Historic, archived document Do not assume content reflects current scientific knowledge, policies, or practices. . 1 Pe) Wie ee ae eee ae : i » 4 . ~ ‘ 4k 7x . 2 Lie, "aed ee Y ¥ ~/- ire ¥.* ‘ Ce bad , ri = * ale BUREAU OF ENTOMOLOGY—BULLETIN No, 106. \ | L. O. HOWARD, Entomologist and Chief of Bureau. ‘THE LIFE HISTORY AND BIONOMICS OF SOME q NORTH AMERICAN TICKS. BY W. A. HOOKER, F. C. BISHOPP, anp H. P. WOOD. UNDER THE DIRECTION OF W. D. HUNTER. _IssueD SEPTEMBER 7, 1912. om, Ms ane ihe till ane ITTY - Allbe a Laan é , f = aes ere fils *, all sf oie iN Ee 2 * Le il We" i é J 2 at Na | ‘ih » WASHINGTON: GOVERNMENT PRINTING OFFIOE. 1912. ‘ ee eer OF AGRICULTURE, BUREAU OF ENTOMOLOGY—BULLETIN No, 106. L. O. HOWARD, Entomologist and Chief of Bureau. THE LIFE HISTORY AND BIONOMICS OF SOME NORTH AMERICAN TICKS. BY W. A. HOOKER, F. C. BISHOPP, anp H. P. WOOD. UNDER THE DIRECTION OF W. D. HUNTER. Issunp SEPTEMBER 7, 1912. WASHINGTON: GOVERNMENT PRINTING OFFIOE. 1912, BUREAU OF ENTOMOLOGY. L. O. Howarp, Entomologist and Chief of Bureau. C. L. Maruatr, Entomologist and Acting Chief in Absence of Chief. R. 8. Crirrton, Executive Assistant. W. F. Taster, Chief Clerk. H. CHITTENDEN, 2n charge of truck crop and stored product insect investigations. D. Hopxrins, in charge of forest insect investigations. D. Hunter, in charge of southern field crop insect investigations. M. WEBSTER, 1n charge of cereal and forage insect investigations. L. QUAINTANCE, in charge of deciduous fruit insect investigations. EK. F. Pures, in charge of bee culture. D. M. Roaers, in charge of preventing spread of moths, field work. Roiia P. Currie, in charge of editorial work. MasBet CotcorD, 1n charge of library. F. A. W. iB: uN SOUTHERN FieELD Crop INSECT INVESTIGATIONS. W. D. Hunter, in charge. F. C. BisHopp, A. H.-Jennines, H. P. Woop, W. V. Kine, engaged in tick investi- gations. W. D. Pierce, G. D. Smits, J. D. MrircuHett, Harry Prnxus, B. R. Coan, R. W. MoRELAND, engaged in cotton-boll weevil investigations. A.C. Morean, G. A. Runner, 8S. E. Crump, D. C. Parman, engaged in tobacco insect investigations. T. E. Hottoway, E. R. BARBER, engaged in sugar cane insect investigations. E. A. McGreeor, W. A. THomas, engaged in red spider and other cotton insect investi- gations. J. L. WEBB, engaged in rice insect investigations. R. A. Cootzy, D. L. Van Dine, A. F. Conran, C. C. KrumBuaar, collaborators. 2 LETTER OF TRANSMITTAL. U.S. DEPARTMENT OF AGRICULTURE, BuREAU OF ENTOMOLOGY, Washington, D. C., November 29, 1911. Srr: I have the honor to transmit herewith a manuscript entitled “The Life History and Bionomics of Some North American Ticks,” prepared by Messrs. W. A. Hooker, F. C. Bishopp, and H. P. Wood, under the direction of Mr. W. D. Hunter, of this bureau. Ticks are of considerable importance in the United States in two respects. Two species are the sole agents in the transmission of certain impor- tant diseases. One of these diseases is the well-known splenetic fever of cattle and the other the Rocky Mountain spotted fever of human beings. The information contained in this manuscript will be of immediate value in connection with the eradication of the cattle tick and also of the form which transmits spotted fever. The remaining species treated in this manuscript are now of importance principally as parasites of domestic animals. In many localities they present serious problems to the farmer. Moreover, there is a possibility that some of the species not known at present to be carriers of diseases will eventually be found to be of importance in connection with disease transmission. For these reasons full information regarding the life history of the various species is in demand. The work upon which this bulletin is based has extended from 1907 to the present time. The manuscript is intended to be a com- pendium of information regarding ticks which will remain of direct value for many years and make unnecessary the publication of spe- cial bulletins on some of the species treated. . I recommend that this manuscript be published as Bulletin No. 106 of the Bureau of Entomology. Respectfully, L. O. Howarp, Entomologist and Chief of Bureau. Hon. JAMEs WILson, Secretary of Agriculture. CONTEN ES: : Page. IIe Re Te DOP R ih oes ie in wp oe Ro Dee Oe iba bake «2 oso 13 manne ie eminence a eh eo 8 SIS 2 Sue's Leap VL .'e 14 Byetematic position and classification: .. 2.0... -. i ee ae ee 14 Collecting, preservaae, and mountmg !). 50.20.5202 848 2b ee ol 17 PiemAgM Te, SIN OTEOD 4 8209S 2025 2822s Seu ys eens a IDOE... - 18 Pintary cn tile wiclomical study of ticke_. 22:2. SPY 20 POs Guicets LMCI oe ot LPR cy tg 21 Getieral life history... -:.22...-2:: 24 erent behead ee a Bae kegel ee elt el lo ee 25 EMR teh oe a a oa en ea hee oe ee ene ease 32 Penne rer) rare Onan a en Se tn a as eens 33 ene RE rs i ee ee Sh aye eg Boe ae aan ete 34 menos empoved in atudies Of ticks...2-. 2.22... 222i... 2. he ents 35 ORES SEEDS tlc os eee i le SRO oh eta op an eR ae 4] ane TIOTENR St Teo tye ke Se Ss aos - ae eat eae AE 44 Rena tient wn ne Arvanidee 25. Jo Se 2 2 2 ek ee ee gece es 45 ot ee Lo ee arg ieee aee oaae ea A eR oh a a 45 Tew tek, Argus mintatus Woclt. 