douglasi, V-13-1949 (E.W. Jameson, Jr.). SAN BERNARDINO CO.: Seven Oaks, 1 female DIAGNOSIS. Female. A long-legged tick from "gray squirrel," V-9-1936. SHASTA CO. : (Figs. 220, 2211, separable from the closely Manzanita Lake, Lassen National Park, 1 male related marine bird tick, L. signatus, by and 2 females from nest of 5. lateralis, hypostome arising from a short median exten- IX-20-1940 (T. Aitken). SIERRA CO.: 3.7 sion of basis capituli (Fig. 224); extension mi. N, 2.2 mi. W Hobart Mills, 3 nymphs from originating at inner bases of palpi, taper- Eutamias amoenus, date? (A.M. Barnes). ing anteriorly, with mildly rounded shoulders before reaching base of hypostome. Unfed female 2.7-2.8 mm long, with a broad, Ixodes (Pholeoixodes) holdenriedi Cooley almost straight posterior margin. Capitulum (Figs. 214-217; Map 16) lacking defined cornua. Porose areas large, almost confluent. Auriculae (Fig. 224) Ixodes holdenriedi Cooley, 1946, Pan-Pac. lateral, bulbous protuberances continuous Entomol. 22:103, fig. 1. (female). with narrow, dorsoventral ridge on each side. Palpi broad, short; segments 2 and 3 DIAGNOSIS. Female. Capitulum (Fig. fused. Hypostome (Fig. 224) short, spatu- 214) with very short, bluntly pointed late, with denticular formula 4/4 at corona, cornua. Porose areas large and bordering 3/3 in distal third, 2/2 in basal two- posterolateral margins of basis capituli. thirds. Scutum (Fig. 220) slightly longer Palpi long with a length-width ratio of than wide; widest at midlevel; scapulae about 3.6:l; outer margins straight; segment rudimentary; cervical grooves well defined, 1 with a small convex ventral plate (Fig. shallow, broad, extending to posterolateral THE TICKS OF CALIFORNIA 59 margins. Coxa I (Fig. 225) with ill- TU0LU"E CO.: N Yosemite National Park, 1 defined, rounded, internal spur (Fig. 225). female from G. tephrocotis, VII-1937 All coxae with external spurs (Fig. 225). (Herms). Trochanters 1-111 with small, acute spurs. Engorged specimens bilobed posteriorly. Male. Legs as in female. Posterior --Ixodes (Ixodes) jellisoni Cooley and Kohls margin of body more strikingly squared than (Figs. 228-231; Map 17) in female (Figs. 218, 219). Differs from closely related I. signatus in having seg- Ixodes jellisoni Cooley and Kohls, 1938, ment 4 of palpi arising subterminally on U.S. Pub. Health Serv. Rept. 53:1619, the ventral aspect of segment 3, and in an fig. 2 (female); Cooley and Kohls, 1945, untoothed, plate-like hypostome, pointed U.S. Pub. Health Serv., Natl. Inst. apically rather than bluntly tipped (Fig. Health Bull. 184:41, fig. 15 (male). 223). Capitulum lacking cornua and auriculae. Palpi with segments 1-3 fused (Fig. DIAGNOSIS. Female. Differs from 223). Coxae as in female, but coxa I and closely related L. pacificus in having I1 with short internal spurs. Ventrally, a hypostome narrowly rounded apically rather fringe of heavy spines bordering posterior than bluntly rounded and file 3 of denticles margins of adanal plates and anal plate, extending not more than half length of hypo- extending anterolaterally to spiracular stome (Fig. 231). Denticle formula 313 plates (Fig. 222). apically and 212 to base, instead of 414 Nymph. Cornua distinct. Auriculae distally, then 313 and 212 to base, as in (Fig. 226) lateral, triangular. Palpi (Fig. pacificus. Cornua short (Fig. 230) or 226) short, broad, with segments 2 and 3 absent, as opposed to absent in pacificus. fused; segment 1 lacking anterior and pos- Auriculae appearing only as faint lateral terior spurs. Hypostome (Fig. 226) with swellings (Fig. 231). Scutum usually longer denticles 212; shape as in female. Scutum than broad, less often subcircular (Fig. lacking scapulae and lateral carinae. Coxae 228). Genital aperture at level of coxae with external spurs, but only coxa I with 111, as opposed to between coxae IV in distinct, smaller, inner spur (Fig. 227). pacificus. Coxa I (Fig. 229) with internal Larva. Capitulum and hypostome similar spur long, slim, pointed; external spur to those of nymph. Scutum lacking scapulae almost rudimentary. Coxae 11, 111, and IV and lateral carinae. Coxae with external with external spur small. spurs, but internal spur present only on Male. Very poorly known, but separable coxa I. from A. pacificus by presence of a posterior spur on segment 1 of palpi. This yellowish-brown bird tick is Nymph and larva undescribed. recorded by Keirans and Clifford (1978) only from the Gray-crowned Rosy Finch, Leuco- I. jellisoni is a rather rarely encoun- sticte tephrocotis, the type host; the Cliff terea tick occurring on a variety of rodents Swallow, Petrochelidon pyrrhonota; and the in California. It is also recorded from Prairie Falcon, Falco mexicanus. Kohls Utah by Johnson (1966). (1947) found numerous specimens, including The 20 recorded California collections larvae, nymphs, and adult females, infesting of L. jellisoni were made in the central to nestling Cliff Swallows, but obtained males southern parts of the State, and most were only from crevices near nests. He specu- taken from October to January. Pocket mice lated that the males may be nonfeeding, as and kangaroo rats are the usual hosts. is the case with males of some other Ixodes species. CALIFORNIA RECORDS. ALAMEDA CO.: In addition to the high montane records Calaveras Dam, from Perognathus californi- from California, 1. howelli is known from -cus: 1 female, X-24-1932; 1 male, 1 female, Colorado, Montana, and Texas. X-11-1940, 2 females, XI-14-1940; 2 females, XI-12-1944 (all R. Holdenried); 1 female, CALIFORNIA RECORDS. MONO CO. : 1.2 mi. X-24-1942 (Longanecker); 2 females, XII-6- E, 0.7 mi. S White Mountain Peak, 1 nymph 1940, 2 females, XII-7-1940 (F.C. Evans). from Leucosticte tephrocotis, VII-13-1973 FRESNO CO.: Junction of Hwys. 180 and 145, (R. Johnson); 0.1 mi. E, 0.2 mi. N White ?adults from Dipodomys nitratoides, XII-26- Mountain Peak, 1 female from 4. tephrocotis, 1952 (R.A. Bodostian and G.F. Augustson). VI-27-1974; Ellery Lake, Tioga Pass Huy, 1 KERN CO.: Bakersfield, 2 females from E. female from k. tephrocotis, VII-14-1974. nitratoides, XII-3-1963 (D. Roberts); Lerdo 60 BULLETIN OF THE CALIFORNIA INSECT SURVEY

SOC. Wash. 24:201, pl. 11, figs. 7-12 (female, male); Hooker, Bishopp, and Wood, 1912, U.S. Dept. Agric., Bur. Entomol. Bull. 106:82, pl. VI, figs. 1-3 (nymph, larva).

DIAGNOSIS. Female. Unfed (Fig. 2331, approximately size of I. pacificus, about 2 X 1.32 mm. Engorged, nearly circular, reaching 16 mm (Cooley and Kohls, 1945). Capitulum (Fig. 238) with distinct, rounded cornua; posterior margin of basis capituli almost straight; porose areas with deep pores; auriculae (Fig. 239) weakly defined, not apparent. Palpi (Fig. 238) short, broad, not more than 2.5 X longer than broad; segment 1 with flattened ventral plate (Fig. 239). Hypostome (Fig. 239) rounded apically, with nearly parallel sides, subtended basally by short rounded shoulders; denticles coarse, 2/2. Scutum (Fig. 235) longer than broad; surface punctate, with punctations most obvious mesad of lateral carinae; cervical grooves not apparent. Coxae all with short external spurs; coxa I (Fig. 239) with moderately MAP 17. Distribution in California of: long, slim, internal spur, blunt apically. Ixodes jellisoni Cooley and Kohls, solid Male (Fig. 232). Body size approxi- circles; I. kingi Bishopp, half-circles. mately 2.5 X 1.7 mm. Capitulum (Fig. 236) with short, distinct cornua and almost straight posterior edge dorsally; auriculae (Fig. 237) mild, lateroventral saliences. Grid, 2 females from E. nitratoides, I-15- Hypostome (Pig. 237) rounded, or bilobed 1965 (W. Mulback). KINGS CO.: W Antelope, apically; denticles coarse, 3/3 apically, 1 female from Ground Squirrel, IV-2-1932 then 2/2 to base; median and lateral denti- (RML P8037, type). LOS ANGELES CO.: Lake cles discrete, similar in size and shape. Hughes, 1 female from unrecorded host, Ventrally with shoulders subtending base of 11-24-1947. MONTEREY CO.: Henrv Sands hypostome (Fig. 237). Palpi short, broad; Canyon, 3 females from Dipodomys elephanti- segment 1 with flattened ventral plate as E, 111-15-1956 (K. Murray); Hastings in female. Scutum (Fig. 234) with lateral Reservation: 1 female from E. californicus, carinae evident and punctations most promi- IV-12-1940 (J.M. Linsdale); 1 female from nent along inner aspects of carinae. Coxae Thomomys bottae, XI-26-1941; 1 female each as in female. Genital aperture between from Perognathus sp., XI-22-1950 and XI-28- coxae 11. 1950. RIVERSIDE CO.: Temecula, 1 female Nymph. Basis capituli and scuta1 shape from Dipodomys agilis, XI-11-1966 (R.M. (Fig. 240) similar to female, except that Davis). SAN BERNARDINO CO.: Loma Linda, 1 auriculae present as ventrolateral saliences female from E. agilis agilis, XII-19-1955 (Fig. 241) and segment 1 of palpus with a (R.D. Lee). SAN MATE0 CO.: San Hateo, 1 ventral spur or horn, extending postero- female from cat, XI-29-1949 (R. Maynard). laterally. Hypostome (Fig. 241) rounded apically; denticular formula 2/2. Coxae as in female, but inner spur of coxa I rela- Ixodes (Pholeoixodes) kingi Bishopp tively short (Fig. 241). (Figs. 232-241; Map 17) Larva. Hypostome rounded; denticular formula 2/2. Palpi short, broad; segment 1 Ixodes pratti Banks, 1908, U.S. Dept. with posteroventral horn, or spur but no Agric., Bur. Entomol. Tech. Ser. 15:27, anterior horn. pl. IV, figs. 1, 4, 5 (pro parte, misid.). The female of L. Mngi resembles L. Ixodes kingi Bishopp, 1911, Proc. Biol. sculptus but is more massive and has a much THE TICKS OF CALIFORNIA 61

stouter hypostome. I. texanus females share separate. Hypostome (Fig. 247) large, many characteristics-with L. kingi, but the mildly notched apically, with moderate-sized former lack an internal spur on coxa I, a basal denticles; lateral denticles large, cornua, and heavy scuta1 punctations, as pointed; median denticles small, blunt well as differing in other features. crenulations. Scutum oval, convex; pseudo- I. kingi is widespread in the United scutum discernible primarily by darker color States, occurring from coast to coast and rather than by surface structure. Ventral from Canada to Mexico. West of the Rocky median plate 3X as long as anal plate. Hountains, collections are mainly recorded Coxae essentially as in female. from pocket gophers, kangaroo rats, and mice Nymph. Keirans and Clifford (personal (Greg son 197 1) . communication) consider the nymph of this species undescribed. In their judgment, CALIFORNIA RECORDS. INYO CO.-SAN the nymph described as 2. neotomae by Cooley BERNARDINO CO.: Death Valley, 1 female in and Kohls (1945, p. 36) "is not neotomae Nuttall tick collection, identified by J.E. but is apparently based on a slidelnounted Keirans (see fig. 168 in Nuttall and very small female of some Ixodes sp." Warburton (lgll), p. 174). KERN CO.: Walker Larva undescribed. Pass, Thomomys, 1-1-1891, 1 female (A.K. Fisher). MONTEREY CO.: Bastings Reserva- I. neotomae closely resembles 1. spini- tion: 5 larvae from Neotoma fuscipes, palpis and to a somewhat lesser degree 1. 11-14-1946; 1 larva from E. fuscipes, peromysci. IV-4-1946. I. neotomae has been collected most frequently from woodrats and rabbits (Table 14). Since the closely related 1. spini- --Ixodes (Ixodes) neotomae Cooley palpis also occurs commonly on rabbits as (Figs. 242-249; Map 18) well as rodents, the differentiation of these two species may be unusually dif- Ixodes neotomae Cooley, 1944, Pan-Pacif. f icult. Entomol. 20:7, fig. 1, 2 (male, female).

DIAGNOSIS. Female (Figs. 244, 245). Unfed, relatively small, about 1.5 X 1.02 mm. Capitulum (Fig. 248) with distinct, short cornua and relatively small, well separated porose areas. Auriculae (Fig. 249) long, retrograde, acutely pointed horns directed posteromedially. Palpi over 3.5 X longer than wide, with lateral profile lines straight; segment 1 (Fig. 249) ventrally with posterolaterally directed sharp spur. Hypostome (Fig. 249) subtended basally by a round-shouldered anterior projection of basis capituli; narrowly rounded apically; denticles 4/4, 3/3, then 2/2 at base. Scutum (Fig. 244) length about iths total 5 length of body in unfed female, very broadly rounded behind. Coxa I with short external spur, much longer internal spur. Other coxae lacking internal spurs, with progressively shorter external spurs on 11, 111, and IV. Genital aperture at level of poa- terior margins of coxae 111. Male (Figs. 242, 243). Small, about 1.0 X 0.7 mm. Capitulum (Fig. 246) with short, distinct cornua joined by straight posterior margin of basis capituli. Auri- culae (Fig. 247) short, pointed, or rounded MAP 18. Distribution in California of: lateral protuberances at same level as Ixodes neotomae Cooley, solid circles; L. ventral surface of basis capituli. Palpi ochotonae Gregson, half-circles; 1. short, broad, with segments 1-3 distinctly peromysci Augustson, rectangles. 62 BULLETIN OF THE CALIFORNIA INSECT SURVEY

With the exception of a collection external spur. Superficially, some speci- reported from New Mexico by Keirans and mens appear to lack coxal spurs. Clifford (19781, the recorded distribution Male. Body approximately 2.2 X 1.5 mm. of 3. neotomae is restricted to California, Basis capituli lacking cornua and auriculae. primarily from coastal counties from north- Palpi short, broad. Hypostome short, broad, ern to southern ends of the State. Collec- rounded distally; denticles irregular, tions are recorded for every month except transverse, small crenulations. Coxae as May. We have not examined the specimens in female. Genital aperture between coxae recorded here as nymphs of A. neotomae, and 111. we consider such records of dubious vali- Nymph. Anterior spur absent on segment dity. 1 of palpi, posterior spur present, tapering to point, directed posterolaterally (Fig. 257). Palpi short, thick. Cornua small. Ixodes (Pholeoixodes) ochotonae Gregson Posterior margin basis capituli slightly (Figs. 250-257; Map 18) sinuous (Fig. 256). Auriculae faint. Hypo- stome rounded apically, dentition 2/2 (Fig. Ixodes ochotonae Gregson, 1941, Canad. 257). Scutum about as long as wide, but Entomol. 73:224, pl. XVI, figs. 1-11 posterolateral margins converging sharply (male, female, nymph, larva). before terminating in rounded posterior extremity; lateral carinae distinct (Pig. DIAGNOSIS. Female. Unfed, approxi- 255); coxae similar to those of female. mately 2.2 X 1.4 nrm, approximately same size Larva. Palpal article 1 with anterior as I. pacificus. Capitulum (Pig. 252) spur sharp, equiangular; posterior projec- without cornua; porose areas large, well tion rounded. Cornua projecting laterally. separated; auriculae (Fig. 253) essentially Auriculae absent. Hypostome denticles 2/2. absent; palpi long, over 3.5 X longer than Internal spur of coxa I broadly angulate. wide; distal fifth of hypostome tapered; pointed apically, denticles 3/3 or 3/2 The female of 3. ochotonae resembles apically, then 2/2 to base (Fig. 254). closely I. angustus and I. holdenriedi. I. Scutum (Fig. 250) length-width ratio vari- angustus differs in having a hypostome wiTh able, but always longer than broad. All denticles 3/3 to the base. In I. holden- coxae with small external spurs; coxa I riedi, the hypostome tapers in its apical (Fig. 251) with internal spur subequal to half, terminating in a point. The male of

TABLE 14. California host records of Ixodes neotomae Cooley

Host or Source Tick Collections

~~ ~~ ~~ ~ ~ Adults NJrmphs Larvae Sylvilagus audubonii 1 - - Sylvilagus bachmani 3 - - Sylvilagus sp. 2 - - Lepus californicus 2 - - "Jack rabbit" 2 - - "Rabbi t 7 1 1 "Deer mouse'' 1 - - Neotoma fuscipes 10 4 - Neotoma lepida 3 - - Neotoma sp. 2 1 - Urocyon cinereoargenteus 1 - - Unrecorded 3 1 - THE TICKS OF CALIFORNIA ochotonae resembles angustus, but the latter has stronger blunt denticles on the hypostome arranged in 5 transverse rows. The nymph is separable from angustus, which has both anterior and posterior spurs on segment 1 of the palpi. I. ochotonae is known primarily from pikas and woodrats. Its distribution includes most of the western United States and British Columbia, Canada.

