RICAN IXUDOIDE VOLUME | - TICKS OF THE SUDAN BY HARRY HOOGSTRAAL S,GAMBIA PORTUGUESE *” GUINEA r~ ASIERRA eo LEONE, “% ETHIOPIA COAST La S¢, *, > \\ v KENYA RUANDA URUNDI NORTHERN RHODESIA SOUTHERN RHODESIA ECHL eas UNION SOUTH G2 WEST : AFRICA OF {> SOUTH AFRICA Wvunem HAAN AFRICAN IXODOIDEA I. TICKS OF THE SUDAN (WITH SPECIAL REFERENCE TO EQUATORIA PROVINCE AND WITH PRELIMINARY REVIEWS OF THE GENERA BOOPHILUS, MARGAROPUS, AND HYALGMMA) Harry Hoogstraal Head, Department of Medical Zoology U. S. Naval Medical Research Unit No. 3 Cairo, Egypt Fee ee OT ae Research Report NM 005 050.29.07 Date of Publication, 1956 With 372 Figures on 103 Plates The opinions or assertions contained herein are the private ones of the writer and are not to be construed as official or reflecting the views of the Navy Department or the naval service at large. I TO COLONEL WILLARD VAN ORSDEL KING, PH.D. A VETERAN MEDICAL ENTOMOLOGIST AND PIONEER WORKER ON TICK VECTORS OF ROCKY MOUNTAIN SPOTTED FEVER. FROM WHOM I HAVE LEARNED MUCH OF THE METHODS OF SCIENTIFIC INQUIRY. HEIL EEL IV TABLE OF CONTENTS Page DEDICATION TAPE EOP ICONDLENTS..,crcis cis.seecremerre wealewiiectes sane Beleosteise el MAPS AND ITLUSTRATIONS............ porte are Seer a niaven :s.cisiare 8 se lor Namopoudatiaserecce.teccccccece sloleteicloisietololeislelevetete 12 DIE CV aia OMS efelalstelenerererereie ereveversietete’ « pieloleveloicicioisleloleleleie\oioios 12 FOREWORD, BY DR. C. B. PHILIP Sieteieforcl clo ierererevo otealepeteiene Stayekere)= 13 PNTRODUGC TION svesctereioisielare ¢ Sobocga0e elotolesoKeteloloiel cfereyclereialeielererete 15 Object and scope of this and related projects. joddoooc 15 History Of uthis. pro jeCiGnrcem «ieccen cis ne eieceiefotete re 6.4 0 sisteser lO ACKNOWIGEDGEMENTS 0.5) oinisiais)s\c)s's\ nish aisjajeleiesetats aie fioooocddoooooas J) LISTS OF TICKS REPORTED FROM THE SUDAN......... Alolb Good 26 pecies and subspecies records by Province....... aisfeyerni 20 Family Argasidaec.....csssee. SOTO MICs Bieiiatetoveleia epetene 26 Family EXOGTCaS . o:s.sleeisieterstarelecieieien ote Sdeadcoasong00 5 CU Species attributed in error to the Sudan......... seletete OE Previously reported names used in error, changed, or synonymized........e. SOIC ITYROOO OOO OO AON pioievers VOLE Names herein changed, synonymized, or proposed. shsloloke . 30 Keys to families and genera of adults.........ceeeeees 39 Keys to developmental stages.......... Starets gs sieveisieisie seen LZ SEEC LEO PREC ORDO. ic aioyaierselataetateinte sietetatee Biotelefefeteteiorciciers SAgegna 45) Manner of data presentation..........seeeess efoleielsleleielors 43 Equatoria Province records.....-.sscceee odddddcbados 43 Distribution in the Sudan............. olslovsloiclolers S060 (vb DUST UU A OMeketotenetaiolateieiciekeiaierfolareleteleteleterelersleleiolelsleleterelelere 45 HO Sie Shetererekeneletorstevelelclersiere so oCoD od GDO0D0OGDG0G0000000000 HH) Bil Oil Opaygayerssaretet share e.nictetere wiete’s.« S Seite ct enh ee-ceisecincies GO Disease relations. .c.. exsleyskoke okexetete Gees SSeS sfaveteraysysyeln/sO RROMATICS a toloreielelevereicleiere Slererelohe semctetste ss loffeletetert sie aire sletosveres GO TASTU AA CAC OMA imtroduetlon.camce. seen coe sioteveraserotetersyels SOD COCOOOUOSEE: Key to Sudan species of Rhipicephaluss....sssesse0. 586 Rhipicephalus, appends culllacus es ohare crac c « . 600 Rhipicephalus EINOMA ec SRN earerctine Meer terns «ae OL Rhipicephalus bequaerti....... aXayete” ofovototeveterereetorve 06.6 OL Rhipicephalus compositus........ Nev eteNereKaverarerartere Berean OL3 Rhi Heise Cis pidaclic os. “ieee rae eee x 5 oleBul ice us Rhipl Gephatius. 7OuStINebUS .... sacecee rc cdewerotcne ccs 636 Rhipiceph LCE pPhalUS \SVERUSIS CVETIUS Iter olererer otolsrcicverereete' so « cOL2 Rhipicephalus Koc hile. - ne etree Soe oto 6.69055 Rh picepialuselonclcoxatus. srerrimns moon... 661 Rhtpreephaius: loncushe se cc. eee eee Bae IAS O15) Rhi picephalus Sie tn te ee aronelevers eieiee OVO Rhipicephalus PSREAVUISIe 'aoreversyetereretero srotololeretsyarererctevete »-s.0 OTK, Rhipicephalus Sanguineus sanguineus.......e.++.+++- O80 Rhipicephalus SUMP SOM stare alatcrereeie alaloverarertiareve ciereve 6 « « 120 Rhipicephalus SIMUSWSIUMUS. . «cer aVelavevehe- eteware Bra toteKers S60 (8 s 5s Page Rhipicephalus sims SenegalensisS....ecceeeeeee 154 Rhi ice us SUICELUS Tc cere rere ce ET ice us SUPCTETItUS....