0...) 22. Aen eee dees - 52 46 we GL ED SLs eG oe ee ae ne alle ge pete neemen Neste MWg Fe ies 5s ge Maia aa 46 PS Cae a ae te ene aE RL ee Lea 47 Pecgea pears) chit aiitean 2S oS, a ee wee aw T- - ee MER et ee ee ee oe Ts oa cnt oA SRR a 48 WUC Oe See ee en a ane LE ee OM 59 Pierre MIME IAMCES: 2 52.4 a2 52 to ne SLE Cee 60 ieremeree cnn me ee et CSR ED. Laks SU 60 EE es) ale ee a een oe SC eat el ee 61 TS TS 50 PS pe le eae es eyo tae aie AA ae pelea eae ane A, Ae a 61 The spinose ear tick, Ornithodoros megnini (Dugés)........-.----.----- 61 coll Teh 2. hss teed ee ee eet a olpet hal anaemia i a ogee 62 Host relationship. . Ato. fas kets dh ieee ee ah gm tapes 9 eco ty a 62 Siepueavisiesl distribution: Dich ot et aRla caf m Reli bens decay yp betsy 62 SERN! A See ere he ka renee en 64 aE Ree ra ee or TN ee was 5 a wee Ss oe ws 68 eee AUPE INIENS Sool ial tte Sk es i go a an +S 68 MEM NNCREE Benes te en SF So ee ne we Se mom < 69 Oh eee le dl ee eS See een RE 69 Development of the Ixodide............ : 69 ee EPET TIER ee arin Ci i ote PELE < ois Lien Did nd. w nonce sins 4 oes 7 The black-legged tick, Iwodes scapularis Say...........-.--.--------- 76 Os OAC DN IE ae belle edie a EY One ey yee 76 Host relationship. . : 7 Geographical distribution:. Pe eee nok! oP, vt & eee 77 REE S Se a ee nT Sa ie A hits win nce oe 0 eet ck Cia ns ws 77 ems a tics scree stirs ct teks POPROIE RBUSOA . 6. 3. 81 Senne, TNO TAMER... 52 Salcedo 2 aoe SANS RRS . 81 ME ee sae oe Sat Le. A Tee PER TROON 5 cones 81 en pe Pte cee nace acess aah wees 81 ————e ee ae So ee a = I ia a Sea a Rta Nine em cya 6 LIFE HISTORY OF NORTH AMERICAN TICKS. Development of the Ixodide—Continued. Page. Genus Ixodes—Continued. iti’ The rotund tick, dzodes king? Bishopp-2- 42204 8) seas 2 ee ee ee 82 Descriptive. satis abe Syste oc See ne 82 Host:relationship,..): 2 cceqsdes sephegi ie See Tere ee ae ee 90 Geographical distributions. 22.252 24. 5) tes eae ee 90 Ethe Mistory 2c52 oji< aceisie ne eee Stee ee 90 . Tater ce yele ss Cou. ct cee Ose te ae eres ae 2 CaN ete ee 95 Hconomic importance: 0322.23.48 cee Sao ee ee 96 Natural. control: . 2 cc. 2. 5s So a ee ee 96 The bird tick, Hemaphysalis chordeilis Packard.................-- eS Tae 97 Descriptive. oo. .2o cle - Oo has ae Oe ee ee ee ee 97 Host relationship... 2... 222 s03.02t eee ee a ee ee 97 Geopraphical distribution: 222252522 oes eae eee 98 Tite history... s2 i) ooo SEAS Cees Ba a ee 98 Life cycle: s.. 352.2002 22r ese ee ee eee 101 Economic importances..\a.252 2 = 254d 22524 fy Ss) eee eee 101 Natural controls.....2082 2 Seer ee ee ee eee 102 Artificial comtroly'¢.4 4.4: S36 5425.9 Sch ee ee ee 102 Genus Rhipicephalus....:.25.9-22es5auc0 Gas ok ee eee eee 102 The brown dog tick, Rhipicephalus sanguineus (Latreille) (teranus Banks). secusees bec Soe bey eee Oe ee ee eee ee 102 Descripves. <5 eeee ace soe ee SEs ee ee ee eae 103 Host relationships... sce es 5.2 22 se oe eee eee eee 103 Georraphical distabutioni. 24.45.22 oo ee ae eee 104 Wate history: 5.2 yee ns 0 es eee ee a ne 105 Dafoe cyeleceo. «bee si 0s SNS Rs ene ray in nee ae 110 Beonomic importamee. 50 042 oo eee eke te Men nl ye oe 110 Natunaleontrolecsecc a: fe cute is Sas a het a een en eae 110 Antiietal control: 2. 5 lee coe Jabs bee & eee age re ee ea 111 Genus Marearopus ®. 22) bo 2 cee ee. cols oe rere ee ee een ene EE The North American cattle tick, Margaropus annulatus (Say). .....-- 11 Deseriptives: ..2cck oes Boe en ee eee 111 Host relationship’. :' 2. ee ee ee eae 2 Georeraphical distributions: -2225-222--5 252 satiate tary Bl fo on MSN 112 Life historys22 24 oe ee ea ee 112 Life cycles.) sc 4 oe SC ak Ee reece 116 Economic importance, natural control, and artificial control...... 117 The Australian cattle tick, Margaropus annulatus australis (Fuller)... 117 Deseriptives2.<.c..- ch Set Skis toPod. ho. Deg) eee ea 117 Host relationship: <5 5-06 aces oo ee ee eee 117 Geographical: distribution:-.. 22.55... 2 eee ts See ee 118 Life history. 20:0. 0.55249) 32) bin ae ee a el Pe eee) <2 118 Life cy Cleec 2.2528 oh 2 8 a ee ee ee oe 122 Biconomic importance jo.) oe eee eee oe 122 Natural control 2< =2 - 25.22 She eS: oe ee ee 122 Artificial control 3. c.22.5.0. ae ee ee ee eee eee 123 CONTENTS. Development of the Ixodiaze—Continued. mM eM ie ok eae sca w crt aeaetyecse ver ose ees The gopher-tortoise tick, Amblyomma tuberculatum Marx...........--. Ts oie onl abel Plo int ole pg lias he apg ag > elroy She iki ee ae eA aR eek Soi Belek ta cyt ty qe ae eae np Cppmemaemnican GismeIDINIIOR. 52535 ee eee keer enero eg daeuee Pi Be Coes ee eee SiS oS. Se eee re te ae 2a Pe ee ein a ae eae. ok eee ches «% mma MONTANE tice eae eae on le te te bee ks epee Cee eA bee fot er Pan a ee ad ne 2 Got ein 2 an o's ce The iguana tick, Amblyomma dissimile Koch.............-.--------- Loe er rh cee pepepaeeiete iil papi heh aco pela Melee ebe eieda ms sae Co ee Tn pad a ry Cele, SAL Gaal OO aca Oa ay Sey Geographical distribution...............-. Sls eile I hcp ERMC rae ees ee Ta ee ee eke. Ride | Bo os kia Si Bier eee eee eee) oper) Pees Se hy See ee See creme: SILLINONIIICE oy. 5 2 Se a So Se Se oo a bon doen ooo nse wu DEER ee ce eee Pe. Lares a eee oa ee CESS knw od a2 The Gulf Coast tick, Amblyomma maculatum Koch............