CALIFORNIA RECORDS. INYO CO. : 3 mi. S and 5 mi. W Big Pine Creek, 1 female from Ochotona princeps, VI-7-1942 (D. F. Hoff- meister). MENDOCINO CO. : Hopland Field Station, near Hopland, 2 nymphs from Urocyon cinereoargenteus, IV-21-1975 (R.S. Lane). ORANGE CO.: 5.8 mi. E Trabucco, 1 nymph from Neotoma fuscipes, XII-20-1965 (J.S. Fowler). PLUMAS CO.: Plumas Eureka State Park. 1 female from Eutamias townsendii, V-6-i981 (T. Schwan) ; specific localities not recorded: 2 larvae, 2 nymphs from N. fuscipes, 111-24-1949 (E.W. Jameson, Jr.7; 16 nymphs from Neotoma sp. 111-7-1949; 1 nymph from Neotoma sp., 111-6-1949. MAP 19. Distribution in California of Ixodes pacificus Cooley and Kohls, solid --Ixodes (Ixodes) pacificus Cooley and Kohls circles. Western Black-legged Tick (Figs. 258-267; Map 19) ridges (Fig. 261); ventrally, basis capituli Ixodes californicus Banks, 1908 (nec Banks, with salient posterior point at midline. 19041, U.S. Dept. Agric., Bur. Entomol. Palpi short, broad; segment 1 with small Tech. Ser. 15:24, pl. 2, fig. 12. ventral tooth. Hypostome (Fig. 261) promi- Ixodes pacificus Cooley and Kohls, 1943, nent, large, with large, laterally project- Pan-Pacif. Entomol., 19:140 (female, ing lateral denticles, particularly near male, nymph); Allred, Beck, and White, base; small median denticles as crenulated 1960, Brigham Young Univ. Sci. Bull., diagonal lines. Coxae essentially as in Biol. Ser. 1:16 (larva). female. Nymph. With definite cornua (Fig. 266). DIAGNOSIS. Female. Unfed, small, red- Auriculae (Fig. 267) small, lateral, shelf- brown with black legs (Figs. 258 to 259). like projections or bluntly triangular, not Body length about 2.6 mm. Hypostome arising greatly different from more acutely tri- from a smoothly tapered, median projection angular auriculae of I. spinipalpis. Dorsal of basis capituli; hypostome broadly rounded scutum (Fig. 264) shape variable, from sub- apically; denticles typically 4/4 apically, circular to bluntly angulate at postero- 313 to level proximal to midlength of hypo- lateral fourth with almost straight converg- stome, then 212 to base (Pig. 263 ). Cornua ing margins before curving again to form absent (Fig. 262) and auriculae not appar- rounded posterior margin. Eypostome (Fig. ent, or present as small ridges (Fig. 263). 267) narrowly rounded apically; denticles Palpal length-width ratio about 3.5:l. 3/3, then 2/2 to base. Palpi long; segment Palpi and trochanters of legs lacking spurs. 2 longer than 3. Palpi and trochanters of Coxae each with a small external spur. legs lacking spurs, segment 1 of palpi with Coxa I with a strong, acuminate inner spur a faint ventral plate. Coxae basically as extending over coxa I1 (Fig. 263). Genital in adult female, but internal spur of coxa opening near level of coxae IV (Fig. 258). I weaker (Fig. 267). Male (Fig. 258). Small, about 2.2 mm Larva. Spurs absent on palpi article long; dark brownish black. Cornua absent 1. Hypostome widest at midlevel, and nar- (Fig. 260), but auriculae present as lateral rowly tapered, rounded apically; denticles 64 BULLETIN OF THE CALIFORNIA INSECT SURVEY

212 to base except for short distal 313, poor yield of adults reported by the authors with third file composed of 3-4 denticles. indicates that a vital factor, nutritional, Basis capituli with angular auriculae form- environmental, or innnunological, was inimi- ing ventrolateral margins of basis. Five cal to normal development. pairs of scuta1 setae; 4 pairs of central dorsal setae; central dorsal seta 1 50-65 I. pacificus is known from California, um long. Seven pairs coarse marginal dorsal Oregon, Washington, Idaho, Nevada, and Utah, setae, bluntly tipped and minutely f imbri- and in British Columbia. In California it ated, 1.5-2 X as long as pair of supple- is recorded from 50 counties at elevations mentary setae; 4 pairs of marginal ventral ranging from sea level to over 2,150 m. As setae. Coxa I with prominent acute inner might be expected for a species in which spur; coxae I and I1 with small external the adults are so vulnerable to desiccation, spurs. most collections have been made in coastal areas. Adults of I. pacificus are very similar Among 3,000 specimens of adult 2. &- to the relaGvely rare I. jellisoni. -ficus collected in California, the majority Females of the latter differ Tram pacificus were taken from November through May. in having the hypostome vefY narrowly Larvae and nymphs were most commonly found rounded apically; the basis capituli has no from March through June, but they, as well visible transverse sutural line, and the as the adult, have been collected in every tarsi are shorter (tarsi I 0.54 mm long, season. The irnmature stages occur comonly compared to 0.84 mm in pacificus); and the on alligator lizards (Gerrhonotus spp.), genital aperture is at the level of coxae fence lizards (Sceloporus spp.), and ground- 111. The male of jellisoni has a distinct inhabiting birds, and less frequently on a posterior spur on segment 1 of the palpi variety of small rodents and lagomorphs. instead of the mild ventral tooth seen in In the most thorough study of its kind ever pacif icus. reported for deer, Westrom (1975) reported This is the most commonly encountered collecting larvae and nymphs most abundantly Ixodes species in California, and the most in the spring, but in lesser numbers at all important economically and to human health. seasons, from 23 of 71 deer examined in Adult ticks feeding on man may produce pain- Mendocino County, indicating that large ful sores which are slow to heal. Occasion- mammals may play a more important role as ally their bites are followed by sensory hosts of the immature stages than is disturbances near the affected area. Nymphs generally realized. probably attack man more frequently in Arthur and Snow (1968) implicated 55 California than indicated by the record. species of vertebrates as hosts for 2. Three collections of nymphs embedded in the pacificus, and 52 species are recorded here skin of man were made in 1981, primarily from California (Table 15) , including because of an increased awareness of the several not previously listed. Dogs are problem. Domestic livestock and wild game common hosts of the adult in coastal animals are commonly bitten. Nelson (1973) regions. On deer, Westrom (1975) reported has reported tick paralysis caused by I. it as second only to Dermacentor occiden- pacificus in a dog in California, and Hughes talis as an acarine parasite in Mendocino et al. (1976) isolated a spotted-fever-group County, finding it on 66% of the 71 deer rickettsia from the tick in Oregon. Lane examined. Man and horses are frequently et al. (1981) cite evidence that A. pacifi- bitten by adult females. We identified 2. -cus in Mendocino County is associated with pacificus from 11 of 167 bears (Ursus a rickettsia of the spotted fever group. americanus) examined by D. Graeber from 1974 Epidemiological evidence points to the to 1978 for ectoparasites in Yosemite implication of I. pacificus in the transmis- National Park, and from 28 of 73 bears sion of Lyme disease of man in northern examined by Graeber in Sequoia National Park California and Oregon (Steere and Malawista, from May to December 1980. 1979; Naversen and Gardner, 1978). Arthur and Snow (1968) reared I. &- ficus in the laboratory using guinea pigs --Ixodes (Ixodes) peromysci Augustson as hosts. They found that a relative humi- (Figs. 268-279; Map 18) dity of approximately 90% was essential to survival of all stages. Under the culture Ixodes peromysci Augustson, 1939, Bull. S. conditions reported, the life cycle requires Calif. Acad. Sci. 38:142, pl. 30, figs. less than one year, which reflects the con- 1-5; pl. 31, figs. 1-9 (female, male, dition occurring naturally, although the nymph, larva). THE TICKS OF CALIFORNIA 65

DIAGNOSIS. Female (Fig. 269). One of separable by their somewhat larger size, our smallest ticks, body about 1.3 X 0.8 mm hypostome dentition greater than 313, and in unfed state. Basis capituli (Pig. 271) horn-like retrograde auriculae. Hales of with faint, well separated porose areas; peromysci resemble closely neotomae and cornua distinct, posteriorly directed, spinipalpis, but the internal spur of coxa pointed projections; posterior dorsal margin I is not acuminate, and the ventral median straight, salient; auriculae prominent, plate is less than 3X longer than the anal posteriorly directed stout spurs (Fig. 273); plate. The nymph of peromysci is distin- palpi over 3X longer than wide; basal palpal guished from species of similar appearance segment with prominent posteroventral spur such as 2. pacificus and L. spinipalpis by (Fig. 273); hypostome long, narrow, narrowly its small size. rounded terminally, with denticles 313 in I. peromysci is known only from Cali- apical half, then 212 to base (Fig. 273). fornia. All the collections confirmed by Scutum longer than wide (Fig. 271). Legs us have been from the Channel Islands, where of moderate length. Coxae with external the primary host appears to be Peromyscus spurs short but distinct; internal spur only maniculatus . on coxa I, short. Genital aperture between coxae IV. CALIFORNIA RECORDS. SANTA BARBARA CO. : Male (Fig. 268). Relatively rarely Santa Barbara Island: 11 males, 16 females, recorded. Very small, with body, or idio- 6 nymphs, 1 larva from Peromyscus manicu- soma, 0.96-1.1 mm long. Capitulum (Fig. latus, V-(28-30)-1939 (Augustson); 3 females 270) with well developed cornua, pointed from 1. maniculatus, VI-12-1978 (Storrer and directed posteriorly. Auriculae mild and Trager); 4 females and 1 male from 2. saliences (Fig. 272). Palpi short and maniculatus, 111-(8,9)-1979; 1 female from broad. Hypostome (Fig. 272) broadly and -P. maniculatus, 111-8-1979 (P.W. Collins) ; deeply indented apically, with lateral 1 female, 1 male from E. maniculatus III-8- large, sharp denticles, in contrast to more 1979 (J. Storrer); 1 nymph from E. manicu- median diagonal rows of crenulations; basal latus, 111-6-1979 (J. Storrer). VENTURA denticles large, relatively blunt, and CO.: Anacapa Island: 1 female from 2. s- directed posteriorly. Scutum (Fig. 270) culatus, X-3-1978 (J.M. Greaves); 3 collec- with distinct punctations, more prominent tions of 1-2 nymphs each fromz. maniculatus in area bordering and posterior to pseudo- anacapae, X-(1-3)-1978; 5 collections of scutum. Ventral plates (Fig. 275) with 1-13 nymphs each from Rattus rattus, distinct punctations. Median ventral plate X-(1-3)-1978; 1 nymph from Gerrhonotus about 2.5 X longer than anal plate. All multicarinatus, X-1-1978 (D.L. Johnson and coxae with distinct, short external spurs; S .E. Miller). coxa I with longer, bluntly pointed internal In addition to the above records, the spur (Fig. 274). Rocky Mountain Laboratory of the National Nymph (Fig. 279). Body length from tip Institute of Allergy and Infectious Diseases of scapulae 0.65-0.70 mm. Capitulum not lists 1 nymph of 1. peromysci from Neotoma subtriangular dorsally (Fig. 276). Cornua fuscipes collected IV-4-1933 at Pasadena, and posterior margin of capitulum as in Los Angeles County. J.E. Keirans (personal female. Auriculae bluntly triangular communication) states that the specimen is lateral extensions (Fig. 277). Capitulum misidentified, and he believes it may 0.25 X 0.16 mm. Hypostome similar to represent the true nymph of 1. neotomae. female, but 0.144 mm long and denticles 313 only near apex, then 2/2 to base. Palpi long, slim, with slightly concave outer Ixodes (Pholeoixodes) rugosus Bishopp margins; segment 1 with small ventral pos- (Figs. 280-291; Elap 20) terior projection (Fig. 277). Scutum subcircular, about 0.42 mm long (Fig. 278). --Ixodes cookei var. rugosus Bishopp, 1911, Coxal spurs similar to those of female. Proc. Biol. SO~.Wash. 24:197. --pl. 11. figs. Spiracular plate oval, 0.11 X 0.07 mm. I 1.-. 2 (female. male). Larva. Subcircular, about 0.5 X 0.45 Ixodes rugosus; Cooley and Kohls, 1945, mm in the unfed state. Scutum as broad as U.S. Pub. Health Serv., Natl. Inst. long, with acute lateral angles. Cornua, Health Bull. 184:122, fig. 44H-K auriculae, and coxae 1-111 similar to those (nymph) of nymph. DIAGNOSIS. Female. Unfed, oval-shaped, Females of I. neotomae and 1. spini- almost 3 mm long, red-brown. Basis capi- palpis, which resemble 1. peromysci, are tuli (Pig. 284) with porose areas large, 66 BULLETIN OF THE CALIFORNIA INSECT SURVEY

TABLE 15. California Records of Ixodes pacificus Cooley and Kohls

Host or Source Tick Collections Nymphs Unrecorded Instars Sceloporus 2. occidentalis 14 - Sceloporus graciosus 3 - Sceloporus sp. 6 - Cnemidophorus sp . 1 Gerrhonotus coeruleus 4 Gerrhonotus E. multicarinatus 8 Gerrhonotus E. scincicauda 2 Gerrhonotus sp. 8 "Lizard" 3 Lophortyx californica 1 Aphelocoma californica 1 Colaptes cafer 1 Cyanocitta stelleri 1 Troglodytes troglodytes 2 Thryomanes bewickii - Hy loc ichla ustulata 3 Wilsonia pusilla 1 Aimophila ruficeps - Aimophila sp. - "Sparrow" 6 Junco oreganus 4 Zonotrichia leucophrys nuttalli 1 Zonotrichia atricapilla 1 Passerella ileaca 1 Melospiza melodia 1 Didelphis marsupialis virginiana 1 Scapanus latimanus - Sorex sp. - -Homo sapiens a "Jackrabbit" - "Cottontail rabbit" - Spermophilus beecheyi 8 Spermophilus sp. 1 Sciurus griseus 1 Perognathus californicus 2 THE TICKS OF CALIFORNIA 67

TABLE 15 continued

~~~ Host or Source Tick Collections

Adults Nymphs Unrecorded Instars Dip od omy s dese rt i 1 - Dipodomys sp. 1 Reithrodontomys megalotis - Peromyscus boylii - Peromyscus maniculatus - Peromyscus truei - Peromyscus sp. - Neotoma fuscipes - Neotoma lepida 1 Microtus californicus - Microtus sp. - Canis familiaris 80 Canis latrans 10 Urocyon cinereoargenteus 5 Urocyon littoralis littoralis 1 "FOX" 3 Ursus americanus 40 Procyon lotor - Mustela frenata 1 Taxidia taxus 1 --Felis catus 5 Felis concolor 7 --Felis rufus 20 Equus caballus 42 "Mu 1eo 1 --Sus scrofa 1 Odocoileus h. columbianus 45 "De er" 16 Cervus nannodes 1 --Bos taurus 12 Vegetation (tick drag) a8 House 4 Unrecorded 108 68 BULLETIN OF THE CAL,IFORNIA INSECT SURVEY

- f nearly parallel lateral margins (Fig. 289). Cornua short. Auriculae curved lateral saliences. Palpi short, broad; segment 1 ventrally with prominent, rounded posterior projection. Hypostome rounded apically; dentition 2/2 except for additional minute denticles on corona (Fig. 290). Scutum (Fig. 289) length and width subequal; lateral carinae distinct, bowed outward at midlevel. Internal spur of coxa I short, rounded (Fig. 291); external spurs of coxae present, small. Larva. Larvae have been recorded but are undescribed.

Adults and nymphs of r. rugosus superficially resemble L. pacificus but are more closely related to I. sculptus, as discussed under that species. I. rugosus is known primarily from skunxs, foxes, weasels, and dogs along the Pacific coast of Washington, Oregon, and California. Gregson (1956) lists 6 collection records from British Columbia from coyotes and skunks.