--+eeeeereeeeee ee 168 Rhipicephalus TLLCUSPIS..ccccccccccccccccccees IZ ce Rhipi phalus SPiers otorstorotare/ossionerers/elsieversinieieiesieleleicle 110 V ANIMALS AND OTHER SOURCES OF TICKS... ...ccccccccccccce ed 19 UALOLUABETO VANCE amtcteinclersiele cleleciersielela cisievetoiorereheielsiersiererhOoe Bahre nin Ghazale Exo valine Clerteretesisieelersioreie ciecieisiersisimcieieienin OS Upper’ Nites Provinces. ss sss e.c.ca(e.cels « ¢ crelsieiiociatelelsene cl? NarLuUreeLOVANCO cicvererelereisisierereicteleteots cies ie cee eee TOOL) Kordohant ProvanCecwrssicieec etic et cet ene come comet oLe. BilweyNalenProvalne Sercreiciecsiesice ne sieeicie cietac me einiersicie eines KassallemProvanCe's ss sfs'sie o.oo o's b mrerevonareteneversne srevevererae re mie eROLO KHALCOUMMETOVLNC EC stele te oc else ol erelotaicverstersbetevers ororererevensiereversieieocl Northern Broviince nc cic.e siele os ene elere eisteatonste eioeerereminreiciemieOce VI CLIMATIC AND BIOTIC FEATURES OF THE SUDAN..............931 RETIN CO Wle ie terecere creteletorera eiore:n eretavercraFanarerorentretare ievavevevelerarcielereicinool: PioralRDUStLOPCUS es ceiesre ererecrseeine es tiene te ec ceererotrelrleemree Ooo) KavnalgDictrctSmand ArOaS srs ctejerereeicici cele cleisiclercielersreisiOle el CKD USL TIL Gl OMlonelerererererecicie ere orereraie rere eieleie oiereicielelelsitere occa VIl INP ERIN ENG Naveretecetescccere tere tere ere rere ere eee ne ee SETILIDIETSTeTe Oe. Ar as TSU CUS orate eters SNe Cree hee on Deen oa ee ee OOe. Argas ref Lexus, ASD Edi cis c.c-ojo 8 cuscee savers wie were eretereeeOOs Ornithodoros annulatus, 0. capensis, 0. foleyi.......864 thodoros lahorensis, 0. moubata.......sceeeecees S04 Ornithodoros savignyi, O. tholozani......sesecceeeee S07 Amblyomma cohaerens, A. lepidum, A. marmoreum (group)867 Amblyomma nuttalli, A. sum, A. rhinocerotis.....868 AmbL VOrma thOLLONL, A. VabLOsatullscciee soweaice «. «+2608 Aponomma exornatum, A. Tatum........cccccccsccccee es 809 Boophilus annulatus, B. decoloratus.........e++s+0+ 869 Boophilus MicTOplus...ssescccscccccscsessssscceceseesdl0 Dermacentor c. circumguttatus, D. rhinocerinus.......870 Haemaphysalis Somes E. Noodi shoodianwaerew > os ncor0 Haemaphysalis houy 0,0,0 e[e)0 6 eCS OTS 0 SUS Ce ST Sere coos coll Haemaphysalis leachii leachii, H. leachii mhsami....871 Hyalomma detritum and H. scupens pe naERS Seen a ae cy All Hyalomma Gromedaril .. wccdsacweue. cles eeen. ooee O70 Gra Page Hy a@loMmma, GXCaV.Atlile cts clcloisteic.c 6 o1te ssicie sieleicicle wc ele cis'eicieissOLO omma Sp. no. L near excavatum.....ccccccccccccess SOL Hyalomma sp. no. 2 near GxCAVALUM. .ceeeeecee cece cee e 0005 Hyalommasmarginatums snc tiee Res eeesu.cw Sladisle os ob ccccece 887 SIGE SOO es ONMARETUNC ALUMelsis cickeis slot sk + s2Gislesforles s © cle oe.c 0 0 0 8090 KOCOS PAINT See tee as 555255 sib sh sts ceeee te sees Coe xodes rasus, .. SChilLingSi.... cc cece eee eee e cence ee OI Margaro FOUTS PNAMOUMESE Toe ci ce Seb o Mess oo0es as 2090 Marge aro WINLHOMIGE EEE SAREE Se eee Dee eek esse Sc 899 ee ee MAVCMTBLUSs +555 5 sss Sees oe see ce bes oe G00 ice us com Situs, R. cuspidatus............... 908 Rhiplcephalus AiStimctus....ccccccecccces wooeeec sees HS Rhipicephalus 8. evertsi, R. longus, R. mblensi......909 Rhipicephalus pravus, R. S- SANZUINEUS.. 22... eee eee ee WOD ice us Simpsoni...... Pe iaoisiicisicrecicleloielereeleisiersini oO. Rhipicephalus S. sims, R. s. senegalensis............911 Rhipicepnalus ipicep Supertri tus, R. tricuspis........ oocnqo0edbl BIBLIOGRAPHY...... BOOS OOOO MICO OOIOOOICOOCIOCIIOOICOC eee cccecee 912 13-14 15.26 27-30 FI 9e 33-34 35-36 37-38 MAPS _AND_ ILLUSTRATIONS SUBJECT PAGE Subdivisions of Ethiopian Faunal Region...... 22 Polatical Divisionsvof Africacicsciselsicicincisencs Cr The Sudan. Principal Collecting Sites....... 25 Ixodidae. Key Morphological Characters...... 38 ALR GuAus. Jo0eAre Key Morphological Characters (0. savignyi)... 50 Key Morphological Characters (0. savigny Nee Gal ARGAS Key Morphological CharacterS...cccccccccccees D4 Peripheral and Lateral Differences........... 57 A. (Argas) poricus and A. (Argas) reflexus.. 58 i. Orcadenus brum Lei ERS eC NERO 81 k. (thie POLAT CES) MEOUC biiciclererciclolerorsioverersiarevoree ag O . (Chiropterargas) COMPUSUS...ccccccccecceee OT mt k. (Carios) VeSpertilionis....sssseceseeee ee 103 ORN| THODOROS MOUBATA Maite wand i Bemal.e'd.)