------ nem ee eo te Oe PLS WA TIES NE ON See Oe Soe, jE a 8 5 1 amie al pt ee a aia Bees pgs ge gre eeerimeesr dist paton > ras OS Soe eee SO ee ee SLE ET kal ee ll ops cle 9 a iN a els ae tame ryrie... ote Ee RTS Oe Ce Rae eee ern MUADOTEAINCE 2 soa gh ee aS re see es 2 os ele ak DEMIR CEMOE 2 20 CR on ne we eee Sieh. “See ee re yee The lone star tick, Amblyomma americanum (Linneus)..-...--..----- CETL oS Latches rental ate alae: catia atag anh 1 lg pies ae on gig a Host relationship. ....... Piece ahah Prat 4 hae Le rel ye iene ae Peemersoiscee Cintriuiiitr 9 2 829 Bo ES ee TON Ee CT Oa Me aE At A At eal Me Gy Si Cael ee ae tn ae BS See vee ee CE Pee EL We. eee CEEIIMOPR go 2. Mes eee eee ec bak PieeR ONENESS tree Let, ink nd Spaeth tees Seat Oe The cayenne tick, Amblyomma cajennense (Fabricius).........---..-- CLP TRI ha hal a Shan SR el SS cal tel Reg nee OE SAT RIL alk i Ral d aet AA lil nie SN Pie ga age aa Seer irae aPLEIEEIO a en et Se f22 Sess ii kesks (TEESE SIS GS ag Tela acta: Pee 9 elie asl ie method Se ale Saar Ca eee Rae re ns ws agian se se Gee aes ee MUGGING SOR DOTUBMOE oon pice oo 6 = cee wees wea eee ese Bee. nats iN Se es eg Sk a ce aime mimic ade SIU CRIIEIRT OR Mts Diet he a wseeeeanceesinn one I i ek ee ew ate weainmieibics The rabbit Dermacentor, Dermacentor parumapertus marginatus Banks. IPT erie kb we a sare ih tid glk oe welsh sain mameenne® Host relationship. .......... Pe A On eee eee tall «aad PMR ep INMsOIN. . 525. hoo chad. Gh SIE LIFE HISTORY OF NORTH AMERICAN TICKS. The North American cattle tick, Margaropus annulatus: Fig. 1.— ‘“Deposited-out” female with eggs. Fig. 2.—Unengorged larva. Page. Fig. 3—Engorged nymph, dorsal view. Fig. 4——Engorged nymph, ventral view. Fig. 5.—Unengorged nymph (balsam mount). Fig. 6.—Fully engorged female, dorsal view. Fig. 7.—Male, ventral view. Fig. 8.—Male, dorsal view. Fig. 9.— Unengorged female, dorsalview. Fig. 10.—Unengorged female, ventral view. Fig.11.—Fully engorged female, ventral view.. The gopher-tortoise tick, Amblyomma tuberculatum: Fig. 1.—Un- engorged larva. Fig. 2.—Unengorged nymph. Fig. 3.—En- gorged nymph. Fig. 4.—Engorged female, dorsal view. Fig. 5.—Females engorging on tortoise’s feet. Fig. 6.—Male, dorsal view. Fig. 7—Unengorged female, dorsal view. Fig. 8.—En- gorged female, ventral view ry . The iguana tick, Amblyomma dissimile: Fig. 1.—Unengorged larva. Fig.2.—Unengorgednymph (balsam mount). Fig.3.— Male, dorsal view. Fig. 4.—Unengorged female, dorsal view. Fig. 5.—Fully engorged female (alive). The Gulf Coast tick, Amblyomma maculatum: Fig. 6.—Unengorged larva. Fig. 7.— Unengorged nymph (balsam mount). Fig. 8.—Male, dorsal view. Fig. 9—Unengorged female, dorsal view. Fig. 10.— Eneorged: female, idorsallvaew. 2eooas ance ee. sec eee ee The lone star tick, Amblyommaamericanum: Fig. 1.—Unengorged larva. Fig. 2.—Unengorged female, dorsal view. Fig. 3.— Male (balsam mount). Fig. 4-—Unengorged nymph (balsam mount). Fig.5.—Male, dorsalview. Fig.6.—Engorged female, frontal view. Fig. 7.—Male, ventral view. Fig. 8.—Un- engorged female, dorsal view. Fig. 9.—Unengorged female, ventral view. Fig. 10.—Fully engorged female, dorsal view. . The Cayenne tick, Amblyomma cajennense: Fig. 1.—Unengorged larva. Fig. 2—Unengorged female, dorsal view. Fig. 3.— Male, dorsal view. Fig. 4.—Unengorged nymph (balsam mount). Fig. 5.—Male, ventral view. Fig. 6.—Unengorged female, ventral view. Fig. 7.—Engorged female, dorsal view. The Pacific Coast tick, Dermacentor occidentalis: Fig. 8.—Male, dorsal view. Fig.9.—Male, ventralview. Fig. 10.—Unengorged female, dorsal view. Fig. 11.—Unengorged female, ventral view. Fig. 12.—Fully engorged female, dorsal view...... Ct: The rabbit Dermacentor, Dermacentor parumapertus margin- atus: Fig. 1.—Unengorged larva. Fig. 2.—Unengorged nymph (balsam mount). Fig. 3.—‘‘Deposited-out” female, dorsal view. Fig.4.—Male, dorsalview. Fig.5.—Unengorged female, dorsal view. The tropical horse tick, Dermacentor nitens: Fig. 6.—Unengorged larva. Fig. 7—Unengorged nymph (bal- sam mount). Fig. 8—Engorged female, dorsal view. Fig. 9.—Male, dorsal view. Fig. 10.—Unengorged female, dorsal view. Fig. 11.—Engorged nymph, ventral view. Fig. 12.— Engorged nymph, dorsal view. Fig. 13.—Engorged female, ventral view. Fig. 14.—Male (balsam mount) .....-... Gemetee 112 124 132 144 152 160 ILLUSTRATIONS, Prate XIV. The Rocky Mountain spotted-fever tick, Dermacentor venustus: Fia. Fig. 1—‘‘Deposited-out” female with eggs. Fig. 2.—Unen- gorged larva. Fig. 3.—‘‘Deposited-out”’ female, frontal view. Fig. 4.—Engorged larva. Fig. 5.—Engorged female, dorsal view. Fig. 6.—Unengorged female, dorsal view. Fig. 7.— Male, dorsal view. Fig. 8.—Male, ventral view. Fig. 9.— Unengorged female, ventral view ........---.-----+--+------ XV. The American dog tick, Dermacentor variabilis: Fig. 1.—Unen- gorged larva. Fig. 2.—Unengorged nymph (balsam mount). Fig. 3.—Engorged nymph about to molt tofemale. Fig. 4.— Engorged nymph about to molt to male. Fig. 5.—Male, dorsal “view (Texas form). Fig. 6—Unengorged female, dorsal view. Fig. 7.