MAP 20. Distribution in California of: California collections of I. rugosus Ixodes rugosus Bishopp, half-circles; A. extend from Humboldt County to gs Angeles sculptus Neumann, solid circles. County, restricted to or west of the coastal mountain range. All instars have been collected most frequently from the spotted depressed, subtending posterior margin of skunk, Spilogale gracilis, although nymphs the basis. Cornua and auriculae not appar- and adults have been found on various other ent or absent. Palpi length-vidth ratio carnivores (Table 16). The adults and 2.5:l. Hypostome (Fig. 285) short (0.45 nymphs have been collected in California mm), spatulate, rounded apically, arising during every season of the year. from a broad-shouldered base; denticles 212, except 3/3 on corona. Scutum longer than broad, widest anteriorly, usually with Ixodes (Pholeoixodes) sculptus Neumann anterolateral rugosities and large, numerous (Figs. 292-301; Map 20) punctations (Pig. 287). External spurs of all coxae small. Internal spur of coxa I Ixodes sculptus Neumann, 1904, Arch. Para- moderately long, thin, and pointed apically sitologie 8:462 (female); Hixson, 1932, (Fig. 288). Genital aperture between coxae Iowa State Coll. J. Sci. 7:36, figs. 2, 111. 3 (male, nymph, larva). Male- Subelliptical (Pig- 280), similar Ixodes aequalis Wherry and Wellman, 1909, in size to unfed female. Basis capituli Entomol. News 20:376 (nomen nudum). (Fig. 282) with posterior margin straight Ixodes aequalis Banks, 1910, Proc. Entomol. to slightly concave. Cornua absent. Auri- SOC. Wash. 12:8. culae curved lateral saliences. Palpal segment 1 with salient posterolateral margin DIAGNOSIS. Female (Fig. 296). Unfed, almost tooth-like in appearance. Hypostome about 2.2-2.6 X 1.4-1.8 mm. Engorged female (Fig. 283) with 3-4 irregularly paired rows 6.3 X 4.5 mm (Cooley and Kohls, 1945). of bluntly pointed, overlapping denticles; Capitulum (Fig. 298) with conspicuous cornua lateral and median denticles of approxi- tending to converge posteriorly and elevated mately same shape and size. Scutum evenly over salient and sinuous posterior margin. rounded at both ends; surface with numerous Auriculae essentially absent. Palpi of shallow punctations (Fig. 286). Coxae as moderate length; length-width ratio not over in female. 3:l. Palpal segment 1 (Pig. 299) with Nymph. Basis capituli broad with short, ventral plate angulate posteriorly. Hypo- THE TICKS OF CALIFORNIA 69

TABLE 16. California Records of Ixodes rugosus Bishopp

~ ~~ ~ Host or Source Tick Collections

Adults Didelphis virginiana 1 Homo sapiens 1 Canis familiaris 3 Urocyon cinereoargenteus 2 Urocyon littoralis santacruzae 1 Procyon lotor 1 Mustela frenata xanthogenys 3 "Weasel" 1 Spilogale gracilis 10 Mephitis mephitis 3 "Skunk" 1 --Felis rufus 1 Unrecorded 1

stome (Fig. 299) bluntly rounded, with sides apically, then 2/2 to base. Coxae as in slightly convex, arising from a tapered female. medial projection of basis with mere sug- Larva. Segment 1 of palpi with both gestions of shoulders; dentition 313 sub- anterior and posterior ventral spurs promi- apically, then 212 to base. Scutum suboval, nent and of subequal size. Lateral margins deeply punctate, with cervical grooves and of basis capituli subparallel; 2 pairs lateral carinae usually distinct. Coxae central dorsal setae. each with short external spur; coxa I (Fig. 297) with long pencil-shaped internal spur. The female I. sculptus resembles I. Genital aperture between coxae 111. rugosus, but it :as a much more prominent Male (Fig. 292). Body oval, 2.5 X 1.7 cornua and lacks anterolateral scuta1 mm. Basis capituli (Fig. 294) with straight wrinkles. It also resembles r. kingi, but to concave posterodorsal margin and cornua is less massive and the posterior margin of small to absent. Auriculae weak lateral the basis capituli is sinuous. The male of ridges. Palpi short, broad; segment 1 with sculptus is similar to rugosus, but the small ventral plate. Hypostome (Fig. 295) hypostome of the latter has more pointed short, with apex rounded or notched; denti- denticles which are overlapping rather than cles short, rounded, widely spaced teeth rounded and widely spaced. The nymph of 2. (almost crenulations), approximately sub- sculptus is separable from I. angustus and equal in size laterally and medially. -I. soricis in that the internal spur of Scutum with numerous large punctations, coxa I is considerably longer than the particularly laterally. Coxae as in female. external spur. Nymphs of rugosus and kingi Genital aperture (Fig. 293) between coxae lack the anterior spur of palpal segment 1 11. seen on sculptus, but in some specimens of Nymph. Capitulum (Fig. 300) with large, the latter the anterior spur is rounded and pointed cornua diverging posteriorly; pos- similar in appearance to rugosus. terior margin slightly sinuous; auriculae I. sculptus is widely distributed in large, rounded saliences. Palpal segment 1 the central and western United States and ventrally with prominent anterior and pos- Canada, where it occurs commonly on burrow- terior projections (Fig. 301). Hypostome ing rodents and in their nests. (Fig. 301) rounded apically; denticles 3/3 70 BULLETIN OF THE CALIFORNIA INSECT SURVEY

In California, I. sculptus is rather Ixodes eudyptidis var. signata Neumann, generally distributed in the coastal range 1904, Arch. Parasitologie 8:451. of the central part of the state, and in Ceratixodes signatus; Banks, 1908, , U.S. the Sierra Nevada from Kern County to Lassen Dept. Agric., Bur. Entomol. Tech. Ser. County. Collections are primarily from l5:21. ground squirrels, Spermophilus spp., and their burrows (Table 17). Adults as well DIAGNOSIS. Female. Spider-like appear- as nymphs may be found throughout the year, ance (Fig. 303), with long, thin legs. Body but collections of larvae have been recorded elongate, engorged size 8 X 3.5 mm. Capi- only during the summer. tulum (Fig. 306) broad, without cornua, although posterolateral corners salient. Porose areas extensive and almost confluent Ixodes ( ScapNxodes) signatus Birula on median line. Evenly curved lateral (Figs. 302-309, Map 21) extensions of basis capituli form auriculae (Fig. 307). Hypostome (Fig. 307) broadly Ixodes signatus Birula, 1895, Izv. Imp. rounded apically; dentition 3/3 distally, Akad. Nauk, ser. 5, 2:357, table 1, then 212 to base; not arising from median figs. 10-13 (female); Nuttall and extension of the basis capituli. Scutum Warburton, 1911, Ticks: a monograph of (Fig. 303) oval but variable. Cervical the Ixodoidea 2:263, fig. 263 (nymph, grooves always distinct. Coxae with exter- larva); Gregson, 1954, Canad. Entomol. nal spurs, but lacking internal spurs (Fig. 86:275, fig. 1 (male). 308). Spiracular plate nearly circular; Ixodes arcticus Osborn, 1899, in Jordan et macula eccentric; goblets numerous and al., Acarina fur seals and fur-seal small. Genital aperture at level of coxae islands, North Pacific Ocean, pt. 3, 11. 9:553, fig. 2. Male (Fig. 302). An elongate narrow Ixodes parvirostris Neumann, 1901, M&n. SOC. bodied tick with elongate, oblong scutum Zool. Fr. 14:284. having nearly parallel lateral margins.

TABLE 17. Records of Ixodes sculptus Neumann, 1904

Host or Source Tick Collections

Adults Larvae Unrecorded Instars Lophortyx sp. 1 - - -Homo sapiens 1 "Jackrabbit" 1 Eutamias merriami 1 Marmota sp. - Spermophilus beecheyi 102 Spermophilus beecheyi douglasii 2 Spermophilus beldingi 6 Spermophilus lateralis 13 Spermophilus oregonus 2 Neotoma fuscipes 1 Martes americana 1 Mustela frenata 3 Vegetation 4 Unrecorded 10 THE TICKS OF CALIFORNIA 71

the Pigeon Guillemot is the only other host recorded here. All stages of the tick have been collected in every season.

CALIFORNIA RECORDS. DEL NORTE CO.: Castle Rock, Crescent City, 1 nymph and 1 larva from Phalacrocorax pelagicus, date? (H.S. Fullon). HUMBOLDT CO.: Flat Iron Rock, 10 females, 13 nymphs, 45 larvae from rocks, VI-22-1972 (a. Johnson). MARIN CO.: Fort Baker: 2 nymphs from 2. pelagicus, IV-24-1957 (H. Tingle); 1 female from "cormorant ,I' IV-15-1957 (Fontaine and M. Tingle); 1 female and 1 nymph from Phala- crocorax penicillatus, date? (Shepard); Stinson Beach, 1 female from "cormorant," IX-27-1939 (Hems); Tomales Bay, 8 females, 23 nymphs, 4 larvae from Phalacrocorax sp., XII-16-1920 (M.E. Davidson); Point Tomales, 10 males, 8 females, 23 nymphs, 1 larva from rocks, 11-26-1979 (G. Ulrich). MONTEREY CO.: Pacific Grove: 1 female from E. pela- &us, 1894 (Nuttall and Warburton); 3 females, 1 nymph, 1 larva from E. penicillatus, 1894, Nuttall collection, identification verified by 5.13. Keirans; 2 females MAP 21. Distribution in California of: from 2. pelagicus, XII-30-1905; unidentified Ixodes sipnatus Birula, half-circles; 2. location, 1 nymph from E. pelagicus, VIII-2- soricis Gregson, rectangles; _I. spinipalpis 1927, Bishopp collection. SAN FRANCISCO Hadwen and Nuttall, solid circles. CO.: Farallon Islands: 3 males, 28 females, 8 nymphs, 1 larva from abandoned nests of -P. pelagicus, V-6-1949 (D.G. Hanna); 24 Capitulum (Pig. 304) lacking cornua and females, 3 nymphs, 1 larva, XI-18-1949 (E.S. auriculae, palpi with numerous stout spines; Ross and D. Kelly); 1 larva fromg. penicil- segment 3 spatulate and reflexed, with inner -* latus VIII-9-1982 (R. Henderson). SANTA distal flange and two stout distal spines; BARBARA CO.: Prince Island, 9 females, 3 segment 4 in some specimens invisible in nymphs, 1 larva from Cepphus columba, dorsal view, due to subapical ventralinser- VII-23-1977 (Kelley and T. Schwan). SANTA tion. Hypostome reduced, truncate and con- CLARA CO.: Los Gatos, 1 female from cave mediodistally (Fig. 305); denticles "orchard," 111-13-1954 (L. Haitozo); San rudimentary. Coxae as in female. Jose, 1 nymph from "Cormorant," 111-12-1954 Nymph. Brownish-black nymphs elongated (Binnewies) . as in female and resembling it closely in other respects, including shape of capitulum (Fig. 309) and coxal armature (Fig. 309). Ixodes (Pholeoixodes) soricis Gregson Larva. Similar to female in shape of (Figs. 310-313; Map 21) capitulum, armature of legs, and shape of anal groove. Ixodes soricis Gregson, 1942, Canad. Ento- mol. 74:137, figs. 1-9 (female, nymph, I. signatus is a large tick which may larva); Gregson and Kohls, 1952, Canad. be distinguished from the closely related Entomol. 84:185, fig. 1 (male). bird tick, r. howelli, as described under the latter species, as well as by the fact DIAGNOSIS. Female (Fig. 310). Pale that the normal hosts of howelli are terres- yellowish white. Very small: unengorged trial rather than marine birds. body to tip of scapula 1.2 X 0.48 mm (Greg- I. signatus is known from California, son, 1942). Body with numerous appressed Oregon, Washington, Alaska, Canada, Japan, setae. Capitulum (Fig. 312) with cornua and eastern Russia. very small; auriculae not apparent. Palpi In California, r. signatus is known convexly curved on both inner and outer primarily from cormorants and their nests; margins (Fig. 313); less than 3X longer than 72 BULLETIN OF THE CALIFORNIA INSECT SURVEY broad. Hypostome (Fig. 313) pointed api- collections of 1 to numerous specimens are cally; sides evenly tapered to apex; denti- recorded from only 6 California counties, tion 313; denticles long, sharp. Scutum although the combined ranges of the recorded (Pig. 310) length-width ratio over 4:3; hosts cover all but the southeastern region scapulae short; lateral carinae distinct, of California. long, straight; longitudinal furrows on posterior margin visible at magnification CALIFORNIA RECORDS. ALAMEDA CO. : Straw- of 1OOX. All coxae (Pig. 311) with external berry Canyon, 1 nymph from Scapanus w- spurs and barbed setae; internal spur on -’manus V-25-1955 (W.C. Russell). MARIN CO.: coxa I subequal to external spur. Genital Fort Baker: 1 female from Sorex sp., XI-15- opening (Pig. 311) at level of interface 1957 (M. Tingle); 7 nymphs from Sorex sp., between coxae I11 and IV. VI-29-1943 (R. Holdenried); Camp Mendell, 1 Male. Extremely small: 0.97 (to tip of female from Sorex trowbridgii, X-10-1942 scapula) X 0.64 mn (Gregson and Kohls, (R. Holdenried); Fort Baker, 1 female from 1952). Basis capituli without cornua or -Sorex vagrans, 1-9-1943 (Holdenried and auriculae; posterior dorsal margin straight Burroughs). MONTeREY 0.: Hastings Reser- to slightly convex. Palpi short, bulbous. vation, from Sorex ornatus: 1 female, V-2-39 Hypostome reduced, slightly notched api- (R. Holdenried); 2 females, XII-16-39 (J.A. cally, with 4 transverse rows of irregular Gray); 5 females and 2 larvae, 1-13-40; 2 crenulations. Scutum convex, narrower collections of 1 female each 11-11-40; 1 anteriorly than osteriorly. Ventrally female, XII-17-41; 1 female, 1-17-56. anal plate about frds as long as median PLUMAS 0.:Rock Creek Dam, 2 females, 4 3 plate. Genital opening at level of anterior nymphs, 5 larvae from 5. trowbridgii, V-30- margins of coxae 111. Coxae similar to 1950 (E.W. Jameson); E Rock Creek Dam, 2 those of female, but internal spur of coxa nymphs from 2. trowbridgii, VI-1-1950 (E.W. I slightly larger than external spur. Jameson); Quincy and unrecorded locations, Nymph. Basis capituli with very small 33 collections from 2. trowbridgii, 35 cornua and no auriculae. Hypostome apex females, 6 nymphs, 3 larvae, X-31-48 - rounded; dentition 212 until 111 near base. XI-2-50 (E.W. Jameson). SAN MATE0 CO.: no Segment 1 of palpi ventrally with prominent specific locality given, 3 larvae from 2. anterior and posterior spur-like projec- vagrans, 11-21-73 (R.B. Eads). SONOMA CO.: tions. Scuta1 length about 0.4 mn. Coxal no specific locality given, 1-6 specimens spurs similar to those of female. each from 3 Scapanus latimanus, 1976-77 Larva. Palpal segment 1 with ventral (cited by Yates et al., 1979). anterior and posterior spurlike projections as in nymph. Scutum 0.257 X 0.28 mm (Gregson, 1942). --Ixodes (Ixodes) spinipalpis Hadwen and Nuttall -I. soricis closely resembles I. angus (Figs. 314-319; Map 21) -’tus from which the female is distinguished by its smaller size and the presence of Ixodes dentatus vat. spinipalpis Hadwen and furrows on the posterior margin of the Nuttall, in Nuttall, 1916, Parasitology scutum; the male is separable from angustus 8:301, figs. 6, 7, 8 (female, nymph, by its smaller size and by its more reduced larva). hypostome with irregular crenulations. Ixodes diversifossus Bishopp, 1912, Proc. Nymphs of soricis are smaller than those of Biol. SOC. Wash. 25:30, pl. 1, figs. angustus, and the scutum lacks the lateral 7-12 (nec Neumann, 1899) (female, male). carinae of the latter. The only other male tick in California as small as r. soricis DIAGNOSIS. Female. Closely resembling is I.peromysci, a species easily separable erom sci but larger; unfed female 1.89 by such features as its strong cornua, lieX 1 w 26 mm Cooley and Kohls 1945). Capi- heavily toothed hypostome and insular dis- tulum (Pig. 315) with distinct cornua. tri bution. Auriculae (Fig. 317) posteriorly directed, I. soricis is primarily a parasite of retrograde horns. Segment 1 of palpi with shreGs in the western United States and short, broadly pointed ventral spur with Canada. In California it has been collected unequal sides (Fig. 317). Hypostome arising from 3 species of shrews, Sorex trowbridgii, from acuminate medial projection of basis -S. vagrans and 5. ornatus. The only other capituli; narrowly rounded apically, with host recorded in the state is the broad- pointed denticles larger laterally than footed mole, Scapanus latimanus. The 51 medially; 4/4, 3/3, then 2/2 to base (Fig. THE TICKS OF CAL,IFORNI.A 73