./ciclcl cislole/aletelele a\orelalero\elayoreiereieiat ting Dilshrl DubsOM'csie <:cleveleloicterelelovolsic/ sere’ sieieis| oiersicioseye oi 0 Spermatophone cic aisieis «ia'e c/sic's'vlels e\e'sielolelciaisicioisieleintoo man MaxvAleicierclel al sreleletolotevelaleverovere/eleleteratefaleloleeielG Tarrvad-—Nympliaili MOL taleric/cials ls clals wiolelslalete sisieleieiciel go Q Internal Organs....ceseseeesecceeccceseece eld? 2 Sagittal Views, Unengorged and Engorging...158 apitulum. | Vertical SectloOn. Jebel Borkoly \ ¢ Letti Bosin + Berber KASSALA ie° Atboroe Ed Domer Kororo Tae 7 o Goz Reqebs KHARTOUM.” Wod Roiyo*Hoshe < Sennor ‘KOR DOF AN’ +E! Obes exbureey ‘ Loke Ros Amer* Gebelein Wod el Noi / /NILE\ Zor f”_| *Nyolo *Muhagoriyo \ eDetomi > Ler Renk Roseres | \ Koolib Hillss *Umm Berembeito } ‘ * "Idd e| Ghonom * Jebel Umm Dono ‘ *Heibon Nubo Mise = Jebel Tobuli =, “Polo Kone Tabongo* * *Sofono-.5 oper ws Ne Pethiiceowoe sh) = Goluol? Nyong Forest Gogrial* Mesnra el Rea CP Neue ee ne oa bal” BARR” "EL GHAZAL. a \ F E-T, A 10: 'P ‘ : + i {Gas UPPER. \NILE | uk Fodiot Akobo Post Sioa sTon} < J *Molek )\ | A . / Loke Nyobers: * “La Sage <5 | sf Rumbeke — ju \ sPibor Post . \ : \ *. oS Yirol “lel ay eye Toll Post ) Tombes in See oTerokeko ‘A Es R smerids Jubosf Mongolia ( Kopoetor \ «Lirongu \ 'e, sYombio is *R jot, Bo TA eK Lo Be Tonite .xatire »Nogichot & [hee SEN ei Ss Kole porie “*Nimules. Figure 3 THE SUDAN. PROVINCES AND PRINCIPAL COLLHOTING SITES OF TICKS REPORTED HEREIN (For additional Equatoria Province collecting localities see Figures 321 and 322) PLATE III 5 25 = *uepng Worsy oureu STYY Jepun poder YsITz J *eTquuoTysenb pxooed 2 *uBpng WOT pLOoeL AJTTBOOT 9YTUTJEp YSATI x *spiooel meu pue peyst—tqnd ATsnotasid y *TeTsejuU sues uo (G7G6T Teetysd00y) ya10der *9O0UTAOIg WOLF poss YSITI + AreuTwtTeid 10yJ ydooxe Suepng WOLF pLOO9I YSITI yy x X X X X Xx re) 0 0 TAuSTAeS *O 0 fe) 0 ) 9) fy) 0 + % Byeqnol *O SOUYUOGOHLINUO 0 0 0 0 0 0 0 0 xx (*ds *¥) x X 0 0 0 0 0 0 + STUOTTT}A1edsea °¥ 0 0 0 0 0 0 0 0 xx SUXOTJOI *y x X X x x X x X x snotsied °¥ HH HH 0) (@) (@) (0) 0 (6) HH snsnjuoo *y 0 0 0 0 Xx X 0 0 + Tydumiq °¥ 0 ) fo) 0 0 ) 0 0 HH Tyenoq *y SvyouyV NUGHLYON WAOLUVH VIVSSY unduva Nvyaoquoy ere er ae eae VIUOLVADdE SUI dads SQW OU AONTAOUd aVQISVONV ATINVA ATONTAOWd AG SCHOOGU SaToadSans CNY Salouds NVGANS SHL WOYS OSLYOddY SHOIL 40 SLSI1 Jah 0 (6) + + aK xX xX x. Xx O O 0) O Hi (0) eH HH KH 0) O 0) ) (0) 0 J + J ) é + 0 ne + + 0 X ) ) O + x te xX Xx x 6) 0 ) 0 0) 0 (8) (0) xX O fe) 0) ) (e) (0) J 8) Rs ) ) 6) ) ) 0 0) O KH fo) ) 0) 0) Ry fe) O 0 # fe) ) 0) O ) ) ) 0) # 0 ) xX x xX x X X xX Q 6) 0) 0) @) @) (0) 0) HH SNYBIOTOOep *q snjetnuue °g SET 18 a0 Ong umyel °y unjsuioxe *y Suezesyoo °y VWWOATENWNY SAS ASS GIIN YIN ‘TWZVHOD NUGHLWON WAOLUVHM VIVSSYA UNUVd NVEOGUOX Gog uagan Ta wnyg ViNOLvNba Sa OadS nn nn er SSS SS ce lS ES SD S@w OW AONTAOUd SS aS SS SS SSS SS SA aVQOOXT ALINVA = 24s ) fe) ) fe) ) re) fe) ) x eyelred *H 0 He 0 0 0 aK 0 x Hx TuesyuMmM TTYORST °H - + + + Xx ne X x X TryoeeT TTyoweT *y 0 0 0 0 0 5 +3 +3 +3 Tog *H 0 0 0 0 0 0 0 xx xe Tpooy Tpooy *H 0 0 0 0 0 0 0 0 xx Tyrewnbeq °H 0 0 0 0 0 0 0 + X JEST Topo *y SITVSKHdAUVW@VH 0 ) 0 ) 0) 0 @) 0) 3 snutieoouTyd °q 0 0 0 0 0 0 0 0 HH snzeVpyNsUMIITO °d °q HOLNGDIVNUAC @TIN SIN ‘TWZ¥HO NUGHLWN WAOLUVH VIVSSV UVC NVGOQOY ang umaan Ta uve VINOLVNta SAL OGIS SCw OY AONT AOU aVQI@MxXI ATINVA = 280 0) 0 0 0 (0) 0 ) 0) HH STUOT[T}ACGSOA °T 0 0 0 O 0 0 0 0 or xoTduys xeTduys *T 0 0 0 0 0 0 0 0 #e TSSUTTL TOS °T 0 0 0 0 0 0 0 0 ie seToedsqns, snsei *T 6) 6) O O 6) O O HH HH STsuetqoifeu *T 0 0 0 0 0 0 0 0 HH sndTedtavo *T Se GO xen fe) O O HH Hi HH HH HH Hi umyeoundy °y HH HH HH HH HH RK KH HH HH sedtynt *y HH HH HH HH HH ) ) ) ) UMyeuTsre °H HH HH HK HH HH O ) ) O umsseaduy °y HH HK He HH HH ) @) fe) 0 imyeyTeduyT °H HH RH HH HK HH (0) 9) ) 0 TTrepeuoip *H or or Hx oe oe 0 0 0 0 UNzTIYSP *H HH HH KH HH HH fe) ) ) ) UNYBABOXS *H VWWNOTVAH STIN IN ‘TVZVHD NUSHLYON WAOLUVHY IVSSYH UNdEVd NYMO@OY ana waaan Ta wya VIuoLvnta Sal odds Sau Oa TONTAO Ud SVGI@X ATIWVA o Oe Ie) fel 6) © ©. (0o (i © oy eo © oO 12 © ji CG fiom o oS oF iO) © O° .o) (CO iH © on Ome So © i C1 oO: o> oO S (eo) oO je) (oe) oo ms NUAHIHON WNOLYVHY WIVSSVH Unduvd NvaOdHOY oO tas] io) oO Oo oO oO GON WIN TIWZWHD anld widdh Ia wnya VINOlvnta TsueTUMM SNSUCT SNYBXOOTSUOT TYoOH TSJIoAS TS}JISAS snypOUuTYSTp snyepTdsno sn} Tsoduoo Tateenbaq Tplousre snyeTno tpuedde a S 0 ie He dst orted Taos a TpTel “Wl SNdGCOUVODUYN Sa 04ds S@uOORY ANT AOUd aVGid0xI XW 0 0 0 0 0 fe) 0 0 ore TpnentTe *y Saaqoxl XIGNAddV *pepntouy JOU ere sjeytTeu usTeLOJ OF YISUBI} UT STBUTUB WOIy pue sUSpreD TeoTSOTOOY woly spzooey :qION @) fo) 0) 0) 0 ) 0) ) HH (eda) x) 0 0 0 0 0 0 ) Hit 0 STdsnotiy *y 0 0 ) 0 fo) 0 Hi 0 xx sn}tizzedns °y 0 0 0 0 0 0 0 #H 0 snyeoqns *y 0 0 0 0 0 0 0 xx “x S}sueTesoues SnUTS *y x 0 Xx + X me x x x SMTS SNUTS °Y 0 0 0 0 0 0 0 0 xx Tuosduts *y X X X X X X X X X Sneutmsues