—Male, dorsal view (Oregon form). Fig. 8.—Unengorged female, ventral view. Fig. 9.—Male, ventral view. Fig. 10.— Engorged female, dorsal view. Fig. 11.—Engorged female, RE I ee adh ao isia'n a age Sa wwe oid 2 TEXT FIGURES. . The fowl tick, Argas miniatus: Distribution in the United States and earns ae es es PO a eas kaa a cia debe es . The spinose ear tick, Ornithodoros megnini: Distribution............. . The black-legged tick, Ivodes scapularis: Distribution in the United SOE RT sar el neo SWE Rit saat aD a od ier a ta eC ie . The rotund tick, Jvodes kingi: Distribution in the United States . . . The rabbit tick, Hemaphysalis leporis-palustris: Distribution in the SE SE Swe ile ded Slo et Se ies 1 Baa oe eS po . The brown dog tick, Rhipicephalus sanguineus: Distribution in the de a ath a. 5 . The North American cattle tick, Margaropus annulatus: Distribution Ee ER AO eat hancacnccmiceeanaecawe . The gopher-tortoise tick, Amblyomma tuberculatum: Distribution... . . The gopher-tortoise tick: Vesica biloba...................--.-.-.-- . The Gulf Coast tick, Amblyomma maculatum: Distribution in the en Se Ee ee at eS Saline Anke Warten . The lone star tick, Amblyomma americanum: Distribution in the ee ee BE aes a oe cn oe ne xl omnes oa S we wm . The Cayenne tick, Amblyomma cajennense: Distribution in North and nal A I Otte ub aed cba d monde ad~ ass iceumecmacacccs . The rabbit Dermacentor, Dermacentor parumapertus AAO Dis- ERE Sy Ey Sp rye = eee ee . The Rocky Mountain spotted-fever tick, Dermacentor venustus: Dis- Deremrneemn era ieer Nigsatert TATOO... a na ccc Gan cdacenenensmecace dim . The Pacific Coast tick, Dermacentor occidentalis: Distribution in the Me atte ie ia Cds, Do Us dies ncn oklebictGauseeschsonss . The American dog tick, Dermacentor variabilis: Distribution in the I eet aN hae bine o kmmbeeenieba . The tropical horse tick, Dermacentor nitens: Distribution in North ne TET WEE RIMS ee els aenkGuexeuedenuce 11 Page. 164 192 7 6 =e ee ee a ee 6 EE ——— es a eC THE LIFE HISTORY AND BIONOMICS OF SOME NORTH AMERICAN TICKS. INTRODUCTION, During the course of investigations of the biology of the North American fever or cattle tick, conducted by the Bureau of Entomology, with headquarters at the field laboratory at Dallas, Tex., many other species of ticks were met with, some of which are of considerable economic importance. In view of their importance as ectoparasites and the part that several are known to play in the transmission of disease, and in view of the further fact that comparatively little was known of their life history and bionomics, it was decided that the investigations should be extended to include as many of the species as possible. During 1905 a few notes were made upon ticks other than the cattle tick (Margaropus annulatus), which was at that time being studied. These miscellaneous notes were consolidated with those of 1906 during the winter of 1906-7 and published in Bulletin 72 of this bureau, together with the results of studies of the cattle tick. In 1907 fur- ther attention was given to those species which could be obtained for study. During the summer of that year valuable information was received from Prof. C. P. Lounsbury, who visited the laboratory. In 1908 the study of various species was taken up more extensively. The work was confined principally, however, to those species occur- ring in Texas. In 1910 an investigation of the Rocky Mountain spotted-fever tick (Dermacentor venustus) was undertaken by the bureau in cooperation with the Montana Agricultural Experiment Station and the Bureau of Biological Survey. Incidental to the work on the spotted-fever tick studies were undertaken on the life his- tories and habits of a number of species of ticks occurring in the western United States, and several species not herein discussed are at the present time being studied. Our knowledge of the life his- 13 aa. LIFE HISTORY OF NORTH AMERICAN TICKS. tories of some of these species is rather complete, but it seems desir- able to withhold from publication the results of investigations of these species until further data are accumulated. This tick investigation has been conducted, throughout, under the general direction of Mr. W. D. Hunter, from whom the writers have received many helpful suggestions. Mr. W. A. Hooker had direct charge of the investigation from its inception until September 15, 1908, when Mr. F. C. Bishopp took direct charge of the investiga- tion and has carried it forward. Mr. H. P. Wood’s connection with the work has been continuous since November, 1907. Mr. J. D. Mitchell and Mr. F. C. Pratt (now deceased) contributed valuable assistance by collecting material and making notes on host relations and economic status. Mr. W. V. King, acting under the direct super- vision of Prof. R. A. Cooley, of the Montana Agricultural Experiment Station, has, by the collection of several hundred lots of ticks in the Northwestern States, supplied a large amount of information on the hosts of Dermacentor venustus and other species and some information upon the geographical distribution of these species. Prof. Cooley has also aided materially in