317). Scutum (Fig. 315) oval, widest behind gest a potential significance in the circu- midlevel, reaching more than half length of lation of a variety of microorganisms mong body in unfed female. All coxae with short its hosts, and it has been found naturally external spurs. Coxa I (Fig. 318) with infected with Powassan virus in South long, slim internal spur. Genital opening Dakota. It is distributed throughout the between coxae IV. western United States and Canada, occurring Male. Idiosoma 1.4-1.5 mm long. Capi- on lagomorphs and rodents. Immature ticks tulum (Fig. 314) with distinct cornua, pos- occur also on birds. teriorly converging lateral margins, and straight posterior margin between cornua. In California the 10 recorded species Hypostome large, slightly notched apically. of hosts include a variety of rodents and Lateral denticles large, sharp, but more lagomorphs as well as terrestrial birds median denticles in diagonal rows of pointed (Table 18). Adults have been taken in to rounded crenulations; basally with pair fall, winter, and spring, nymphs during all of large, relatively blunt, posteriorly seasons. The geographic distribution in directed teeth (Fig. 316). Scutum (Fig. California includes both the southern and 314) with pseudoscutum variably indicated northern coastal ranges as well as the by darker color and smaller punctations than central Sierra Nevada and the arid south- on remainder of shield. Coxae essentially eastern region. as in female- Ventral median plate 3X longer than anal plate. Nymph. Capitulum with distinct cornua Ixodes (Pholeoixodes) texanus Banks and short, bluntly triangular auriculae (Figs. 320-329; Map 22) (Fig. 319). Palpi long, slim, with outer margins concave. Hypostome (Fig. 319) Ixodes pratti Banks, 1908, U.S. Dep. Agric., narrowly rounded apically, widest at mid- Bur. Entomol. Tech. Ser. 15:27, pl. IV, level; denticles long, pointed, 313 in figs. 1, 4, 5 (pro parte, misid.). apical third, then 212 to base. Scutum Ixodes texanus Banks, 1909, Proc. Entomol. broadly oval, rounded caudally. Internal SOC. Wash. 10:172, figs. 16, 17 spur of coxa I about twice as long as exter- (female); Cooley and Kohls, 1945, U.S. nal spur. Pub. Health Serv., latl. Inst. Health Larva. Capitulum with only rudimentary Bull. 184:77, figs. 30, 31 (male, cornua. Auriculae present but reduced. nymph); Allred, Beck, and White, 1960, Palpi long, slim. Hypostome narrowly Brigham Young Univ. Sci. Bull. Biol. rounded apically, with denticles 313, then Ser. 1:22, figs. 28-30 (larva). 212 basally. Scutum as broad as long. Coxae I and I1 with spurs as in nymph, but DIAGNOSIS. Female. Sclerotized parts smaller. yellowish brown. Basis capituli (Fig. 326) with cornua indistinct to moderate and with Females of 1. peromysci have auriculae posterior margin slightly sinuous. Porose which are posteriorly directed spurs rather areas deep, irregular, well separated. than retrograde horns as inL. spinipalpis. Palpi short, wide, rugose; segment 1 with Females of 5. neotomae have the genital small ventral plate, somewhat angular pos- atrium at the level of coxae 111 or 111-IV terolaterally. Hypostome flanked basally rather than at the level of coxae IV. L. by angular humps (Fig. 3271, rounded api- neotomae females also have the spur of cally; dental formula 2/2 except apically, palpal segment 1 more acutely pointed and where smaller denticles may be 515. Lateral with equal sides. Xales of neotomae do not denticles coarse, pointed; median denticles have quite as large basal teeth on the hypo- smaller, more rounded. Auriculae absent. stome as in spinipalpis, nor is the pseudo- Scutum (Fig. 324) length and width subequal; scutum demarked by a zone of punctations. lateral areas wrinkled. Coxae (Fig. 325) The similar males of peromysci are much with external spurs vestigial to absent; smaller than the males of spinipalpis. coxa I with short, rounded inner spur. Nymphs of the very similar peromysci differ Genital aperture at level of posterior from spinipalpis in having the internal and margins of coxae I11 (Fig. 325). external spurs of coxae I1 subequal, as well Male. Basis capituli (Fig. 322) with as by the smaller measurements of peromysci. cornua indistinct to absent; posterior I. spinipalpis is of no demonstrated dorsal margin sinuous; auriculae absent. econ&inic or public health importance. Its Palpi short, wide, rugose; segment 1 as in wide host range and distribution muld sug- female. Hypostome (Fig. 323) flanked i 74 BULLETIN OF THE CALIFORNIA INSECT SURVEY

TABLE 18. California Records of Ixodes spinipalpis Radwen and Nuttall

Host or Source Tick Collections

Nymphs Larvae Pipilo fuscus 1 - Pipilo erythrophthalmus - 2 Zonotrichia atricapilla - 1 Sylvilagus bachmani 2 6 Sylvilagus sp. 1 1 Lepus californicus - - "Rabbit" 1 1 Perognathus formosus 2 - Peromyscus maniculatus 2 2 Peromyscus sp. 2 - Neotoma fuscipes 2 - Neotoma lepida 3 1 Neotma sp. 1 - Microtus californicus 2 1 Unrecorded 1 1

basally by angular humps as in female and I. texanus closely resembles L. hearlei, slightly notched apically; denticles faint, but ?he female of the latter is smaller, transverse rows of crenulations. Ventral less robust, and lacks the angular humps median and anal plates of subequal length. subtending the base of the hypostome. Simi- Genital aperture at about level of coxae larly, the male and nymphs of hearlei lack 11. Coxae (Fig. 321) with external spurs humps on either side of the hypostome. as low ridges. Coxa I with internal spur I. texanus is distributed throughout very short, rounded. Trochanters without the United States and Mexico. In Canada it spurs. is common in British Columbia but less Nymph. Basis capituli of California common in the east. In the west it is specimens lacking cornua, with posterior primarily found on mustelid hosts, but dorsal margin nearly straight (Fig. 328). raccoons are frequently infested in the Palpi short, thick, rugose; segment 1 east. ventrally with plate protruding posterolaterally as a rounded flange (Fig. 329). In California, recorded distribution of Hypostome (Fig. 329) broadly rounded; dental -I. texanus is primarily from interior moun- formula 2/2, except apical area with a third tainous areas. Specimens collected from file of about 3 teeth. Basally, hypostome pine martens near Lake Tahoe appear quite flanked by angular humps or shoulders (Fig. typical of 2. texanus, but larvae, nymphs 329) best seen on ventral side. Scutum and females collected in southern California broader than long and broader anterior to from skunks and the opposum are atypical midlevel; rugose anterolaterally. Coxal and are only provisionally placed in this spurs similar to those of female. species. Larva. No spurs on palpal segment 1. Hypostome somewhat oval in outline; denti- CALIFORNIA RECORDS. FRESNO CO. : King's cles 2/2 to base, plus 1-2 apical teeth in River, 1 female from gray squirrel, VII-1896 file 3. Coxae without external spurs; (V.L. Kellogg), (in Nuttall collection, internal spur of coxa I bluntly pointed. identification verified by J.E. Keirans). THE TICKS OF CALIFORNIA 75

I,.-- I Y Y Y ,. ". ,. .* ". Barrow Straite, 1850-1851, 2: appendix, ccx, fig. 10 (female); Neumann, 1899, Mdm. SOC. Zool. Fr. 12:126, 127 (male, nymph); Nuttall, 1912, Parasitology 5:60, fig. 9 (larva). Hyalomma Pickard-Cambridge, 1876, Proc. 2001. SOC. London 17:261, pl..XIX, fig. 3. Ixodes borealis Kramer and Neuman, 1883, in Nordenskib'ld, Vega Expeditiones Vetensk 3:527. -_~1. 43. Ixodes hirsutus Birula, 1895, Isv. Imp. Akad. Nauk, ser. 5, 2:356, pl. I, figs. 7-9. Ixodes putus procellariae Schulze, 1930, Sitzungsber. Abh. Naturforsch. Ges. Rostock 2:120-124.

DIAGNOSIS. Female (Figs. 332, 333). A large, long-legged tick. Body length, including capitulum, 3.3 mm, increasing to 11 X 7 mm in engorged state (Nuttall and Warburton, 1911). Dorsal and ventral body prominently clothed with long hairs. Capi- tulum (Fig. 336, 337) without cornua or auriculae. Porose areas large, distinct, MAP 22. Distribution in California of: separated by narrow median strip of cuti- Ixodes texanus Banks, solid circles; r. cula. Palpi with segment 4 relatively uriae White, rectangles; L. woodi Bishopp, elongate and visible from above; segment 1 ha1f-circ les . relatively well developed. Hypostome rounded apically; dentition 212; subequal size; teeth bluntly pointed. Scutum (Fig. 332) wider anteriorly, broadly rounded caudally. Cervical grooves broad, straight, NEVADA CO.: Sagehen Creek Field Station, prominent, diverging posteriorly. Coxae from Martes americana: 1 female, VII-18- without spurs. Genital aperture (Fig. 333) 1979; 1 nymph, VII-22-79; 2 nymphs and 1 at level of interval between coxae I1 and female, VII-29-1979; 1 nymph, VIII-4-1979; 111. 1 nymph, VIII-17-1979; 6 females, IX-19- Male (Figs. 330, 331). Large: length, 1979, 3 nymphs and 13 females, 11-1-1980; 1 exclusive of caudal spines, 3.16 mu, width female, 11-5-1980 (W. Zielinski). ORANGE 2.49 mm (Cooley and Kohls, 1945). Capitulum CO.: Dana Point: 1 female from Didelphis (Fig. 335) without cornua or auriculae. virginiana, X-29-1951; 2 females and 50 Palpi with segment 1 large and visible io larvae from Spilogale gracilis, X-22-1952; dorsal view; segment 4 arising on ventral San Juan Capistrano: 1 female and 3 nymphs surface -11 removed from apical point of from Mephitis mephitis, X-4-1951; 1 female, segment 3. Hypostome (Pig. 335) reduced, 9 nymphs, 40 larvae from 5. gracilis, X-18- broad, bilobed, with vestigial denticles. 1951. SAN BEWARDINO CO.: Seven Oaks, 1 Scutum (Fig. 330) punctate; cervical grooves female from gray squirrel, V-9-1936. SHASTA prominent, diverging posteriorly. Anal, CO.: Lassen Natl. Park, 2 females and lmale adanal, and epimeral plates bearing caudal from Eutamias s. frater, IX-20-1940 (T. fringe of prominent spines (Fig. 334). Aitken). TULARl3-CO.: Mt. Whitney, 2 females Coxae essentially as in female. Genital from unidentified host, VI-19-1915. aperture at level of posterior margins of coxae I. Nymph. Capitulum and hypostome (Fig. Ixodes (Ceratixodes) uriae White 338) similar to those of female. Scutum (Pigs. 330-339; Map 22) (Fig. 339) widest anteriorly; cervical. grooves prominent. Coxae without spurs. --Ixodes uriae White, 1852, in Sutherland, Larva. With only 1 pair of posthypo- Journal of a voyage in Baffin's Bay and stomal setae; 4 pairs of scuta1 setae. 76 BULLETIN OF THE CALIFORNIA INSECT SURVEY

I. uriae occurs on marine birds and in internal spur as long or longer than exter- theiy nests in both northern and southern nal spur. Genital aperture at level of hemispheres in boreal, subpolar, and polar coxae 111. regions. It has been recorded from Europe, Male. Small: body 2.22 mm long to tip Asia, Australia, South America, and Canada, of scapula, width 1.44 mm (Kohls, 1950). as well as from Alaska, Oregon, and Cali- Capitulum (Fig. 342) without cornua. Auri- f ornia. culae broad lateral saliences, Palpi short, broad; lacking ventral projections on seg- Eley (1977) recorded --I. uriae from 10 ment 1. Hypostome (Fig. 343) notched api- of 12 species of seabirds nesting on off- cally; denticles as discrete crenulations; shore rocks of Del Norte and Humboldt files in 4/4 arrangement with no overlapping counties. The birds found infested were of rows. Scutum with well-defined puncta- Oceanodroma furcata, the Fork-tailed Petrel; tions throughout; cervical grooves absent. -0. leucorhoa, Leach' s Petrel; Phalacrocorax Coxae (Fig. 341) similar to those of female, auritus, the Double-crested Cormorant; p. but inner spur of coxa I not as slim. Ven- penicillatus, Brandt's Cormorant; p. pela- tral median plate somewhat longer than anal gicus, the Pelagic Cormorant; Lams occidew plate. Genital aperture at level of inter- -'talis the Western Gull; Uria aalge, the val between coxae 11 and 111. Common Murre; Cepphus columba, the Pigeon Nymph. Capitulum with short cornua; Guillemot; Ptychorhamphus aleuticus, auriculae small, broadly pointed projections Cassin's Auklet; and Lunda cirrhata, the (Fig. 348). Palpi short, broad; segment 1 Tufted Puffin. Eley also collected 2 female without anterior spur, but with strong pos- --I. uriae from Lutra canadensis, the River terior spur directed laterally more than Otter, on a beach 1.5 Im south of Trinidad, posteriorly (Fig. 348). Hypostome short, Humboldt Co., on V-2-1970. The latter host rounded apically; denticles 2/2 to base association is considered accidental. (Fig. 348). Scutum slightly wider than Harald Johnson collected 1 male, 9 females, long; widest at midlevel; lateral carinae and 12 engorged nymphs from Flatiron Rock, faint. Coxal spurs (Fig. 349) similar to Trinidad Head, Humboldt Co., on VI-22-1972. those of female, but internal spur of coxa We have examined the male of this collec- I wider and longer than external spur. tion; all others vere destroyed during virus Larva. Undescribed. isolation attempts . Females of L. woodi resemble 1. angustus and r. ochotonae, but differ in having 2/2 Ixodes (Pholeoixodes) modi Bishopp dentition throughout the length of the hypo- (Figs. 340-349; Map 22) stome and prominent ridges on the dorsal side of the basis capituli, which restrict Ixodes angustus modi Bishopp, 1911, Proc. the posterolateral extensions of the porose Biol. Soc. Wash. 24:205, pl. 11, fig. areas. The hypostome of the male woodi 13 (female); Cooley and Kohls, 1945, differs from that of angustus, in which the U.S. Pub. Health Serv., Natl. Inst. crenulations are indistinct. Nymphs of Health Bull. 184:163, fig. 656-1 angustus differ from woodi in having an (nymph); Kohls, 1950, J. Parasitol. anterior spur on segment 1 of the palpi. 36:65, fig. 1 (male). Nymphs of ochotonae differ from woodi in that the posterior spur of palpal segment 1 DIAGNOSIS. Female (Fig. 344). Capitu- is directed more posteriorly than laterally, lum (Figs. 346, 347) without cornua or and in having a weaker internal spur on auriculae. Porose areas moderate in size, coxa I. well separated. Palpi up to 3.8 X longer --I. woodi is primarily found on woodrats, than wide; segment 1 with laterally project- Neotoma species, and their nests. Keirans ing suboval ventral plate. Hypostome borne and Clifford (1978) list collections from on tapered median anterior projection of Alabama, Arizona, California, Colorado, basis capituli; apex narrowly rounded; den- Indiana, Kansas, Oklahoma, Texas and Utah. ticles 2/2, with lateral denticles larger --I. woodi is known from skunks as well as than medians, Ventral basis capituli elon- woodrats and their nests in foothill and gate, smooth, lacking lateral constrictions. low mountainous regions of central and Scutum (Fig. 345) distinctly longer than southern California. wide, oval, widest caudad to midlevel. Scapulae large, triangular. All coxae with CALIFORNIA RECORDS. KERN CO.: 2-4 miles short external spurs; coxa I with slender SW Glenville, 1 nymph from Neotoma fuscipes, THE TICKS OF CALIFORNIA 77

11-(17-18)-1959 (A.M. Barnes). MADERA CO. : Rhipicephalus sanguineus (Latreille) San Joaquin Experimental Range, 2 nymphs (Figs. 350-356; Map 23) from 1. fuscipes, XI-14-1950 (O'Neal and N. Brown Dog Tick Cohn). MONTEREY CO. : Hastings Reservation: 4 nymphs from Spilogale gracilis, XI-19-38; Ixodes sanguineus Latreille, 1806, Genera 4 nymphs from E. fuscipes, 1-11-39 (R. crustaceorum et insectorum 1:157. Holdenried); 2 nymphs from E. fuscipes, Rhipicephalus sanguineus; Koch, 1844, Arch. 111-28-39 (R. Holdenried); 2 nymphs from Naturgesch. 10:239; Hooker et al., 1912, Spilogale gracilis, 11-26-40; 1 nymph from U.S. Dept. Agric., Bur. Entomol. Bull. -N. fuscipes, XI-13-40; 1 nymph from Mephitis 106:102 (all instars described); Cooley, mephitis, X-26-44; 1 nymph from 2. fuscipes, 1946, U.S. Pub. Health Serv., Natl. 1-10-46; 3 nymphs from E. fuscipes, 11-2-46; Inst. Health Bull. 187:24. 11 nymphs from E. fuscipes, 111-3-46. NAPA Rhipicephalus s t igmat icus Gers t Hc ker , 1873, CO.: 10 miles S Honticello, 2 nymphs from Gliederthiere, p. 496. -N. fuscipes, 111-17-1957 (R.O. Schuster) . Rhipicephalus texanus Banks, 1908, U.S. PLUMAS CO.: 2.4 miles E Virgilia, 1 female Dept. Agric., Bur. Entomol. Tech. Ser. and 2 nymphs from 1. fuscipes, X-25-1961 15 :34. (A.M. Barnes). RIVERSIDE CO.: 3 collections from San Timoteo Canyon: 2 nymphs from 5. DIAGNOSIS. Female. Body (Pig. 351) fuscipes, 1-8-1952; 1 female from nest of oval to elliptical; widest posteriorly. -N. fuscipes, 1-8-1952; 2 nymphs and 1 larva Unfed tick small: 2.4-2.7 mm long from tips (presumably of same species) from nest of of scapulae X 1.44-1.68 mm wide (Cooley, -N. fuscipes, 1-27-1952 (Ryckman et al.). 1946a). Capitulum hexagonal (Fig. 353). SAN LUIS OBISPO CO.: 1 mi. N, 4 mi. E Santa Cornua present as salient corners. Porose Margarita, 3 nymphs from E. fuscipes, X-31- areas small, with long axis longitudinal. 1951 (R. Murray). SANTA BARBARA CO.: Van- Palpi short, broad; segment 1 with ventral, denberg Air Force Base, 1 female from E. subtriangular plate and curved, irregularly fuscipes, XI-10-1980 (H. Madon). branched, parallel setae on inner margin;

Genus Rhipicephalus Koch

Rhipicephalus Koch, 1844, Arch. Naturgesch. (Berlin) 10: 238. Rhipicephalus Nuttall and Warburton, 1911, Ticks: a monograph of the Ixodoidea 2:122. Rhipicephalus Cooley, 1946, U.S. Pub. Health Serv., Natl. Inst. Health Bull. 187:23. Rhipicephalus Hoogstraal, 1956, African Ixodoidea 1:583.