Sneutnsues °y 0 0 0 0 0 0 0 0 x snaead *Y €°9U05) 5 Saye dao a tee @UN SUN ‘WZVHOD NUGHLYON WAOLUVHT WIVSSYH uNgyd NVdOGUON anqq wmaan Ta unya VIuOLWNda cage cess —_———— SduOOdU AONTAOUd ————— aVCLdOXI XTINVA - 31- SPECIES ATTRIBUTED IN ERROR TO THE SUDAN CRNITHQDOROS MEGNINI (Duges) (now Otobius megnini). Balfour (1906) Stated: "Mrs. Broun .... recognized the spinose nymph of the ear tick, Ornithodorus Megnini from the Sudari*. There is little doubt that this was an erroneous identi fication. O. megnini, listed for the Sudan (King 1911), was considered an error by King (1926). Nevertheless, the possi. bility that this species may be introduced into the Sudan should be considered. A tick infesting the ears of cattle, it has been introduced into South Africa (Bedford 1925,1932B) from America. It has also established itself in Nyasaland (Wilson 1950B), Northern Rhodesia (Morris 1933), Southern Rhodesia (Jack 1942), southern Belgian Congo (Schoenaers 1950, 1951A), and on Madagascar (Biick 1948B, Hoogstraal 1953E). AMBLYOMMA HEBRAEUM Koch, 1&4. Listed for the Sudan (King 1911) Yater deleted (King 1926). This species does not occur in the Sudan. EM(NH) files contain records of A. hebraeum from the Sudan (H. H. King legit, 1911), but corresponding specimens cannot be located in the collection. A specimen tube in the same institution contains material of both A. variegatum and A. hebraeum and a label (in Hirst's handwriting) = ERELE GDS them as such. Another almost illegible label in the vial reads "Taufikia, Sudan, 1909, H. H. King". There is a remote chance that the A. hebraeum specimens were removed from imported cattle but it is more likely that these are South African specimens that were somehow mixed with Sudan specimens. Museum records show that the specimens were presented by members of the Committee for Entomological Research. A note by Nuttall in his collection logbook states that the A. variegatum-hebraeum of King (1911) is A. lepidun. APONOMMA LAEVE Neumann, 1899, Listed from the Sudan by King (1911 . This is a nonAfrican species name (Theiler 1954B). The species in the Sudan is actually A. latum (Koch, 1844). - 32- BOOPHILUS AUSTRALIS (Fuller, 1899). Listed from the Sudan by Bal. our (1911H) and King (1911). This name, a synonym of B. microplus (Canestrini, 1883), which is not known from the Sudan, probably refers to misidentified material of B. decoloratus. It is less likely that it refers to B. annulatus. HAEMAPHYSALIS CAICARATA Neumann, 1902, was reported from Roseires, “Blue Nile Province, by Neumann (1910A). From his description and figure it is evident that this material represents H. houyi Nuttall and Warburton, 1915 (Hoogstraal 1955D). King ) did not list H. calcarata in his reports on Sudan ticks, and, although he collected specimens of H. houyi, they had been identified as H. leachii. HAEMAPHYSALIS ERINACEI Pavesi, 1884, which was reported from the an (Hoogstraal 1954B) was later (1955C) deleted. This re- cord was due to an early erroneous identification. HYALOMMA AEGYPTIUM (Linnaeus, 1758). A11 Sudan Hyalomma ticks pre- viously have been lumped under this name by King (1911,1926), O'Farrell (1913A,B), and others. This species does not occur in the Sudan. RHIPICEPHALUS BURSA Canestrini and Fanzago, 1877. The report by Weber (1943), from 8700 feet elevation in the Imatong Mountains of Equatoria Province, is based on material (kindly loaned by Dr. J. Bequaert) that Dr. Theiler and I have found to be R. kochi. 5 RHIPICEPHALUS CAPENSIS Koch, 1844. I have been unable to find evi. dence to support Zumpt's (1942B) statement that this tick oc curs in the Sudan, - 33- PREVIOUSLY REPORTED NAMES USED IN ERROR, CHANGED , OR SYNONYMIZED . This is not a list of synonyms of names now used; these may be found in taxonomic papers cited throughout this report. It is rather a list of names that have appeared in earlier papers on Sudan ticks or for which the Sudan has been listed as the source, but that subsequently have been changed or synonymized under names appearing in the present report. ARGAS MINIATUS Koch, 1844, mentioned by Balfour (1906) from Khartoum, ~——is a synonym of Argas persicus (Oken, 1818). AMBLYOMMA PETERSI Karsch, 1878, is believed by Schulze (1932A) to be Synonymous With A. rhinocerotis (de Geer, 1778). King (1926) and almost all other previous authors have applied the name petersi to this species, Schulze's findings are tentatively accepted in the present work, though specialists are not in complete agreement. AMBLYOMMA WERNERI WERNERI Schulze, 1932A, said to originate from ordofan, is synonymized under A. nuttalli Donitz, 1909. DERMACENTOR RHINOCEROTIS (de Geer, 1778), was used by previous au. thors, including King (1911,1926), probably in error for Dermacentor rhinocerinus Denny, 1843. The species rhinocero. tis described by de Geer is actually an Amblyomma according to Schulze (1932A), whose decision is tentatively accepted here, though specialists are not in complete agreement. HAEMAPHYSALIS sp. nov. Hoogstraal (1954B). Later described as H. bequaerti Hoogstraal, 1956(A). HYALOMMA BRUMPTI Delpy, 1946. Listed by Hoogstraal (1954B) but herein changed to H. impeltatum Schulze and Schlottke, 1930, on the basis of toate resulting from examination of type material. 