several ways, including the sending of a number of specimens of Ixodes kingi, which were used in the studies of that species published herein. Several correspondents, particularly in the Western States, have furnished a large number of lots of ticks which have been of much value in the work. Mr. C. E. Hood, Mr. G. N. Wolcott, and Mr. G. W. Hood are responsible for a few of the counts of eggs, and these gentlemen, as well as Mr. E. A. McGregor and Mr. J. Jacobs, have assisted in the summarization and tabulation of records.! CONCERNING TICKS IN GENERAL. SYSTEMATIC POSITION AND CLASSIFICATION. The ticks form a superfamily of the order Acarina, and are closely related to the mites which produce scab, mange, and itch. The superfamily Ixodoidea, to which all the ticks belong, is composed of two families, namely, the Argaside, represented in this country by 2 1 Credit for taking the photographs used in making the plates in this bulletin should be given as follows: H. P. Wood: Plate I, fig. 1. F.C. Bishopp: Plate I, fig. 6; Plate III, fig. 2; Plate XIV, fig. 5. W.E. Hinds: Plate II, fig. 1; Plate VI, fig. 10; Plate VII, figs. 1, 5, 6; Plate VIII, fig. 5; Plate XI, fig. 3; Plate XIII, fig. 14. j W. A. Hooker: Plate I, figs. 4, 5; Plate III, figs. 1, 3, 4; Plate V, figs. 1, 4; Plate VI, figs. 11, 12, 15; Plate VII, figs.2, 4; Plate VIII, figs.1, 2; Plate IX, figs. 1-8; Plate X, figs.1-10; Plate XI, figs. 1, 4, 5, 7, 8, 9; Plate XII, figs. 1, 4, 7; Plate XIII, figs. 1-13; Plate XIV, fig. 2; Plate XV, figs. 1-4, 10, 11. G.N. Wolcott: Plate I, figs. 2,3; Plate II, fig. 2; Plate III, figs. 5-8; Plate IV, figs. 1,9; Plate V, figs. 2,3, 5-8; Plate VI, figs. 1-9, 13, 14, 16,17; Plate VII, figs. 3, 7-10; Plate VIII, figs. 3, 4, 7-11; Plate XI, figs. 6, 10; Plate XII, figs. 2, 3, 5, 6, 8-12; Plate XIV, figs. 1, 3, 4, 6-9; Plate XV, figs. 5-9, SYSTEMATIC POSITION AND CLASSIFICATION. 15 genera and 6 (possibly 7) species, and the Ixodide, represented by 8 genera, 36 species, and 3 varieties, as follows: Argas (2 species in the United States). Ornithodoros (4 species in the United States). Rhipicephalus (1 species in the United States). Margaropus(1species in the United States). Rhipicephalinz; Hemaphysalis (2 species in the United States). Dermacentor (8 species and 1 variety in the United States). Ixodes (16 species and 2 varieties in the United States). Ceratixodes (2 species in the United Teodre:.. a. States). Aponomma (1 speciesin the United States). Amblyomma (5 species in the United States). Argasidze | Ixodoidea... Ixodide .. The type locality of 12 of the recognized species occurring in this country is outside of the United States. Of these 12 species 2 were described by Linneeus (1758), 2 by Fabricius (1794), 1 by Leach (1815), 3 by Koch (1844), 1 by Guérin (1849), 1 by Cambridge (1876), and 2 by Dugés (1883). Many of the names of supposedly new species have proved to be synonyms. Nymphs have occasionally been described as new species. The sexual dimorphism and the variation in the amount of blood en- gorged at the time the specimens were collected account for many of the synonyms. Of the 8 species described by Say in 1821 all but 3 are synonyms or remain unrecognized, while of the 5 described by Fitch (1872) all are synonyms or remain unrecognized. Of the 9 described by kerri (1868, 1869, 1872) 5 are recognized as distinct species. The first work upon iis classification of the ticks of this country was that of Dr. Marx (1893), whose studies were soon after ter- minated by his death. In 1896 Dr. L. G. Neumann published the first part of his revision of the Ixodoidea of the world, the last part of which appeared in 1901, but which has been followed from year to year by a series of ‘*Notes.”” Unfortunately this valuable work, in its original form, is accessible to only a few. However, a concise summary has recently been published (1911). The first great step in the classification of the ticks of thiscountry was taken in 1901, when the important work by Drs. Salmon and Stiles, on the cattle ticks of the United States, was published. With the appearance, in 1908, of the revision of the Ixodoidea of the United 16 LIFE HISTORY OF NORTH AMERICAN TICKS. States, by Mr. Nathan Banks, a work became available by means of which most of the ticks of this country can be readily identified. Since this work appeared several new species have been described by Messrs. Banks (1910), Stiles (1910), and Bishopp (1911b),?so that at present 42 species and 3 varieties representing the genera Argas, Ornithodoros, Amblyomma, Aponomma, Ceratixodes, Dermacentor, Hemaphysalis, Ixodes, Margaropus, and Rhipicephalus, are known to occur within our borders. A valuable monograph of the Ixodoidea is being published by Nuttall, Warburton, Cooper, and Robinson. Two parts, dealing with the Argasidae, the classification of ticks, and the genus Ixodes, have been issued. A detailed illustrated account of the general structure of ticks, to which reference should be made, has been given by Salmon and Stiles (1901, pp. 387-398). It may be well, however, to give a brief descrip- tion of the so-called capitulum or head, bearing the palpi and the