Type of genus: Rhipicephalus sanguineus (Latreille, 1806).

DIAGNOSIS (California species only). All instars inornate, with eyes and 9 festoons. Basis capituli hexagonal, with tri- angulate lateral projections (Pig. 303). Palpi of adults and nymphs short, with segment 1 barely, or not at all, visible in dorsal view (Fig. 353). Adults with coxa I bifid. Adults and nymphs with distinct anal groove posterior to anus (Fig. 355). Larvae with 1 pair of posthypostomal setae; scutum with 3 pairs of setae; 8 pairs of marginal dorsal setae, of which 4 pairs anterior to MAP 23. Distribution in California of sensilla sagittiformia; 2 pairs of central Rhipicephalus sanguineus (Latreille), solid dorsal setae. circles. 78 BULLETIN OF T?E CALIFORNIA INSECT SURVEY segment 2 with similar setae. Hypostome The brown dog tick, also known as the long, clavate, with distinct corona and 313 kennel tick, is the only representative of dentition. Coxa I with external spur thin- the large genus Rhipicephalus in the United ner and more acutely pointed than internal States. Hoogstraal (1956) states that this spur; coxae 11-IV with small rounded exter- tick now occupies practically all countries nal spurs. Coxae I1 and 111 with internal between 50°1 and about 35%. As the name spurs as small rounded mounds. Spiracular implies, it is a common pest on dogs. Its plate short, comma-shaped. habits differ from those of other ticks Male. (Fig. 350). Elongate oval, widest occurring on dogs in California in that behind middle. Size similar to that of typically it is a resident of kennels or unfed female. Capitulum shape similar to homes housing dogs, rather than the exterior that of female. Coxae as in female, but environment. It is a 3-host tick in which coxa IV with small internal spur (Fig. 352). the larva, nymph, and adult each engorge Genital aperture at level of anterior region once, typically on the dog, dropping off of coxae 11. Anal, adanal, and accessory after each feeding to seek harborage in the plates distinct (Fig. 352). Spiracular kennel or home. Eggs are laid in cracks in plate elongate, comma-shaped. the housing, often in the upper parts of Nymph. Basis capituli (Fig. 354) hexag- the structure. Other hiding places, such onal in dorsal view. Anal groove (Fig. 355) as under the edge of rugs, furniture, etc., in form of arc bordering posterior of anus. may be infested. In subtropical areas or Coxa I (Fig. 356) with external spur widely in heated structures , breeding may occur separated from smaller internal spur; coxae throughout the year. Nuttall (1915) I1 and 111 with small external spurs (Fig. reported that at a 3OoC temperature the life 356); coxa IV with external spur rudimentary cycle of &. sanguineus could be completed or absent. Other diagnostic features as in as little as 63 days. He also found that given for genus. adults may survive without food up to 568 Larva. Basis capituli hexagonal in days. dorsal view. Other diagnostic features as In the United States, the brown dog tick given for genus. is important primarily as a blood sucking nuisance of dogs, but it has tremendous Adults of 2. sanguineus may be distin- potential as a carrier of disease agents. guished from all other ticks occurring nor- It is a vector of the protozoan parasite mally in California by the hexagonal basis Babesia canis, causing canine jaundice. capituli. A cattle tick, Boophilus annula- This pathogenic protozoan is common in many %, which at one time was introduced occa- tropical regions, and although it is uncom- sionally over the southern borders of the mon in the United States, it has been state, has a hexagonal basis capituli, but reported from California (Levine, 1973). is easily distinguishable by a lack of mar- Outside of the United States, &. sanguineus ginal festoons and presence of prominent has been associated with a variety of patho- transverse ridges on the short palpi. gens, including Babesia & of equines in Nymphs of s. sanguineus may be most easily central Asia; E. gibsoni of canines in confused with some species of Dermacentor, India and China; Rickettsia conori , causing from which they differ in having the basis boutonneuse fever in man in southern Europe capituli hexagonal dorsally, with the and Africa; and Rickettisa siberica, the anterolateral margins converging abruptly causal agent of Siberian tick typhus in man. to the base of the hypostome, in contrast In Mexico, E. sanguineus has been implicated to the more gradual convergence of the in the transmission to man of rickettsiae antero-lateral margins in Dermacentor. of the spotted fever group, and Burgdorfer Rhipicephalus nymphs also differ from Derma- et al. (1975, 1978) reported it as a vector centor in possessing a much smaller palpal in Mississippi of Rickettsia rhipicephali. segment 1 which is typically not visible The causal agent of canine rickettsiosis , dorsally and forms only a small angular Ehrlichia canis, is transmitted by s. E- plate ventrally; the palpal apices are more guineus in the Middle East, Africa, and the sharply triangular dorsally, and the anal Orient . groove is more distinct. The latter charac- R. sanguineus in the United States is ter is best observed in unfed, unmounted almost exclusively a parasite of dogs; only specimens. Larvae differ from the similar- rarely has it been found attacking man. appearing larvae of Dermacentor in having 4 But in some countries it is found on a rather than 3 marginal dorsal setae anterior variety of domestic and wild mammals as well to the sensillae sagittiformia. as on birds, and in certain areas attacks THE TICKS OF CALIFORNIA 79 man readily. It probably represents a through April, but in all probability breed- species complex. ing is continuous throughout the year in The majority of California collections San Diego and Imperial counties and in warm are associated with the large population housing in which infested dags live. As centers, but it appears that infestations seen in Table 19, California records of the may be encountered wherever dogs are main- brown dog tick have been made almost exclu- tained. Adult ticks have been collected in sively from dogs or their housing. A single every month; nymphs have been collected in collection of an adult female tick is every season. Collections of larvae are recorded from an unidentified bird from Kern relatively few, with no records for January County.

TABLE 19. California Records of Rhipicephalus sanguineus (Latreille)

Host or Source Tick Collections

Adults Nymphs Larvae Unrecorded Instars Bird 1 - - - Homo sapiens 1 1 - Canis familiaris 53 22 5 Kennel 4 1 1 House 13 5 1 Vegetation 1 - - Miscellaneous 1 - - 1 Unrecorded 20 5 1 24

Literature Cited

Allred, D. M., D. E. Beck, and L. D. White Misc. Publ. Entomol. SOC. Amer. 8: 1960 Ticks of the genus Ixodes in Utah. 161-376. Brigham Young Univ. Sci. Bull., Banks, N. Biol. Ser. 1: 1-42. 1904 Some Arachnida from California. Allred, D. M., and E. J. Roscoe Proc. Calif. Acad. Sci. 3: 331-369. 1956 Life history of the tick Dermacen- 1908 A revision of the Ixodoidea, or -tor parumapertus in Utah. J. Para- ticks, of the United States. U.S. sitol. 42: 516-522. Dept. Agric., Bur. Entomol. Tech. Anastos, G., and C. N. Smith Ser. 15; 61 pp. 1957 The male, nymph, and larva of 1909 Three new ticks from the United Ixodes brunneus Koch (Acarina: States. Proc. Entomol. SOC. Wash. Ixodidae). J. Parasitol. 43: 10: 170-173. 535-541. 1910 New American mites (Arachnoidea, Arthur, D. R. Acarina). Proc. Entomol. SOC. 1960 Ticks: a monograph of the Ixodoi- Wash. 12: 2-12. dea, part V. Cambridge Univ. 1912 New American mites. Proc. Entomol. Press, London; 251 pp. SOC. Wash. 14: 96-99. Arthur, D. R., and K. Snow Bequaert, J. C. 1968 Ixodes pacificus Cooley and Kohls, 1946 The ticks, or Ixodoidea, of the 1943: its life history and occur- northeastern United States and rence. Parasitology 58: 893-906. eastern Canada. Entomol. Amer., Audouin, V. J. n.s. 25: 73-232. 1826 Expl. P1. Arachnides Egypte. In Berlese, A. Descr. de l'lgypte, Paris 1: 182- 1888 Acari Austro-Americani. Boll. 186, pl. 9 (cited by Oudemans, della SOC. Entomol. Italiana 20: 1936). 171-223. Augustson, G. F. Birula, A. 1939 A new Ixodes (Acarina: Ixodidae). 1895 Ixodidae novi vel parum cogniti Bull. S. Calif. Acad. Sci. 38: Musei Zoologici Academiae Caesareae 141-147. Scientiarum Petropolitanae. Izv. Ault, S. R., and K. D. Elliott Imp. Akad. Nauk, ser. 5, 2: 353- 1979 Spider predation (Araneae: Salti- 364. cidae) on Ornithodoros coriaceus Bishopp, F. C. (Acarina: Argasidae), with a survey 1911 Some new North American Ixodidae, of the predators of the genus m- with notes on other species. Proc. thodoros. J. Med. Entomol. 15: Biol. SOC. Wash. 24: 197-208. 570-571. 1912 A new species of Dermacentor, and Bacha, W. J., Jr. notes on other North American Ixo- 1957 The life history of Otobius lago- didae. Proc. Biol. SOC. Wash. 25: philus. J. Parasitol. 43: 560-565. 29-37. Balashov, Y. S. Bishopp, F. C., and H. L. Trembley 1968 Bloodsucking ticks (Ixodoidea) : 1945 Distribution and hosts of certain vectors of diseases of man and North American ticks. J. Parasi- animals. Nauka Publ. House, Lenin- tol. 31: 1-54. grad; 319 pp. Trans. and ed. 8. Bishopp, F. C., and H. P. Wood Hoogstraal and R. Tatchell, 1972, 1913 The biology of some North American

81 82 BULLETIN OF THE CALIFORNIA INSECT SURVEY

ticks of the genus Dermacentor. 1938 The genera Dermacentor and Otocen- Parasitology 6: 153-187. -tor (Ixodidae) in the United Boyer, K. M., R. S. Munford, G. 0. Maupin, States, with studies in variation. C. P. Pattison, M. D. Fox, A. H. Barnes, U.S. Pub. Health Serv., Natl. Inst. W. L. Jones, and J. E. Maynard Health Bull. 171: 1-89. 1977 Tick-borne relapsing fever: an 1944a Ixodes neotomae, a new species from interstate outbreak originating at California (Acarina: Ixodidae). Grand Canyon National Park. Amer. Pan-Pacif. Entomol. 20: 7-12. J. Epidem. 105: 469-479. 1944b Ixodes ozarkus n.sp. and Ornitho- Brinton, L. P., D. E. Beck, and D. M. Allred doros aquilae n.sp., with notes on 1965 Identification of the adults, -0. talaje and 0. kelleyi (Ixodoi- nymphs and larvae of ticks of the deal. J. Parasitol. 30: 287-294. genus Dermacentor Koch (Ixodidae) 1946a The genera Boophilus, Rhipicephalus in the western United States. and Haemaphysalis (Ixodidae) of Brigham Young Univ. Sci. Bull., the New World. U.S. Pub. Health Biol. Ser. 5; 44 pp. Serv., Natl. Inst. Health Bull. Brunetti, 0. A. 187: 1-54. 1965 Tick paralysis in California deer. 1946b Ixodes holdenriedi, a new species Calif. Fish and Game 51: 208-210. of tick from a pocket gopher in Burgdorfer, W., L. P. Brinton, W. L. California (Acarina: Ixodidae). Krinsky, and R. N. Philip Pan-Pacif. Entomol. 22: 103-104. 1978 Rickettsia rhipicephali, a new Cooley, R. A., and G. M. Kohls spotted fever group rickettsia from 1938 Two new species of ticks (Ixodes)

the brown dog tick, Rhipicephalus~~ from California (Acarina: Ixodi- sanguineus. In J. Kazar, R. A. dae). U.S. Pub. Health Serv. Rept. Ormsbee, and I. N. Taresevich, 53: 1616-1621. eds. , Rickettsiae and rickettsial 1940 Two new species of Argasidae diseases, pp. 307-316. Veda Publ. (Acarina: Ixodoidea). U.S. Pub. House, Bratislava, USSR. Health Serv. Rept. 55: 925-933. Burgdorfer, W., D. J. Sexton, R. K. Gerloff, 1941a Three new species of Ornithodoros R. L. Anacker, R. N. Philip, and L. A. (Acarina: Ixodoidea). U.S. Pub. Thomas Health Serv. Rept. 56: 587-594. 1975 Rhipicephalus sanguineus: vector 1941b Further new species of Ornithodoros of a new spotted fever group from bats (Acarina: Argasidae). rickettsia in the United States. U.S. Pub. Health Serv. Rept. 56: Infect. hun. 12: 205-210. 910-914. Camin, J. H., and R. W. Drenner 1943 Ixodes californicus Banks, 1904, 1978 Climbing behavior and host-finding Ixodes pacificus n.sp. and Ixodes

of larval rabbit ticks (Haema-~ conepati n.sp. (Acarina: Ixodidae). physalis leporispalustris). J. Pan-Pacif. Entomol. 19: 139-147. Parasitol. 64: 905-909. 1944a The genus Amblyomma (Ixodidae) in Canestrini, G. the United States. J. Parasitol. 1890 Prospetto dell'acarofauna italiana. 30: 70-111. Pt. 4. Famiglie: Tetranychini, 1944b The Argasidae of North America, Ixodini, Argasiai. Padova, pp. Central America and Cuba. her. 427-540. Midland Naturalist, monogr. 1; 152 Clifford, C. M., and G. M. Kohls PP. 1963 Observations on the life cycle of 1945 The genus Ixodes in North America. the tick Argas brevipes Banks, U.S. Pub. Health Serv., Natl. Inst. 1908. J. Parasitol. 49: 527. Health Bull. 184: 1-246. Clifford, C. M., D. E. Sonenshine, E. L. Curtice, C. Atwood, C. S. Robbins, and L.E. Hughes 1891 The biology of the cattle tick. 1969 Tests on ticks from wild birds J. Comp. Med. Vet. Arch. 12: 313- collected in the eastern United 319. States for rickettsiae and viruses. 1892 Parasites, being a list of those Amer. J. Trop. Med. Hyg. 18: 1057- infesting the domesticated animals 1061. and man in the United States, J, Cooley, R. A. Comp. Med. Vet. Arch, 13: 223-236. 1936 Ornithodoros parkeri, a new species Davis. G. E. w on rodents. U.S. Pub. Health Rept. 1959 Ornithodoros parkeri: distribution f 51: 431-433. and host data; spontaneous infec- p: ." THE TICKS OF CALIFORNIA 83