5 Wh S RHIPICEPHAIUS FAICATUS Neumann, 1908, is a synonym of R. lo Neumann, 1007 (aumpt 1942B,1950A). R. falcatus was listed by King (1926) from the Sudan. Specimens in Sudan Government collections identified as this species by King refer actual. ly to R. longus, R. supertritus, R. simus senegalensis, and R. sanguineus. . falcatus was originally rather sanguineus vaguely access = the complex group in which it falls, and frequently the name has been used indiscriminately. RHIPICEPHAIUS LUNULATUS Neumann, 1907. This name was used (Hoog- straal 19525) on the basis of Santos Dias! (195@,1952C) assertion that this is a valid species, distinct from R. tricuspis Donitz, 1906. Dr. G. Theiler, however, has studied this question so thoroughly (1955 correspondence) and con. firmed her earlier findings (1947) so convincingly that R. lunulatus is herein used in synonymy under R. tricuspis. RHIPICEPHAGUS MACROPIS Schulze, 1936(C) is a synonym of R. san. neus 27 neus Latreille, 1806, according to Zumpt ° e original specimens of R. macropis came from dogs in Port Sudan (Sudan) and Aden (Arabia). RHIPICEPHAWUS PUNCTATISSIMUS Gerst&cker, 1873, is a synonym of are ineus s ineus Latreille, 1806, according to Zum é 7 ctatissimss was listed by King (1908, 1911) from the a ane antos Dias (1952H,1953A,B) considers this to be a subspecies of R. sanguineus , and to be the same as R. sulcatus, but he has not examined type material or reared series. (2?) RHIPICEPHAIUS SHIPLEYI Neumann, 1902. Type locality: "Soudan possibly meaning Anglo-Egyptian Sudan (or "French Soudan", i.e. French Equatorial Africa). Zumpt (1943A,1950A) has synonymized R. shipleyi under R. s. sims Koch, 1844. RHIPICEPHAWS SUICATUS Neumann, 1908, The material on which King's Teport of this species in the Sudan was based has pro- ven upon comparison with Dr. Theiler's reared series of R. sulcatus and large numbers of other specimens to be heavily punctate individuals of R. s. sanguineus. R. sulcatus is known, however, by a few more recent specimens from the Sudan. =) Soe NAMES HEREIN CHANGED, SYNONYMIZED, OR PROPOSED . #AMBLYOMMA WERNERI WERNERI Schulze, 1932(A) is synonymized under A. nuttalli Donitz, 1909. *APONOMMA HALLI Tendeiro, 1950, is synonymized under A. exornatum ~~ (Koch, 1824). BOOPHILUS CONGOIENSIS Minning, 1934, is synonymized under B. annuw latus (Koch, Lo4d4). HAEMAPHYSALIS LEACHII HUMEROSOIDES, a subspecies tentatively sug- gested by Theiler (19Z3B), is an elongate form of H. leachii leachii, so far as now demonstrable. HAEMAPHYSALIS MUHSAMI Santos Dias, 1954(E), is a subspecies of H. eachil. ##"HYALOMMA SCUPENSE™ Schulze, 1919, appears from Soviet works to have distinct biological characteristics and minor morphological characteristics intergrading with H. detritum; it is therefore indicated as a "form" of yet unknown taxonomic status. HYALOMMA IMPELTATUM Schulze and Schlottke, 1930, is considered to have priority over H. brumpti Delpy, 1946. H. erythraeum Tonelli-Rondelli, 1932, is also considered to be a synonym of H. impeltatum. HYALOMMA MARGINATUM Koch, 1844, is said by Pomerantzev (1950) to be a Synonym of H. plumbeum (Panzer, 1795). This matter should be investigated by independent workers. *The fact that these names have been modified or are considered synonyms has been mentioned in the preliminary publication on Sudan ticks (Hoogstraal 1954B), but the reasons for these changes are pre— sented only in the present report. ##Non-Sudanese species. = 30= HYALOMMA LEWISI Schulze, 1936, is a synonym of H. truncatum, not of H. excavatum as stated by Delpy (1949B). **HYALOMMA AEGYPTIUM PUNCTATA Schulze, 1919, is considered as a Synonym of F. aegypti ° **HYALOMMA ALBIPARMATUM Schulze and Schlottke, 1930, is raised from the status of a synonym of H. marginatum (= H. savi i) (Delpy 1949B) and of H. truncatum (Fete Raion full species rank; H. impressum 2g Sp anduSe, 1930, is considered as a synonym of H. iparmatum, not of H. truncatum