haustellum, the latter of which consists of the mandibles, mandibular sheaths, and hypostome, which are inserted into the skin of the host. The capitulum is a small subtriangular piece that articulates with the anterior margin and usually within a slight emargination of the corneous shield, or scutum, which in the ixodid female forms the front part of the dorsum and in the male covers the greater part of the body. The hypostome, or labium, which lies underneath the mandibles, is an elongated dart or spatulate structure, which is com- posed of 2 lateral symmetrical halves bearing many hooks or denticles directed backward, so that when embedded in the flesh it can not be forcibly withdrawn with ease. The mandibles, 2 in number, are terminated anteriorly by either 2 or 3 processes, known as apophyses, that are used for piercing the flesh and making an entrance. The mandibular sheaths surround the base of the mandibles and extend backward on the dorsal side of the haustellum. Upon each side of the haustellum, applied closely thereto, are the palpi, which are grooved on the inner margin. There is considerable variation in the size of the individual ticks of a single species, among both males and females. The engorged nymphs also vary in size, the larger usually being prefemales.? Under the several species measurements will be found which indicate the variation. The color, especially of the immature stages, is also quite variable. In engorged larve and nymphs of some species there is a variation from dark brown to pink and pale gray. This is apparently 1 For the classification of the ticks the worker is referred to Mr. Banks’s Revision of the Ixodoidea (1908). Descriptions of adults, aside from their size and coloration, have been omitted in the present work, as they will be found in Mr. Banks’s Revision. 2 See Bibliographical references, p. 205. 3 The terms ‘‘premale” and “‘prefemale” have been used to denote those individuals, not yet molted to adults, in which the sew can be recognized, TS eS LLCU eS a a ee ee ar, ee ee ae eee eee ee. ee ee ee ~~ ere =a SCC. COLLECTING, PRESERVING, AND MOUNTING. 17 due to the proportionate amount of blood, lymph, and inflammatory exudate that has been engorged. The internal anatomy of ticks has been studied by a number of investigators, among whom mention should be made of Heller (1858), Pagenstecher (1861), Williams (1905), Allen (1905), Christophers (1906), Bonnet (1907), Nordenskidld (1908, 1909), and Samson (1909). COLLECTING, PRESERVING, AND MOUNTING. Owing to the fact that very few entomologists or zoologists have had experience with the systematic collection of ticks a few sugges- tions along this line may prove of value. The writers have found that in many instances zoologists have received and handled speci- mens of skins and living animals to which ticks were attached and entirely overlooked these parasites. In other cases the ticks were seen but were not preserved.. The ease with which this class of para- sites may be preserved should encourage zoologists to keep on the lookout for them and to collect all specimens seen. Persons who collect specimens of ticks should record the host, point of attachment, date, and locality. All parts of the host, including the inside of the ears, should be closely examined. Upon the discovery of a female, and before removing it, the collector should search closely for the male, which may be attached near by. A pair of forceps will be found useful in removing the smaller ticks. Those with short mouthparts are readily removed without injury, but many, particularly those of the genus Ixodes, are usually so firmly attached that the body of the tick will be separated from the capitulum unless the latter is firmly grasped. Some ixo- dologists have recommended the application of a penetrating oil and waiting for the tick to loosen its hold, but this will seldom be found necessary. In collecting ticks from small animals which have been shot or trapped, a supply of small cotton bags should be at hand into which the host can be placed as soon as shot and the bag firmly tied to pre- vent the escape of the ticks. In this way specimens may be exam- ined at the collector’s convenience and notes may be made on the habits of the ticks. If the host animal is too large to be bagged it should be examined at once over some white surface, such as white cotton cloth spread over the ground. The importance of immediate examination is emphasized, as the writers have found that larve of the rabbit tick and of other species with short hypostomes commence to leave the host within a very few minutes after the animal is killed. Ticks with long hypostomes, such as Ixodes, are sometimes unable to detach, and therefore remain upon the animal. In a number of 21448°—Bull. 106—12—2 18 LIFE HISTORY OF NORTH AMERICAN TICKS. instances living and dead ticks have been found clinging to the skins of animals which have been nailed on a wall for several days. The collection of ticks from herbage, on which they are awaiting a host, may best be done by dragging a white cloth, preferably of wool, over bushes, grass, etc. An ordinary insect beating net may also be