tion with relapsing fever spiro- Hafniae. (Cited by Nuttall et al., chetes. U.S. Pub. Health Sen. 1908.1 Rept. 54: 1345-1349. Fairchild, G. B., G. M. Kohls, and V. J. 1940 Ticks and relapsing fever in the Tipton United States. U.S. Pub. Health 1966 The ticks of Panama. In R. L. Serv. Rept. 55: 2347-2351. Wenzel and V. J. Tipton, Ectopara- 1941 Ornithodoros parkeri and relapsing sites of Panama, pp. 162-219. fever spirochetes in Utah. U.S. Field Mus. Nat. Hist., Chicago. Pub. Health Serv. Rept. 56: 2464- Fe Idman-Muhsam, B . 2468. 1973 Autogeny in soft ticks of the genus Davis, G. E., and A. J. Mamos Ornithodoros (Acari: Argasidae). 1956 An atypical Ornithodoros hernsi J. Parasitol. 59: 536-539. from Utah (Ixodoidea: Argasidae). Fischer de Waldheim, G. J. Parasitol. 42: 293-296. 1823 Notice sur 1'Argas de Perse (Malleh JhgBs, A. de Mianeh) decrit par les voyageurs 1876 Turicata de Guanajuato. El Reper sous le nom de punaise venimeuse torio de Guanajuato (Newspaper), de Miana. Mem. SOC. Imp. Nat. April 25. (Cited by Megnin, P. Hoscou 6: 269-283. (Cited by 1885 Les Argas du Mexique. J. Nuttall et al., 1908.) Anat. Physiol., Paris, 21: Garcia, R. 460-47 5. ) 1963 Studies on the ecology of Ornitho- 1884 Turicata y garrapata de Guanajuato. -doros coriaceus Koch (Acarina: La Naturaleza (Mexico) 6: 195-198, Argasidae). Ph.D. diss. , Univ. pl. 4. California, Berkeley; 177 pp. Gastfriend, A. 1955 New host records for the immature stages of the tick Dermacentor pl. 3, fig. 2. parumapertus. J. Parasitol. 41: Dunn, L. H. 63-65. 1933 Observation on the host selection George, J. E. of Ornithodoros talaje Guerin in 1971 Drop off rhythms of engorged rabbit Panama. Amer. J. Trop. Med. 13: ticks, Haemaphysalis le oris alus 47 5-483. -tris (Packard, 18-y Eads, R. B., G. C. Menzies, and G. G. Ixodidae). J. Med. Entomol. 8: Hightower 461-479. 1956 The ticks of Texas, with notes on Gerstiicker, A. their medical significance. Texas 1873 Gliderthiere (Insekten, Arachniden, J. Sei. 8: 7-24. Myriapoden und Isopoden). Baron Easton, E. R., and R. L. Goulding Carl Claus von der Decken's Travels 1974 Ectoparasites in two diverse in East Africa, 3: Scientific habitats in western Oregon: I. Results, Part 2; 542 pp. Leipzig Ixodes (Acarina: Ixodidae). J. and Heidelberg. Med. Entomol. 11: 413-418. Gregson, J. D. Eklund, C. H., G. M. Kohls, and J. M. 1941 Two new species of ticks from Brennan British Columbia (Ixodidae). 1955 Distribution of Colorado tick Canad. Entomol. 73: 220-228. fever and virus-carrying ticks. 1942 A new species of tick found on J. Amer. Med. Assoc. 157: 335-337. shrews (Ixodes soricis n.sp.1. Eley, T. J. Canad. Entomol. 74: 137-139. 1977 Ixodes uriae (Acari: Ixodidae) from 1954 The male of Ixodes signatus Birula a river otter. J. Med. Entamol. (Acarina: Ixodidae). Canad. Ento- 13: 506. mol. 86: 275-277. Fabricius, J. C. 1956 The Ixodoidea of Canada. Canada 1787 Mantissa insectorum sistens species Dept. Agric. Sci. Serv., Entomol. nuper detectas adiectis synonymis, Publ. 930; 92 pp. observationibus, descriptionibus, 1971 Studies on two populations of emendationibus, vol. 2; 382 pp. Ixodes kingi Bishopp (Ixodidae). Hafaiae. (Cited by Robinson, 1926.) Canad. J. 2001. 49: 591-597. 1794 Entomologica systematica, etc., Gregson, J. D., and G. M. Kohls Vol. 4. Ticks, pp. 425-435. 1952 The male of Ixodes soricis Gregson 84 BULLETIN OF THE CALIFORNIA INSECT SURVEY

(Acarina: Ixodidae). Canad. Ento- Dept. Agric., Bur. Entomol. Bull. mol. 84: 185-188. 106; 239 pp. Groocock, C. M., W. R. Hess, and W. J. Hughes, L. E., C. M. Clifford, R. Gresbrink, Gladney L. A. Thomas, and J.E. Keirans 1980 Experimental transmission of 1976 Isolation of a spotted fever African swine fever virus by rickettsia from the Pacific Coast Ornithodoros coriaceus, an argasid tick, Ixodes pacificus, in Oregon. tick indigenous to the United Amer. J. Trop. Med. Hygiene 25: States. Amer. J. Vet. Res. 41: 513-516. 519-594. Hunter, W. D., and W. A. Hooker GuBrin-MBneville, F. E. 1907 Information concerning the North 1849 Description de 1'Argas talaje. American fever tick, with notes on Rev. Mag. 2001. 1: 342-344, pl. 9. other species. U.S. Dept. Agric., Hews, W. B. Bur. Entomol. Bull. 72; 87 pp. 1916 The pajaroello (0. coriaceus Koch), International Commission of Zoological with special reference to life Nomenclature history and biting habits. J. 1978 Opinion 1107. Bull. 2001. Nomen- Parasitol. 2: 137-142. clature 35: 88. Hems, W. B., and C. M. Wheeler Jessup, D. A. 1935 Tick transmission of California 1979 Tick paralysis in a grey fox. J. relapsing fever. J. Econ. Entomol. Wildlife Dis. 15: 271-272. 28: 846-855. Johnson, D. E. Hixson, H. 1966 Ticks of Dugway proving ground and 1932 The life history and habits of vicinity and their host associa- Ixodes sculptus Neumann (Ixodidae). tions. Utah Acad. Proceedings 43: Iowa State Coll. J. Sci. 7: 36-42. 49-66. Hokama, Y., and J. A. Howarth Johnson, H. N., and J. Casals 1977 Dry-ice ((202) trap for efficient 1972 Arboviruses associated with marine field collection of Ornithodoros bird colonies of the Pacific Ocean coriaceus (Acarina: Argasidae). region. In A. I. Cherepanov et J. Med. Entomol. 13: 627-628. al., eds., Transcontinental con- Hornsher, P. J., and D. E. Sonenshine nections of migratory birds and 1975 Scanning electron microscopy of their role in the distribution of ticks for systematic studies: fine arboviruses, pp. 200-204. Nauka structure of Haller's organ in ten Publ. House, Siberian Branch, species of Ixodes. Trans. Amer. Novosibirsk. Microscop. SOC. 94: 368-374. Jones, E. K., and C. M. Clifford Hoogstraal, H. 1972 The systematics of the subfamily 1956 African Ixodoidea, vol. 1. Ticks Ornithodorinae (Acarina: Argasi- of the Sudan (with special refer- dae), vol. 5. A revised key to ence to Equatoria Province and with larval Argasidae of the Western preliminary reviews of the genera Hemisphere and description of seven Boophilus, Margaropus and e- new species of Ornithodoros. Ann. lemma). V.S. Dept. Navy, Bur. Med. Entomol. SOC. Amer. 65: 730-740. Surg.; 1101 pp. Karsch, F. 1973 Acarina (ticks). In Viruses and 1879 Zwei neue Arachniden des Berliner invertebrates, ed. A. J. Gibbs, Museums. Mitt. Miinch. Entomol. pp. 89-103. North Holland Publ. Ver. 3: 95-96. Co., Amsterdam. Keirans, J. E., and C. M. Clifford Hoogstraal, H., C. H. Clifford, J. E. 1978 The genus Ixodes in the United Keirans, and H.Y. Wassef States: a scanning electron micro- 1979 Recent developments in biomedical scope study and key to the adults. knowledge of Argas ticks (1x0- J. Med. Entomol. Suppl. 2; 149 pp. doidea: Argasidae). In Recent Koch, C. L. advances in acarology, vol. 2, ed. 1844 Systematische Uebersicht iiber die J. G. Rodriguez, QQ- 269-278. Ordnung der Zechen. Arch. Natur- Academic Press. gesch. (Berlin), 10: 217-239. Hooker, W. A., F. C. Bishopp, and H. P. Wood 1847 Ubersicht des Arachniden systems. 1912 The life-history and bionomics of Niirnberg 4; 136 pp., pl. 1-30. some North American ticks. U.S. THE TICKS OF CALIFORNIA 85

Kohls, G. M. and descriptions of three new 1947 Notes on the tick Ixodes howelli species. Ann. Entomol. SOC. Amer. Cooley and Kohls, with descrip- 58: 331-364. tions. J. Parasitol. 33: 57-61. Kramer, P., and C. J. Neuman 1950 Description of the male Ixodes 1883 Acariden, wahrend der Vega-Expedi- woodi Bishopp (Acarina: Ixodidae). tion eingesammelt. In Norden- J. Parasitol. 36: 65-66. skib'ld, Vega-Expeditiones Vetensk. Kohls, G. M., and C.M. Clifford (Stockholm) 3: 519-529. 1963 Ornithodoros sparnus sp.n., a para- Lane, R. S., R. W. Emmons, D. V. Dondero, site of wood rats, Neotoma spp., and B. C. Nelson and deer mice, Peromyscus spp., in 1981 Ecology of tick-borne agents in Utah and Arizona (Acarina: Argasi- California. I. Spotted fever group dae). J. Parasitol. 49: 857-861. rickettsiae. Am. J. Trop. Med. 1966 Three new species of Ixodes from Hyg. 30: 239-252. Mexico and description of the male Latreille, P. A. of J-. auritulus auritulus Neumann, 1795 Observations sur la vari6t6 des -I. conepati Cooley and Kohls, and organes de la bouche des tiques, -I. lasallei Mendez and Ortiz (Aca- etc. Hag. Encycl. J. Sci. Lett. rina: Ixodidae). J. Parasitol. Arts 4: 15-20. 52: 810-820. 1796 Precis des caracteres g6n6riques 1968 The subgenus Persicargas (Ixodoi- des insectes disposes dans un ordre dea, Argasidae, Argas). 6. A. (I.) naturel; 202 pp. giganteus, n.sp., from wild birds 1806 Genera crustaceorum et insectorum in western United States and secundum ordinem naturalem in Sonora, Mexico. Ann. Entomol. SOC. familias disposita, iconibus Amer. 61: 1113-1116, exemplisque plurimis explicata, Kohls, G. M., and H. Hoogstraal vol. 1; 302 pp. Paris and Argento- 1960 Observations on the subgenus Argas rati. (Ixodoidea, Argasidae, Argas). 2, Levine, N. D. -A. cooleyi, new species, from 1973 Protozoan parasites of domestic western North American birds. Ann. animals and of man, 2nd ed. Entomol. SOC. Amer. 53: 625-631. Burgess, Minneapolis. Kohls, G. M., H. Hoogstraal, and C. M. Linnaeus, C. Clifford 1758 Systema naturae, ed. 10. Regnum 1961 Observations on the subgenus Argas animale; 824 pp. Holmiae. (Ixodoidea, Argasidae, Argas). 5. Loomis, E. C. Study of A. brevipes Banks, 1908, 1961 Life histories of ticks under from birds in Arizona and Califor laboratory conditions (Acarina: nia, U.S.A., and Baja California, Ixodidae and Argasidae). J. Para- Plexico. Ann. Entomol. SOC. her. sitol. 47: 91-99. 54: 869-877. Loomis, E. C., and R. B. Bushnell Kohls, G. M., H. Hoogstraal, C. M. Clifford, 1968 Tick paralysis in California live- and M. N. Kaiser stock. Am. J. Vet. Res. 29: 1089- 1970 The subgenus Persicar as (Ixodoi- 1093. dea, Argasidae-9. Redes- Loomis, E. C., and D. P. Furman cription and New World records of 1977 The pajaroello tick. Univ. Calif. Argas (P.) persicus (Oken), and Div. Agric. Scis., leaflet 2503; 3 resurrection, redescription, and PP- records of A. (E.) radiatus Mackie, T. T., G. W. Hunter 111, and C. B. Railliet, A. (E.) sanchezi Dug&, Worth and A. (5)miniatus Koch, New 1946 In A manual of tropical medicine, World ticks misidentified as A. pp. 62-73. W. B. Saunders, Phila- (E.) persicus. Ann. Entomol. SOT. delphia. Amer. 63: 590-606. Marx, G. Kohls, G. M., D. E. Sonenshine, and C. ?I. 1895 Obituary notice of his work. Proc. Clifford Entomol. SOC. Wash. 3: 195-201, 1 1965 The systematics of the subfamily PI* Ornithodorinae (Acarina: Argasi- da Massa, A. J., and H. E. Adler dae). 11. Identification of the 1979 Avian spirochaetosis: natural larvae of the Western Hemisphere transmission by Argas (Persicargas) 86 BULLETIN OF THE CALIFORNIA INSECT SURVEY

sanchezi (Ixodoidea: Argasidae) Nuttall, G. H. F., and C. Warburton and existence of different sero- 1908 On a new genus of the Ixodoidea, logic and immunologic types of together with a description of Borrelia anserina in the United eleven nev species of ticks. Proc. States. Amer. J. Vet. Res. 40: Camb. Phil. SOC. 14: 392-416. 154-157. 1911 Ixodidae. Ticks: a monograph of Mazzotti, L. the Ixodoidea, part 2: 105-348. 1943 Una nueva especie de Ornithodoros Cambridge Univ. Press, London. en Mexico. Rev. Inst. Salub. 1915 Ticks: a monograph of the Ixodoi- Enferm. Trop. 4: 371-374. dea, part 3. The genus Haema- McIvor, B. C. physalis, pp. 349-550. Cambridge 1937 A new species of Ornithodoros tick Univ. Press, London. from California (Acarina: Ixodoi- Nuttall, G. E. F., C. Warburton, W. F. dea). J. Parasitol. 23: 363-367. Cooper, and L. E. Robinson Murray, A. 1908 Ticks: a monograph of the Ixodoi- 1877 Economic entomology. Aptera. dea, part 1. Argasidae, pp. 1-104. Chapman and Hall, London. 1: 433 Cambridge Univ. Press, London. PP Oken, L. Naversen, D. N., and L. W. Gardner 1818 Sogenannte gif tige Wanze in 1978 Erythema chronicum migrans in Persien. Isis: 1567-1570. (Cited America. Arch. Dermatol. 114: by Nuttall et al., 1908.) 253-254. Oliver, J. H., Jr., and L. P. Brinton Nelson. B. C. 1972 Cytogenetics of ticks (Acari: 1973 Tick paralysis in a dog caused by Ixodoidea), 7. Spermatogenesis in Ixodes pacificus Cooley and Kohls the Pacific Coast tick, Dermacentor (Acarina: Ixodidae). Calif. Vector occidentalis Harx (Ixodidae). J. Views 20: 80-82. Parasitol. 58: 365-379. Neumann I L. G. Oliver, J. E., Jr., P. R. Wilkinson, and G. 1896 Revision de la famille des 1x0- M. Kohls dides. I. Argasines. M6m. Soc. 1972 Observations on hybridization of 3 Zool. France 9: 1-44. species of North imerican Dewa- 1897 Revision de la famille des Ixo- centor ticks. J. Parasitol. 58: dides. N6m. SOC. Zool. Fr. 10: 380-384. 324-420. Osborn, H. 1899 Revision de la famille des Ixo- 1899 List of insects hitherto known dides. M6m. Soc. Zool. Fr. 12: from the Pribilof Islands. IX. 107-294. Acarina. In D. S. Jordan et al., 1901 R6vision de la famille des Ixo- Fur seals and fur-seal islands, dides. H6m. Soc. Zool. Fr. 14: north Pacific Ocean, part 3: 553- 249-372. 554. Washington. 1904 Notes sur les Ixodid&, 11. Arch. Ouderans, A. C. Parasitologie 8: 444-464. 1936 Kritische Historisch Overzicht der 1905 Notes sur les Ixodid&, 111. Arch. Acarologie 3: 746-797. Amsterdam. Parasitologie 9: 225-241. Packard, A. S., Jr. 1907 Notes sur les Ixodid&, V. Arch. 1869 List of hymenopterous and lepidop- Parasitologie 11: 215-232. terous insects collected by the Nuttall, G. H. F. Smithsonian expedition to South 1912 Notes on ticks, 11. (1) New species America, under Prof. James Orton (Amblyomma, Raemaphysalis). (2) (appendix to report on Articu- Ixodes putus: description of the lates). Ann. Rept. Peabody Acad. hitherto unknown larval stage. Sci. (Salem, Mass.) 1: 56-69. Parasi tology 5 : 50-60. Philip, C. B., J. F. Bell, and C. L. Larson 1915 Observations on the biology of 1955 Evidence of infectious diseases Ixodidae, part 2. Parasitology 7: and parasites on a peak population 408-456. of black-tailed jack rabbits in 1916 Notes on ticks, IV. Relating to Nevada. J. Wildlife Mgmt. 19: 225- the genus Ixodes and including a 233. description of three new species Philip, C. B., and G. E. Davis and two new varieties. Parasi- 1940 Relapsing fever: data implicating tology 8: 294-3370 Ornithodoros hermsi as a vector in THE TICKS OF CALIFORNIA 87