as stated by Feldman_-Muhs am CISELY. *#*HYALOMMA HUSSAINI Sharif, 1928, is treated as a valid species (not a synonym of H. excavatum - cf. Delpy, 1949B) and most probably the proper type species of the subgenus Hyalommina, the previously designated type species of which (H. rhipi— cephaloides Neumann, 1901) is no more than a misforme iH. excavatun. *#HY ALOMMA TURANICUM Pomerantzev, 1946, is raised to full species rank and considered to be the same as H. rufipes glabrum Delpy, 1949. HYALOMMINA (Subgenus): See H. hussaini above. MARGAROPUS REIDI SP. NOV. is described from males, females, and nymphs. *RHIPICEPHALUS CUSPIDATUS Neumann, 1906, is shown to be actually different from the tick described and illustrated under this name by Zumpt (1950A) in his review of this genus. RHIPICEPHALUS PSEUDOLONGUS Santos Dias, 1953(D) is a synonym of R. Tongus Neumann, 1907. *The fact that these names have been modified or are considered synonyms has been mentioned in the preliminary publication on Sudan ticks (Hoogstraal 1954B), but the reasons for these changes are presented only in the present report. ##*Non-Sudanese species. - 37 - PALPUS na SEGMENT 4... se seoenreeennn a HY POSTOME _---... SEGMENT 3 Son rer CAPITULUN ... SEGMENT 2 .. BASIS CAPITULI-=7~ Secon INT ERNAL COXAL a EXTERNAL SPURS CERVICAL PIT ~~ CERVICAL GROOVE EYE __ PUNCTATIONS ~- LATERAL GROOVE Va GENITAL APRON ...-GENITAL GROOVE ASS externa COXAL SPUR MAL... COXA Fr TROCHANTER | _POSTEROMEDIAN GROOVE aaa _PARAMEDIAN GROOVE ---SPIRACULAR PLATE ANUS -ANAL GROOVE -ACCESSORY SHIELD ants ~~ ADANAL SHIELD LA J ieee SUBANAL SHIELD _. PARMA s 5 “POSTANAL MEDIAN GROOVE _.. FEMUR .. TIBIA ... PRETARSUS =--.=- TARSUS ...... PAD (PULVILLUS) CLAW ~:PGROSE AREA ap AL aes A ‘SCUTUM 1@ Figures 4 and 5, o&, dorsal and ventral views Figures 6 and 7, 9, dorsal and ventral views KEY MORPHOLOGICAL CHARACTERS HYPOTHETICAL co and @ ticks, FAMILY IXQDIDAE (Characters used in keys in this report, except for self-explanatory terms, such as "caudal appendage" of certain Boophilug males. Addi- tional characters used in fuller descriptions of individual species are explained where they first occur). PLATE IV - 38 = KEY TO FAMILIES AND GENERA OF ADULTS* MALES AND FEMAIES A. Dorsum covered by leathery integument lacking scutum. Mouthparts ventral. Eyes absent, or present in lateral folds. Spi- racular plates small, anterior of coxa IV. CSOLE Ticks)... . PS Na Dorsoventral Groove \ Eye - \- Spiracle hb DEN 1 Supracoxal fold Metatarsus Sebi a= > Tarsus — A ¢ Dorsal humps —--§-“* Subapical dorsal — protuberance a) Figure 10, &, genital area Figure 11, 9, greatly engorged, lateral view (sketch) Figure 12, 9, slightly engorged, lateral view KEY MORPHOLOGICAL CHARACTERS, FAMILY ARGASIDAE Ornithodoros savignyi (Egyptian specimens) PLATE VI - 51 - ARGAS INTRODUCTION Six Argas species occur in the Sudan. Four or possibly five other rare kinds have been recorded elsewhere in Africa. All the Sudan species may bite man, and, except possibly for the bat para. sites, all may cause more or less painful sequelae. Insofar as our present knowledge goes, Argas ticks are rarely known to trans— mit human disease. Birds Semen are the chief hosts of most Argas species. The more widely spread species that parasitize domestic fowls are often serious pests and important vectors of avian diseases, The bat infesting species also range rather widely and m&y possibly disseminate pathogenic organisms among their hosts. Larval Argas ticks, in contrast to the two well-known African argasids, Ornithodoros moubata and 0. savignyi, are all active, suck blood, and often may be observed feeding on oe hosts. Argas nymphs and adults, like those of Ornithodoros, are usually found only by careful searching in niches and concealed resting places in the immediate habitat of their favorite host. Ticks of the genus Argas, unless recently engorged, are usual. ly flatter in profile than Ornithodoros, and can in most instances be easily distinguished from them by the presence of some morpholog— ical differentiation of the peripheral integument of the body. Argas ticks are still surprisingly poorly known in most parts of the world and considerably larger collections, more data on their life history, and more exact collecting data are necessary before the biology and systematics of this genus can be finally settled. The other species of Argas known to occur in Africa, some of which undoubtedly will be faa in the Sudan, are the following; A. aequalis (Neumann, 1901) from Tanganyika. The host is unknown = apparently only the original collection is known. Original. ly