employed. Fruitful results have been found to attend the examina- tion of the dens of animals, nests and regular roosting places of birds, and the ground in the vicinity of resting and watering places of mammals. Pill boxes have been found to be satisfactory receptacles for the ticks when collected. In preserving we have usually used 80 per cent alcohol or a mixture consisting of 60 parts alcohol, 1 part formalin, and 39 parts water. Adult specimens, particularly males with bright color markings, should be mounted on pins, as well as preserved in alcohol. For microscopical study specimens should be mounted in Canada balsam on slides. The contents of the body should first be teased out in hot water, through a slit made at the posterior end of the body. Speci- mens can then be readily cleared by boiling in a 10 per cent solution of caustic potash (KOH), care being taken that the clearing be not carried too far. The method employed by Dr. C. W. Stiles consists in soaking the specimens in from 2 to 5 per cent caustic-potash solution for periods varying from 12 to 96 hours, after which all of the soft body content is removed, and after passing the specimens through water, the alcohols, and xylol or other clearing agents, the specimens are mounted in balsam. ECONOMIC IMPORTANCE. Ticks are of economic importance (1) as agents in the transmission of infectious diseases, and (2) as external parasites, both of man and the lower animals. At least two distinct diseases of man and eight or more of domestic animals are known to be thus transmitted, at least 17 species of ticks bemg involved as carriers. Of these dis- eases one of man and one of cattle occur, and one of fowls is sus- pected to occur, in the United States, while several others would undoubtedly obtain a foothold were they once introduced. It has been estimated by Dr. Mohler (1905) that the cattle tick alone is the source of approximately $40,000,000 annual loss in the United States. Mayer (1906) has estimated the annual loss as nearly $100,000,000. : These parasites are of considerable importance as external parasites, particularly in the Southern States, owing not only to their irritation and great drain upon stock through removal of blood, but also to their indirect effect as well. In one of Theiler’s experiments (1909a) a horse that was infested with Margaropus decoloratus died as a result of infestation from acute anemia due to the withdrawal of blood. ECONOMIC IMPORTANCE, 19 Within 3 days 14 pounds, by weight, of ticks which had dropped from this horse were collected, and this amount represented only about one-half of the total number of ticks which engorged upon it. After dropping, their places of attachment furnish points at which the screw-worm fly (Chrysomyia macellaria) deposits its eggs, the mag- gots from which then readily enter the host. In the Southwestern States the appearance of equines is frequently injured by screw- worms, which gain entrance at the points in the ears where ticks had been attached, burrow, and destroy the supporting cartilage, causing the ears to lop over. This condition is commonly known as ‘‘gotched.”’ Not the least of all is the frequent annoyance which man suffers as the result of the attachment of ticks to his body. The ticks which molt upon the host, instead of having to wait long periods to find another, merely continue sucking blood from the same animal. As a result these ticks reproduce very rapidly and frequently become of much greater importance as external parasites than species which drop to molt. This is the case with the cattle tick. Those which drop to molt have overcome this great disadvantage by becoming more resistant to heat and cold and by having gained the power to withstand much longer periods of fasting. Certain members of the subfamily Ixodine, while not occurring in such great numbers on animals as in the case of species which molt on their hosts—all of which belong to the subfamily Rhipicephaline—are frequently of considerable importance as pests, owing to the fact that the great length of the hypostome permits deep penetration. As the result of this deep penetration by the [xodine, an inflammation is produced which frequently results in suppuration. Often in the attempt to remove ticks belonging to this latter class from the body of the host, the capitulum is separated from the body of the tick and remains embedded in the host. The periods in the life history of ticks of particular importance economically and which should be determined are: Longevity, or the period required forstarvation while awaiting a host; minimum parasitic period, which is used in connection with the preoviposition and incu- bation periods to determine the time that tick-free areas may be used after infested cattle are turned in before the areas become infested; maximum parasitic pervod, or the period required for cleaning the host of all ticks (except males) when kept in tick-free inclosures; preovi- position period and minimum incubation period, used with the mini- mum parasitic period to determine the time that tick-free lots may be used before becoming infested; stage or stages of imbibition of