northern Idaho. U.S. Pub. Health Serdyukova, G. V. Rept. 55: 504-507. 1955 On the question of differential Pickard-Cambridge, 0. characteristics of larvae and 1876 On a new order and some new genera nymphs of Ixodidae. Zool. Zhurnol. of Arachnida from Kerguelen' s 34: 1037-1051. Island. Proc. 2001. SOC. London, Shad, G. A. 17: 258-265. 1958 First report of Ornithodoros PochB, R. M., and J. E. Keirans (Otobius) megnini from pronghorn 1975 Report of a tick, Ornithodoros antelope, Antilocapra americana. rossi (Acarina: Argasidae), from J. Parasitol. 44: 514. the spotted bat, Euderma maculatum Silva-Goytia, R., and A. Elizondo (Chiroptera: Vespertilionidae). J. 1952 Estudios sobre fiebre manchada en Med. Entomol. 12: 503. Mexico, 2. Parasitos hematofagos Pocock, R. I. encontrados naturalmente infec- 1907 Ticks. In Albut and Rolleston, tados. Rev. Med. 32: 278-282. eds., A system of medicine, 2: 187- Sonenshine, D. E. 203. Macmillan, London. 1962 External anatomv of the bat tick Porter, G. S., M. D. Beck, and I. M. Stevens Ornithodoros kelleyi Cooley and 1932 Relapsing fever in California. Kohls (Acarina: Argasidae). J. her. J. Pub. Health 22: 1136-1140. Parasitol. 48: 470-485. Riley, C. V. Sonenshine, D. E., and I. J. Stout 1869 Remarks on the Ixodes bovis. 1970 A contribution to the ecology of Reports on the diseases of cattle ticks infesting wild birds and in the United States made to the rabbits in the Virginia-North Commissioner of Agriculture, Carolina Piedmont (Acarina: Ixodi- Washington, p. 168. dae). J. Med. Entomol. 7: 645-654. Robinow, M., and T. B. Carroll Steere, A. C., and S. E. Malawista 1938 Tick paralysis due to the bite of 1979 Cases of Lyme disease in the United the American dog tick: report of a States: Locations correlated with case observed in Georgia. J. Amer. distribution of Ixodes dammini. Med. Assn. 111: 1093-1094. Ann. Intern. Med. 91: 730-733. Robinson, L. E. Stiles, C. W. 1926 The Genus Amblyomma. Ticks: a 1908 The common tick (Dermacentor monograph of the Ixodoidea, part andersoni) of the Bitter Root 4. Cambridge Univ. Press, London; Valley. U.S. Pub. Health Mar. 285 pp., 7 pl., 130 figs. Hosp. Serv. Rept. 23: 949. Salmon, D. E., and C. W. Stiles 1910 The taxonomic value of the micro- 1901 The cattle ticks (Ixodoidea) of scopic structure of the stigmal the United States. 17th Rept. U.S. plates in the tick genus Dermacen- Bur. Anim. Ind., Washington, pp. -tor. U.S. Pub. Health Mar. Hosp. 380-488. Serv., Hyg. Lab. Bull. 62; 72 pp. Say, T. Tully, J. G., B. F. Whitcomb, H. F. Clark, 1821 An account of the Arachnides of and D. L. Williamson the United States. J. Acad. Nat. 1977 Pathogenic Mycoplasmas: cultivation Sci. Phila. 2: 59-82. and vertebrate pathogenicity of a Schmidtmann, E. T., R. B. Bushnell, E. C. new spiroplasma. Science 195: 892- Loomis, M. N. Oliver, and J. H. Theis 894. 1976 Experimental and epizootiologic Van Peenen, P. F. D., and J. F. Duncan evidence associating Ornithodoros 1968 Piroplasms (Protozoa: Sarcodina) coriaceus Koch (Acari: Argasidae) of wild mammals in California. with the exposure of cattle to Bull. Wildlife Dis. Assoc. 4: 3-8. epizootic bovine abortion in Cali- Westrom, D. R. fornia. J. Med. Entomol. 13: 292- 1975 The population dynamics and dis- 299. tribution of ectoparasites on Schulze, P. Black-tailed deer (Odocoileus 1930 Erster beitrag zu einer Zeckenfauna hemionus columbianus) in Mendocino Danemarks. Sitzungsber . Abh. Co., California. Ph.D. Diss., Naturforsch. Ges. Rostock (1927- Univ. California, Berkeley; 184 pp. 291, 2: 120-124. Wheeler, C. M. 1935 A new species of tick which is a a8 BULLETIN OF THE CALIFORNIA INSECT SURVEY

vector of relapsing fever in Cali- Baffin's Bay and Barrow Straite, fornia. ber. J. Trop. Med. 15: in the years 1850-1851, 2: appen- 435-438. dix, ccviii-ccxi. London. Wheeler, C. M., W. B. Hems, and K. F. Meyer Wright, J. E. 1935 A new tick vector of relapsing 1969 Photoperiodic induction of diapause fever in California. Proc. SOC. in larvae of the winter tick, Der Expmtl. Biol. and Med. 32: 1290- macentor albipictus. Ann. Entomol. 1292. SOC. her. 62: 285-287. Wherry, W. B., and F. C. Wellman Yates, T. L., D. B. Pence, and G. L. 1909 Ticks on the California ground Launchbaugh squirrel. Entomol. News 20: 376. 1979 Ectoparasites from seven species White, A. of North American moles (Insec- 1852 Insects and Aptera. In F. C. tivora: Talpidae. J. Med. Entomol. Sutherland, Journal of a voyage in 16: 166-168. Pia tes 90 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 1

Figs. 1-4. Fig. 1, Dermacentor occidentalis Harx, d, dorsal view; Figs. 2-4, 2. andersoni Stiles, p, dorsal and ventral views and spiracular plate. THE TICKS OF CALIFORNIA 91

leg I 3

-‘\y 1

leg 4

---internal spur, coxa I

-:-external coxal spurs --genital aperture

-&-genital groove

-anus

2 3

--- goblet (aeropyle)

* ------dorsal prolongation 4 92 BUUETIN OF THE CALIFORNIA INSECT SURVEY

Plate 2

Figs. 5-11. Figs. 5 and 6, Dermacentor, 9; capitulum, dorsal and ventral views; Fig. 7, Ixodes, tarsus, leg I; Fig. 8, Ixodes, d, ventral view; Fig. 9, Ixodes , capitulum, ventral view; Fig. 10, Rhipicephalus sanguineus (Latreille), d, posterior idiosoma, ventral view; Fig. 11, -I. angustus Neumann, nymph, capitulum, ventral view. THE TICKS OF CALIFORNIA 93

c- - - corona

--1 ,---palpal segment Ip

I I ---palpus I

t segment I : ---I

'i -- posteroventral spur I rJ------auricula L2 9

-spiracular plate

accessory shield

-----anal groove adanal shield ----anterior spur of palpal segment I postanal - median groove

----posterior spur of 10 palpal segment I

11 94 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 3

Figs. 12-15. Fig. 12, Ornithodoros coriaceus Koch, 9, ventral view; Fig. 13, 0. coriaceus Koch, tarsus, leg I; Fig. 14, Argas brevipes Banks, lateral sutural line and marginal dorsal integument; Fig. 15, 4. persicus (Oken), capitulum, ventral view. THE TICKS OF CALIFORNIA 95

subapical dorsal r--- hood protuberance I

---dorso-ventral groove

preanal groove -- '\ -transverse postanal groove 13

---- postpalpal seta

- posthypostomal seta

sutural line 15

-dorsal cuticula

.ventral cuticula

14 96 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 4

Figs. 16-20. Figs. 16 and 17, Dermacentor andersoni Stiles, larva, dorsal and ventral views; Figs. 18 and 19, Ornithodoros, larva, dorsal and ventral views; Fig. 20, Argasidae, larva, tarsus, leg I. A, apical; AV, apicoventral; B, basal; BV, basoventral; Cd 1, central dorsal seta 1; Md, marginal dorsal seta; MV, midventral; PC, paracapsular; Ph, posthypostomal seta; PL, posterolateral; Pm, posterior median seta; PM, posteromedian; Sc, scuta1 seta; St, sternal setae. THE TICKS OF CALIFORNIA 97

dermal sensilla sensilla porosa posthypostoma I seta

- coxa I internal spur

_e----*coxae ZI 8 ID external spur

16 17

dorsal plate

19 PC

AV Mv PL EJV

20 98 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 5

Figs. 21-24. Argas brevipes Banks: Fig. 21, Dorsum, 9; Fig. 22, Venter, 9; Fig. 23, Capitulum, d; Fig. 24, Marginal dorsal integument, 9. THE TICKS OF CALIFORNIA 99

21 22

23 24 100 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 6

Figs. 25-28. Argas cooleyi Kohls and Hoogstraal: Fig. 25, Venter, 9; Fig. 26, Capitulum, Q; Fig. 27, Tarsus, leg I, 9; Fig. 28, Marginal dorsal integument, d . THE TICKS OF CALIFORNIA 101

25 26

27 102 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 7

Figs. 29-32. Argas persicus (Oken): Fig. 29, Dorsum, 9; Fig. 30, Venter, g; Fig. 31, Capitulum, 9; Fig. 32, Marginal dorsal integument, d. THE TICKS OF CALIFORNIA 103

i? t

29 30

31 32 104 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 8

Figs. 33-38. Argas sanchezi Dughs: Fig. 33, Dorsum, d; Fig. 34, Venter, d; Fig. 35, Dorsum, P; Fig. 36, Venter, 0; Fig. 37, Capitulum, 0; Fig. 38, Posterolateral marginal integument, 0.

33 34 THE TICKS OF CALIFORNIA 105

35 36

37 38 106 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 9

Figs. 39-42. Ornithodoros concanensis Cooley and Kohls: Fig. 39, Venter, Q; Fig. 40, Tarsus, leg I, Q; Fig. 41, Capitulum, Q; Fig. 42, Capitulum and cheeks, 9. THE TICKS OF CALIFORNIA 10 7

39 40

41 42 108 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 10

Figs. 43-46. Ornithodoros coriaceus Koch: Fig. 43, Dorsum, d; Fig. 44, Eyes on anterolateral margin, adult; Fig. 45, Capitulum, adult; Fig. 46, Tarsus, leg I, adult. THE TICKS OF CALIFORNIA 109

43 44

45 46 110 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 11

Figs. 47-50. Ornithodoros dyeri Cooley and Kohls, d: Figs. 47 and 48, Dorsal and ventral views; Fig. 49, Capitulum; Fig. 50, Hypostome detail. THE TICKS OF CALIFORNIA 111

47 40

49 50 112 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 12

Figs. 51-56. Ornithodoros hermsi Wheeler, Hems and Meyer: Figs. 51 and 52, Dorsal and ventral views, 9; Fig. 53, Capitulum, adult; Fig. 54, Genital orifice, 6; Fig. 55, Tarsus, leg I, adult; Pig. 56, Genital orifice, g-

51 52 THE TICKS OF CALJFORNIA 11 3

53 54

55 56 114 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 13

Figs. 57-60. Ornithodoros hermsi Wheeler, Hems and Meyer: Fig. 57, Dorsolateral integument, adult; Figs. 58-60, Capitulum and dorsal and ventral views, larva. THE TICKS OF CALIFORNIA 115

57 58

59 116 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 14

Figs. 61-63. Ornithodoros kelleyi Cooley and Kohls, 6: Fig. 61, Ventral view; Fig. 62, Anterolateral profile; Fig. 63, Capitulum. THE TICKS OF CALIFORNIA 11 7

61 62

63 118 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 15

Figs. 64-69. Ornithodoros parkeri Cooley, 9 : Figs. 64 and 65, Dorsal and ventral views; Fig. 66, Capitulum and genital region; Fig. 67, Capitulum detail; Fig. 68, Tarsus, leg I; Fig. 69, Dorsal integument detail.

64 65 THE TICKS OF CALIFORNIA 119

66 67

68 69 120 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 16

Figs. 70-73. Ornithodoros sparnus Kohls and Clifford: Fig. 70, Ventral view, 9; Fig. 71, Dorsal integument, 6; Fig. 72, Capitulum, Q; Fig. 73, Tarsus, leg I, 9. THE TICKS OF CALIFORNIA 121

70 71

72 73 122 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 17

Figs. 74-77. Ornithodoros stageri Cooley and Kohls, 0: Figs. 74 and 75, Dorsal and ventral views; Fig. 76, Capitulum; Fig. 77, Tarsus, leg I. THE TICKS OF CALIFORNIA 123

74 75

76 77 124 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 18

Figs. 78-81. Ornithodoros talaje (Gu&in-M&eville), Q: Figs. 78 and 79, Dorsal and ventral views; Fig. 80, Capitulum; Fig. 81, Antero- lateral view of body wall and hood. CALIFORNIA THE TICKS OF 125

78 79

80 81 126 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 19

Figs. 82-85. Ornithodoros turicata (Dug&s), 6: Figs. 82 and 83, Dorsal and ventral views; Fig. 84, Capitulum; Fig. 85, Tarsus, leg I. THE TICKS OF CALIFORNIA 127

84 128 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 20

Figs. 86-88. Ornithodoros yumatensis Cooley and Kohls, Q: Fig. 86, Ventral view; Fig. 87, Tarsus, leg IV; Fig. 88, Capitulum. THE TICKS OF CALIFORNIA 129

86 87

88 130 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 21

Figs. 89-94. Otobius lagophilus Cooley and Kohls: Figs. 89 and 90, Dorsal and ventral views, 9; Fig. 91, Capitulum, 9; Fig. 92, Dorsal integument detail, 9; Fig. 93, Dorsal view, nymph; Fig. 94, Capitulum, ventral view, nymph.

89 90 THE TICKS OF CALIFORNIA 131

91 92

93 94 132 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 22

Figs. 95-100. Otobius megnini (Dugh): Figs. 95 and 96, Dorsal and ventral views, 9; Fig. 97, Dorsal integument detail, 9; Fig. 98, Ventral view, nymph; Fig. 99, Capitulum, 9; Fig. 100, Capitulum, nymph.

95 96 THE TICKS OF CAL,IFORNIA 133

98 97

100 99 134 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 23

Figs. 101-104. Amblyomma americanum (Linnaeus): Fig. 101, Dorsal view, d; Fig. 102, Dorsal view, 9; Fig. 103, Capitulum, dorsal view, *; Fig. 104, Capitulum, ventral view, 9. THE TICKS OF CALIFORNIA 135

101 102

103 104 136 BULLETIN OF THE CAT.,IFORNIA INSECT SURVEY

Plate 24

Figs. 105-108. Boophilus annulatus (Say): Figs. 105 and 106, Dorsal and ventral views, 6;Figs. 107 and 108, Dorsal and ventral views, 9. 80 1

901

LET VINXCMIW3 10 SmIL 3H3 138 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 25

Figs. 109-114. Boophilus annulatus (Say) : Fig. 109, Capitulum, dorsal view, 6; Fig. 110, Capitulum, dorsal view, 9; Fig. 111, Coxa and trochanter I, d; Fig. 112, Tarsus, leg 111, 9 ; Fig. 113, Dorsal view, nymph; Fig. 114, Capitulum, dorsal view, nymph.

109 110 THE TICKS OF CALIFORNIA 139

111 112

113 114 140 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 26

Figs. 115-118. Dermacentor albipictus (Packard): Fig. 115, Dorsal view, d; Fig. 116, Dorsal view, 0; Fig. 117, Coxae I-IV, 0; Fig. 118, Spiracular plate, 6. 81 1 LLL

91 1 Sll 142 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 27

Figs. 119-122. Dermacentor albipictus (Packard) , nymph: Fig. 119, Dorsal view; Fig. 120, Coxae I-IV; Fig. 121, Capitulum, dorsal view; Fig. 122, Spiracular plate. THE TICKS OF CALIFORNIA 143

119 120

121 122 144 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 28

Pigs. 123-128. Dermacentor andersoni Stiles: Figs. 123 and 124, Dorsal and ventral views, 6; Figs. 125 and 126, Dorsal and ventral views, 9; Fig. 127, Capitulum, dorsal view, d; Fig. 128, Capitulum, dorsal view, 9.

123 124 THE TICKS OF CALIFORNIA 145

125 126 146 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 29

Figs. 129-134. Dermacentor andersoni Stiles: Fig. 129, Spiracular plate, d; Fig. 130, Spiracular plate, 0; Fig. 131, Coxa I and capitulum, Q; Fig. 132, Ventral aspect with coxae 1-111, nymph; Fig. 133, Capitu- lum, dorsal view, nymph; Fig. 134, Ventral aspect with coxa IV, nymph.

129 130 THE TICKS OF CALIFORNIA 14 7

131 132

133 134 148 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 30

Figs. 135-140. Dermacentor hunteri Bishopp: Fig. 135, Dorsal view, d; Fig. 136, Dorsal view, 9; Fig. 137, Spiracular plate, 9; Fig. 138, Spiracular plate, nymph; Fig. 139, Scutum and capitulum, nymph; Fig. 140, Ventral view, nymph.