described in the genus Ornithodoros but subsequently referred to Argas (Neumann 1908B). = eS A. hermanni (Audouin, 1827) from Egypt. Neumann (1896) noted material Trom Ethiopia and Hoogstraal (1952A) from bird nests in Egypt. We are still studying material referable to this species in view of its possibly unsatisfactory taxonomic criteria and species status. A. striatus Bedford (1932A, 1934) from weaver bird nests, Cape Prov- ince, South Africa. This species is said to be closely related to A. aequalis, but both are in need of comparative biological and morphological studies. A. transgariepinus (White, 1846) from South Africa. A. kochi (New mann, an from Basutoland possibly is a synonym. ~“Berlese (1913) reported specimens biting a child and walking on the wall of a bank in Italy. Hoogstraal (1952A) described biology in Egyptian bat caves. Hoogstraal (1954C) noted presence in Spain. A report on a study of the life cycle, biology, and morphology of each stage will be presented in a forthcoming paper. Subgeneric classifications are not included for other groups in this work because their status is still moot. The issue has been forced in the genus Argas by the necessity for deciding whether to refer to some species generically as esis Ogadenus, or Ar Argas. Studies on this subject are presently under way will be eee more fully elsewhere. - 53 - corey, Nisa 7 ; . /Lotera/ a | Margir @ \upfurned Ver ‘et Fee 20 differentiated periphery 13 (striated) subapical dorsal protuberance hood capitulum genital aperture Supracoxal fold spiracle coxal fold anus ventral paired organ Ly ap SNH, g sas, Vs 14 Figures 13 and 14, Q, Hypothetical Argas species KEY MORPHOLOGICAL CHARACTERS, GENUS ARGAS PLATE XII SST = 1. 36 he KEY TO SUDAN SPECIES OF ARGAS MATES AND FEMALES Pair of transverse, slitlike organs situated just posterior of anus. Body outline generally subcircular or wider than long. (Bat PATASI LES Dea cc cis cs ce cicscloeee eeeececiccccee et Ventral paired organs lacking. Body outline oblong, considerably longer than wide. (Not bat PEPASLEOS Jeeisiciccicccicc coc cccwcisslsiels sancssisiel Legs as long as or longer than body. Hood present. Peripheral differentia. tion of integument and lateral suture LACKINg cccccccccccccccccccccccccccccscccccccccccccche BOUETI Figures 23,24,555 54 Legs shorter than bod Yisieisialcicie stcislcicicisisc ove cle cele cievcieis'e slejs\eicicis|e.e Body outline considerably wider than long. Hood present. Peripheral dif. ferentiation of integument and lateral SLUTS PLAC NE elec aieleieisisiee cclsicsiciceicie cee secieecsincic cAe CONRUOUS Figures 25,20,35)50 Body outline generally subcircular or circular, never as wide as A. confusus. Hood absent. Peripheral integument striated and lateral suture present.....ee+eA. VESPERTILIONIS Figures 7A ELD y Picks) Dorsum with large, depressed polygonal areas, Size large (15 m,. to 22 m. long). (Adults parasitic on mammals).......eeeeeeeA. BRUMPTI Figures 19,20,31,52 Dorsum finely wrinkled. Size moderate (10 mm. long, usually less). (Fowl parasites MCSE ETa e's eile 6 d\eio.6.6.ebleleieiseisiecleleinisioce seseeceee see) =D 5. Figures 15 and 16, as persicus Figures 21 and 22, Argas vespertilionis Figures 17 and 18, = as Tef Lexus Figures 23 and 24, Argas boueti Figures 19 and 20, Arga Ss brumpti Figures 25 and 26, Argas DORSAL PERIPHERAL DIFFERENTIATION (odd numbered figures) AND LATERAL Dorsal and ventral periphery of distinct quadrangular "cells". Postpalpal hairs present. (Chicken parasite, common in Susclearay) io ain oyaiels aiataia lola eierole otetele siete eietarerelevelel (ele /eleieitaetarets A. PERSICUS Figures 15,16,27, 20 Dorsal and ventral periphery finely wrinkled. Postpalpal hairs absent. (Pigeon parasite, very rare in Suet) sale eae REFLEXUS Figures 17,13, 29,30 S confusus INTEGUMENT (even numbered figures) OF ARGAS SPECIES. Note especially the presence of a lateral suture in Figures 16,18,20 and 22 and its absence in Figures 24 and 26. = 50 2 Z ~-o- o ao {50} Q, oO » Heed Sh n = E E10 & 46 DUIMICAS Bia rey, Ay i © 3 fob) to (Le a SS dorsal and ventral views dorsal and ventral views Q> > TSLCUS , XUS y reile Ar irae ARGAS (ARGAS) PERSICUS and ARGAS > > Figures 27 and 28 Figures 29 and 30 (ARGAS ) REF LEXUS Egyptian Specimens PLATE IX Ete) ARGAS (ARGAS) PERSICUS (Oken, 1818). (Figures 15,16,27 and 28) THE POULTRY ARGAS OR FOWL TICK*. LN Oo, SC, EQUATORIA PROVINCE RECORDS 1 Kapoeta domestic chicken Jul (SGC) 19 Torit domestic chickens Dec 684 96 Imurok domestic chickens Jan /ifey 9 Juba domestic chickens Dec 1 ag hd age Torit poultry house Dec eee ee Juba