infec- tion and the stage or stages in which infection is transmitted, i. e., in the case of species involved in disease transmission. °0 LIFE HISTORY OF NORTH AMERICAN TICKS. HISTORY OF THE BIOLOGICAL STUDY OF TICKS. The first studies made of the life history and habits of ticks were those of Dr. Cooper Curtice (1891, 1892a, b) on the cattle tick (Margaropus annulatus) conducted in cooperation with the Texas Agricultural Experiment Station at about the time that Drs. Smith and Kilborne were investigating the réle of that species in the trans- mission of splenetic fever. From the time of these investigations by Dr. Curtice up to 1898 little attention seems to have been given to the biology of ticks other than this species, although a prelimi- nary study was made of Amblyomma variegatum (Hyalomma venustum) in Antigua, by C. A. Barber (1894-95). In 1898 Dixon and Spreull reported studies made of Margaropus decoloratus and the same year Prof. C. P. Lounsbury, the entomologist of Cape Colony, British South Africa, began his classic studies of the Ixodoidea. Since that time Lounsbury has worked out the life history and habits of a large number of South African species. During the course of these studies he has demonstrated the trans- mission by ticks of three distinct diseases of domestic animals and the pathogenicity of at least seven species of ticks and has determined the stages of imbibition and of inoculation of the hosts with the disease-producing organisms. In 1898 Dalrymple, Morgan, and Dodson, of the Louisiana Agricul- tural Experiment Station, published a detailed account of experi- ments relating to the life history of the cattle tick. The information which they furnished upon the longevity of the ‘‘seed ticks” served as a basis for the feed-lot and pasture-rotation system for cleaning stock and pastures of the cattle tick. In 1899 Prof. H. A. Morgan published further information upon the life history of the cattle tick and included data upon the biology of Amblyomma americanum, Dermacentor variabilis (electus), and Ixodes scapularis (not ricinus). In 1899 E. G. Wheler, in England, published an account in which : he reported studies made of Ixodes ricinus and the same year C. J. Pound published notes on the Australian cattle tick (Margaropus annulatus australis). In. 1903, Dr. H. Kossel and his coworkers pub- lished an account of studies of the biology of Jxodes ricinus in a report of investigations made in Germany in which they found it to transmit bovine piroplasmosis. During the course of his investigations of the various protozoan diseases of animals in South Africa, Dr. Arnold Theiler, veterinary bacteriologist to the Transvaal, has added much to our general knowledge of the biology of ticks. Dr. H. T. Ricketts, in connection with his investigation of Rocky Mountain spotted fever, has published information from time to time (1907, 1909a, b) upon the life history and habits of Dermacentor venustus. Prof. R.A. Cooley (1908, 1911) and W. D. Hunter and F. C. Bishopp (1911a, b) GEOGRAPHICAL DISTRIBUTION. 91 have also published studies made of that species. Capt. S. R. Christophers, working in India, has studied Rhipicephalus sanguineus (1907) and made observations on Ornithodoros savignyi (1906). In South America, Dr. F. Lahille, of Argentina (1905), has studied Margaropus annulatus australis and Dr. C. J. Rohr, of Brazil (1909), has conducted extensive investigations on six species. Newstead reported in 1909 on studies made in Jamaica. Among others who have studied the cattle tick are Ransom (1906), Newell and Dough- erty (1906), Schroeder (1907), Hunter and Hooker (1907), Cary (1908), Cotton (1908), and Graybill (1911). Many others have recorded mis- cellaneous observations on different species. Preliminary notes on several species were published by Hunter and Hooker in 1907, and further information has since been published by Hooker (1908, 1909a, b,c). GEOGRAPHICAL DISTRIBUTION. The ticks that commonly attach to the domestic animals have been widely disseminated on the hosts so that now many of them are only limited by climatic zones. Of the several meteorologic factors which control their distribution cold appears to be the most impor- tant, although excessive heat and variation in humidity are also important. Thus in America we find the tick species most abundant in the Tropical and Lower Austral zones and the least so in the Boreal Zone. Several native species, notably Margaropus annulatus, Amblyomma maculatum, and Argas miniatus rarely appear above the Lower Austral Zone, while Amblyomma tuberculatum, A. dissimile, A. cajennense, and Dermacentor nitens occur only in the Tropical or Gulf strips of the Lower Austral Zone. Our two species of Ceratixodes, namely, putus and signatus, apparently occur in the Boreal Zone only. Some species remain attached to their hosts for long periods (par- - ticularly the males) and may be carried great distances. This has resulted in numerous records which must be rejected in determining the normal distribution of a species. The approximate zonal dis- tribution of our species is shown in Table I. 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