135 136 1 OP 6E 1

LE 1 150 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 31

Figs. 141-146. Dermacentor occidentalis Marx: Fig. 141, Dorsal view, 6; Fig. 142, Dorsal view, Q; Fig. 143, Capitulum, dorsal view, d; Fig. 144, Capitulum, dorsal view, p; Fig. 145, Scutum, 0; Fig. 146, Scuta1 detail, adult.

141 142 THE TICKS OF CALIFORNIA 151

143 144

145 146 152 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 32

Figs. 147-152. Dermacentor occidentalis Marx: Fig. 147, Coxa I, 9; Fig. 148, Spiracular plate, d; Fig. 149, Dorsal view, nymph; Fig. 150, Spiracular plate, nymph; Fig. 151, Ventral view, nymph; Fig. 152, Capi- tulum and coxae I, ventral view, nymph.

I47 148 THE TICKS OF CALIFORNIA 153

149

151 152 154 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 33

Figs. 153-159. Dermacentor parumapertus Neumann: Fig. 153, Dorsal view, d; Fig. 154, Dorsal view, Q ; Fig. 155, Coxa I, d; Fig. 156, Scutum, Q; Fig. 157, Coxae I-IV, nymph; Fig. 158, Capitulum, ventral view, nymph; Fig. 159, Spiracular plate, nymph.

153 154 6s 1 LE 1

9s 1 ss 1

z i 156 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 34

Figs. 160-163. Dermacentor variabilis (Say) : Fig. 160, Dorsal view, d; Fig. 161, Dorsal view, 9; Fig. 162, Spiracular plate, *; Fig- 163, Spiracular plate, 9. THE TICKS OF CALIFORNIA 15 7

160 161

162 163 15 8 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 35

Figs. 164-167. Dermacentor variabilis (Say), nymph: Figs. 164 and 165, Dorsal and ventral views; Fig. 166, Capitulum, ventral view; Fig. 167, Spiracular plate. THE TICKS OF CALIFORNIA 159

164 165

-- I E &*+"*+* * -*a:-

166 167 160 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 36

Figs. 168-171. Haemaphysalis chordeilis (Packard): Fig. 168, Dorsal view, 6; Fig. 169, Dorsal view, engorged Q; Fig. 170, Capitulum, ventral view, d; Fig. 171, Capitulum, ventral view, 0. THE TICKS OF CALIFORNIA 161

168 169

170 171 162 BULLETIN OF THE CALIFORNIA INSE(JT SURVEY

Plate 37

Figs. 172-175. Haemaphysalis chordeilis (Packard): Fig. 172, Scutum, Q ; Fig. 173, Capitulum, dorsal view, 9; Fig. 174, Coxae I-IVY d; Fig. 175, Capitulum, dorsal view, nymph. THE TICKS OF CALIFORNIA 163

172 173

174 175 164 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 38

Figs. 176-179. Haemaphysalis leporispalustris (Packard) : Fig. 176, Dorsal view, d; Fig. 177, Dorsal view, 0; Figs. 178 and 179, Capitulum, d, dorsal and ventral views. THE TICKS OF CALIFORNIA 165

176

178 179 166 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 39

Figs. 180-183. Haemaphysalis leporispalustris (Packard) : Fig. 180, Coxa and trochanter IV, d; Fig. 181, Anterior dorsal view, Q; Fig. 182, Capitulum, dorsal view, nymph; Fig. 183, Coxae and trochanters I and 11, nymph. THE TICKS OF CALIFORNIA 167

180 181

182 183 168 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 40

Figs. 184-187. Ixodes angustus Neumann: Fig. 184, Dorsal view, 6; Fig. 185, Dorsal view, Q; Fig. 186, Capitulum and scutum, d; Fig. 187, Capitulum, ventral view, d. THE TICKS OF CALIFORNIA 169

184 185

186 187 170 BULLETIN OF "BE CALIFORNIA INSECT SURVEY

Plate 41

Figs. 188-191. Ixodes angustus Neumano: Figs. 188 and 189, Capitu- lum, 9, dorsal and ventral views; Fig. 190, Capitulum, ventral view, nymph; Fig. 191, Coxa I, nymph.

i THE TICKS OF CALIFORNIA 171

191 172 BULLETIN OF THE CALIFORNIA INSECT SURVEY i

Plate 42

Figs. 192-195. Ixodes auritulus Neumann, Q: Fig. 192, Dorsal view; Fig. 193, Scutum; Figs. 194 and 195, Capitulum, dorsal and ventral views. THE TICKS OF CALIFORNIA 173

192 193

194 195 174 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 43

Figs. 196-199. Ixodes auritulus Neumann: Fig. 196, Scutum, d ; Fig. 197, Coxae I-IV, d; Fig. 198, Coxa I, nymph; Fig. 199, Capitulum, ventral view, nymph. (Figs. 196 and 197 courtesy Keirans and Clifford, 1978. ) THE TICKS OF CALIFORNIA 175

196 197

198 199 176 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 44

Figs. 200-203. Ixodes brunneus Koch, Q: Fig. 200, Dorsal view; Fig. 201, Coxae I-IV; Figs. 202 and 203, Capitulum, dorsal and ventral views. 3 i THE TICKS OF CALIFORNIA 177 d

200 201

202 203 178 BELLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 45

Figs. 204-209. Ixodes brunneus Koch: Fig. 204 and 205, Capitulum, d, dorsal and ventral views; Fig. 206, Scutum, d; Fig. 207, Coxae I-IV, trochanters 11-111, d ; Fig. 208, Capitulum, dorsal view, nymph; Fig. 209, Basis capituli, ventral view, nymph. (Figs. 204-207 courtesy Keirans and Clifford, 1978.)

204 205 THE TICKS OF WIFORNIA 179

206 207

208 209 180 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 46

Figs. 210-213. Ixodes hearlei Gregson: Fig. 210 and 211, Capitulum, 9, dorsal and ventral views; Fig. 212, Scutum, 9; Fig. 213, Capitulum, ventral view, nymph. THE TICKS OF CALIFORNIA 181

210 21 1

212 213 182 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 47

Figs. 214-217. Ixodes holdenriedi Cooley, 9: Figs. 214 and 215, Capitulum, dorsal and ventral views; Fig. 216, Scutum; Fig. 217, Coxae I-IV. (Figs. 214-217 courtesy Keirans and Clifford, 1978.) THE TICKS OF CALIFORNIA 183

214 215

216 217 184 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 48

Figs. 218-223. Ixodes howelli Cooley and Kohls: Figs. 218 and 219, Dorsal and ventral views, *; Figs. 220 and 221, Dorsal and ventral views, 9; Fig. 222, Posterior, ventral view, *; Fig. 223, Capitulum, ventral view, *.

218 219 THE TICKS OF CALIFORNIA 185

220 221

222 223 186 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 49

Figs. 224-227. Ixodes howelli Cooley and Kohls: Fig. 224, Capitu- lum, ventral view, 9; Fig. 225, Coxae and trochanters I and 11, 0; Fig. 226, Capitulum, ventral view, nymph; Fig. 227, Coxa I, nymph. THE TICKS OF CALIFORNIA 187

224 225

226 227 188 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 50

Figs. 228-231. Ixodes jellisoni Cooley and Kohls, 0: Fig. 228, Dorsal view; Fig. 229, Coxa I; Figs. 230 and 231, Capitulum, dorsal and ventral views (tip of hypostome missing). THE TICKS OF CALIFORNIA

228 229

230 23 1 190 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 51

Figs. 232-235. Ixodes kingi Bishopp: Fig. 232, Dorsal view, *; Fig. 233, Dorsal view, 9; Fig. 234, Scutum, 6; Fig. 235, Scutum, 9. THE TICKS OF CALIFORNIA 191

232 233

234 235 192 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 52

Figs. 236-241. Ixodes kingi Bishopp: Figs. 236 and 237, Capitulum, d, dorsal and ventral views; Figs. 238 and 239, Capitulum, 0 , dorsal and ventral views; Fig. 240, Scutum and capitulum, nymph; Fig. 241, Capitulum, ventral view, nymph.

236 237 1PZ OPZ

6EZ 8EZ 194 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 53

Figs. 242-245. Ixodes neotomae Cooley: Pigs. 242 and 243, Dorsal and ventral views, d; Figs. 244 and 245, Dorsal and ventral views, 9. THE TICKS OF CALIFORNIA 195

242 243

244 245 196 BULLETIN OF THE CAT..IFORNIA INSECT SURVEY

Plate 54

Pigs. 246-249. Ixodes neotomae Cooley: Figs. 246 and 247, Capitu- lum, d, dorsal and ventral views; Figs. 248 and 249, Capitulum, 9, dorsal and ventral views. THE TICKS OF CALIFORNIA 197

246 247

249 198 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 55

Figs. 250-253. Ixodes ochotonae Gregson, 0: Fig. 250, Scutum and basis capituli; Fig. 251, Coxae I and 11; Figs. 252 and 253, Capitulum, dorsal and ventral views. TICKS THE OF CALIFORNIA 199

250 251

252 253 200 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 56

Figs. 254-257. Ixodes ochotonae Gregson: Fig. 254, hypostome, 9; Fig. 255, Scutum, nymph; Figs. 256 and 257, Capitulum, nymph, dorsal, and ventral views. THE TICKS OF CALIFORNIA 201

254 255

256 257 2 02 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 57

Figs. 258-263. Ixodes pacificus Cooley and Kohls: Fig. 258, Dorsal view, d, mating with 9, ventral view; Fig. 259, Dorsal view, 9; Figs. 260 and 261, Capitulum, 6, dorsal and ventral views; Fig. 262, Capitu- lum, dorsal view, 9; Fig. 263, Coxae I and I1 and capitulum, 9.

258 259 THE TICKS OF CALIFORNIA 203

260 26 1

262 263 204 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 58

Figs. 264-267. Ixodes pacificus Cooley and Kohls, nymph: Figs. 264 and 265, Dorsal and ventral views; Fig. 266, Capitulum, dorsal view; Fig. 267, Coxae I and capitulum. THE TICKS OF CALIFORNIA 205

264 265

266 267 2 06 BULLETIN OF THE CALIFORNIA INSECT SURVEY I

Plate 59

Figs. 268-273. Ixodes peromysci Augustson: Fig. 268, Dorsal view, d; Fig. 269, Dorsal view, Q; Fig. 270, Scutum and basis capituli, 6; Fig. 271, Scutum and basis capituli, 9; Fig. 272, Capitulum, ventral view, d; Fig. 273, Capitulum, ventral view, 9.

268 269 THE TICKS OF CALIFORNIA 207

270 271

272 273 208 BULLETIN OF THE CALIFORNIA INSECI: SURVEY

Plate 60

Figs. 274-279. Ixodes peromysci Augustson: Fig. 274, Coxae I-IV, d; Fig. 275, Ventral view, d; Figs. 276 and 277, Capitulum, nymph, dorsal and ventral views; Fig. 278, Scutum, nymph; Fig. 279, ventral view, nymph.

274 275 THE TICKS OF CALIFORNIA 209

276 277

278 279 210 BULLETIN OF THE CAL1M)RNIA INSECT SURVEY

Plate 61

Figs. 280-285. Ixodes rugosus Bishopp: Fig. 280, Dorsal view, d; Fig. 281, Dorsal view, engorged 0; Figs. 282 and 283, Capitulum, d, dorsal and ventral views; Figs. 284 and 285, Capitulum, Q, dorsal and ventral views.

280 281 THE TICKS OF CALIFORNIA 211

282 283

284 285 212 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 62

Fig. 286-291. Ixodes rugosus Bishopp: Fig. 286, Scutum and dorsal integument, d; Fig. 287, Scutum, 9; Fig. 288, Coxae I-IVY Q; Fig. 289, Scutum and capitulum, nymph; Fig. 290, Capitulum, ventral view, nymph; Fig. 291, Coxae I-IV, nymph.

286 287 THE TICKS OF CALIFORNIA 213

288 289

290 291 214 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 63

Figs. 292-295. Ixodes sculptus Neumann, 6: Figs. 292 and 293, Dorsal and ventral views; Figs. 294 and 295, Capitulum, dorsal and ventral views. THE TICKS OF CALIFORNIA 215

292 293

294 295 216 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 64

Figs. 296-301. Ixodes sculptus Neumann: Fig. 296, Dorsal view, 9; Fig. 297, Coxa I, Q; Figs. 298 and 299, Capitulum, 9, dorsal and ventral views; Figs. 300 and 301, Capitulum, nymph, dorsal and ventral views.

296 297 THE TICKS OF CALIFORNIA 217

298 299

300 301 218 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 65

Figs. 302-305. Ixodes signatus Birula: Fig. 302, Dorsal view, d; Fig. 303, Dorsal view, 9 ; Figs. 304 and 305, Capitulum, d , dorsal and ventral views. THE TICKS OF CALIFORNIA 219

302 303

304 305 Plate 66

Figs. 306-309. Ixodes signatus Birula: Pigs. 306 and 307, Capitulum, 9, dorsal and ventral views; Fig. 308, Coxa I, Q; Fig. 309, Capitulum and coxa I, ventral view, nymph. THE TICKS OF CALIFORNIA 221

306 307

308 309 222 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 67

Figs. 310-313. Ixodes soricis Gregson, Q: Fig. 310, Dorsal view; Fig. 311, Coxae I-IV; Figs. 312 and 313, Capitulum, dorsal and ventral views. THE TICKS OF CALIFORNIA 223

31 0 31 1

31 2 31 3 224 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 68

Figs. 314-319. Ixodes spinipalpis Hadwen and Nuttall: Fig. 314, Scutum and basis capituli, d; Fig. 315, Scutum, 9; Fig. 316, Capitulum, ventral view, d; Fig. 317, Capitulum, ventral view, 9; Fig. 318, Coxa I, 9; Fig. 319, Capitulum, ventral view, nymph.

314 315 THE TICKS OF CALIFORNIA 225

316 31 7

318 319 226 BULLETIN OF THE CAT.,IFORNIA INSECT SURVEY

Plate 69

Figs. 320-323. Ixodes texanus Banks, 6: Fig. 320, Scutum; Fig. 321, Coxae I-IV; Figs. 322 and 323, Capitulum, dorsal and ventral views. (Figs. 320-323 courtesy Keirans and Clifford, 1978.) THE TICKS OF CALIFORNIA 227

320 32 1

322 323 228 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 70

Figs. 324-329. Ixodes texanus Banks: Fig. 324, Scutum, 9; Fig. 325, Coxae I-IV 0; Figs. 326 and 327, Capitulum, 0, dorsal and ventral views; Figs. 328 and 329, Capitulum, nymph, dorsal and ventral views.

324 325 THE TICKS OF CALIFORNIA 229

326 327

328 329 2 30 BULLETIN OF THE CALlFORNIA INSECT SURVEY

Plate 71

Figs. 330-333. Ixodes uriae White: Figs. 330 and 331, Dorsal and ventral views, 6; Figs. 332 and 333, Dorsal and ventral views, 9. THE TICKS OF CALIFORNIA 2 31

330 33 1

332 333 232 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 72

Figs. 334-339. Ixodes uriae White: Fig. 334, Posterior ventral view, d; Fig. 335, Capitulum, ventral view, d; Fig. 336 and 337, Capi- tulum, 9, dorsal and ventral views; Fig. 338, Capitulum, dorsal view, nymph; Fig. 339, Scutum, nymph.

334 335 THE TICKS OF CALIFORNIA 233

336 337

338 339 2 34 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 73

Figs. 340-343. Ixodes woodi Bishopp, d: Fig. 340, Dorsal view; Fig. 341, Coxae I-IV; Fig. 342, Anterior scutum and capitulum; Fig. 343, Capitulum, ventral view. THE TICKS OF CALIFORNIA 235

340 341

342 343 236 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 74

Figs. 344-349. Ixodes woodi Bishopp: Fig. 344, Dorsal view, 9; Fig. 345, Scutum, 9; Figs. 346 and 347, Capitulum, P, dorsal and ventral views; Fig. 348, Capitulum, ventral view, nymph; Fig. 349, Coxa I, nymph -

344 345 237 THE TICKS OF CALIFORNIA

347 346

348 349 BULLETIN OF THE CALIFORNIA INSECT SURVEY

Plate 75

Figs. 350-356. Rhipicephalus sanguineus (Latreille): Fig. 350, Dorsal view, d; Fig. 351, Dorsal view, 9; Fig. 352, Posterior ventral view and coxae 111-IV, 6; Fig. 353, Capitulum, dorsal view, 9;Fig. 354, Capitulum; dorsal view, nymph; Fig. 355, Anal groove, nymph; Fig. 356, Coxae I-IV, nymph.

350 351 THE TICKS OF CALIFORNIA 239

352 353

354 355 356