poultry house Aug DISTRIBUTION IN THE SUDAN "Throughout the Sudan® (King 1926). In addition to the above. listed Equatoria Province records, two localities have been published in the literature and I have seen specimens from the following places: Upper Nile: Malakal (HH). Bahr El Ghazal: Wau (SVS, HH). Blue Nile: Kosti, Wad el Nail near Singa, Wad Medani (all SG). "Blue Nile Districts” (Archibald 1923). Kassala: Suakin (SCC). Kordofan: Delami (SC). Darfur: Fasher (SVS). Khartoum: Khartoum (SGC, SVS, Gordon College collection, HH). Omiurman (Hj. See also Balfour (1906,1907,1908B,1909,1910,1911A, B,C D,E,G,1912) for spirochete studies on A. persicus from Khartoum, Northern: Dongola (Neumann 1901). Wadi Halfa, Abu Hamed, At- bara, Bd Damer (HH). #In South Africa, called "The Fowl Tampan™ (Theiler 1952A,B). - 59 - DISTRIBUTION A. persicus is now established in most parts of the world between PHT. and 40°S, as a result of transportation of poultry. In Siberia, this tick occurs even farther north than 55°N. (Olenev 1926,1927). In Argentina, 38°S. is its southern limit (Roveda 1940). As an example of the fowl tick's long range spread, it is said to have been introduced into New Zealand from America. Its initial appearance in many parts of the world is believed to have been during early Persian conquests though the species did not necessarily originate in Persia (Robinson and Davidson 1913A). Once introduced, the fowl tick often spreads quickly and widely, as it has done in Argentina where it became a common pest within sixteen years after first reported (Lahille and Joan 1931, Roveda 1940, Lucas 1940). In the United States, after having first been collected in 1872 in southwestern Texas, its dispersion has been"bradual and orderly" (Parman 1926). In other areas it occurs only sporadically. For instance, in Madagascar, A. persicus is said to be restricted to the western coastal lowlands absent from the central uplands (Blick 1935,1948A,C). In Mauritius it is not common or widely spread and seldom appears in large numbers (Moutia and Mamet 1947). The following records are for Africa, Arabia, and outlying islands only. NORTH AFRICA: EGYPT (Savignyi 1826. Audouin 1827, Taschen. berg 187. Neumann 1901,1911. Nuttall et al 1908, Hirst 1914. Mason 1916. Carpano 1929A,B,1935. El Dardiry 1945. Said 1948. Fahmy 1952. Hoogstraal 1952A. Floyd and Hoogstraal 1956. Hurlbut 1956. Taylor, Work, Hurlbut, and Rizk 1956). LIBYA (Zanon 1919. Franchini 1926,1929%. Tonelli-Rondelli 1932A,D. Gaspare 1933. Stella 1938). TUNISIA (Galli-Valerio 1909A,1911B,1914. Comte and Bouquet 1909. Blaizot 1910. Neumann 1911. Langeron 1912, 1921). ALGERIA (Neumann 1901,1911. Brumpt and Foley 1908, Edm. Sergent and Foley 1910,1922,1939. Hindle 1912. Robinson and Davidson 1913A. Donatien 1925. Catanei and Parrot 1926. Foley 1929. Clastrier 1936). MOROCCO (Delano# 1923, Delano& and Lelaurin 1923). WEST AFRICA: NIGERIA (Absence of A. rsicus; Macfie and Johnston 1014. Presence of this tick: Veeco 1943). GOID COAST = 60) = (Stewart 1933,1934). FRENCH WEST AFRICA (Bouvet 1909. Brumpt 1909B. Rousselot 1951,1953B). / LIBERIA: Absence of A. persicus, Bequaert Bn, PORTUGESE GUINEA (Tendeiro 1510 1GCDS Ane 19531954). CENTRAL AFRICA: CAMEROONS (Mohn 1909, Rageau 1953B). BELL GIAN CONGO and RUANDA-URUNDI (Ghesquiere 1919,1921A,B,1922,1928. Schwetz 1927A,B. Bequaert 1930A,B,1931. Gillain 1935. Schoenaers 1951A). No records seen from French Equatorial Africa. EAST AFRICA: SUDAN (Neumann 1901. As A. miniatus: Balfour 1906. Balfour 1906,1907,1908B,1909,1910,1911A ,B,c ,D,5,G,1912. Nuttall et al 1908, King 1908,1911,1921,1926. Archibald 1923. Tonelli-Rondelli 1930A. de Beaurepaire Aragao 1936. Kirk 1938B. Hoogstraal 1954B). ETHIOPIA (Neumann 1911). ERITREA (Franchini 192%. Niro 1935. Stella 1938A,1939A,1940. Ferro-Iuzzi 1948). BRITISH SOMALI. LAND (Drake-Brockman 1913B,1920. Stella 1940). ITALIAN SOMALILAND (Brumpt 1909A. Paoli 1916. Franchini 1925,192%. Niro 1935. Stella 1938A,1939A,1940). KENYA (Anderson 1942A,B. Lewis 1931C,1939A. Piercy 1948. Wiley 1953). UGANDA (Mettam 1932 stated that A. persicus was then not yet reported, but Wilson 195@ lists it as present). J TANGA. NYIKA: ?No records. / SOUTHERN AFRICA: ANGOLA (Howard 1908. Absence in San Salvador: Gamble 1914. Sousa Dias 1950. Bacelar 1950. Santos Dias 195M). MOZAMBIQUE (Howard 1908,190%,1910. Theiler 1943B. Santos Dias 1953B,1954H). "RHODESIA" (Robinson and Davidson 1913A). NORTHERN RHODESIA: ?No records. 7 SOUTHERN RHODESIA (Little 1919,1920. Jack 1921, 1928,1937,1938,1942. Cooper 1944). NYASALAND (De Meza 1918A. Wilson 1950B). SOUTHWEST AFRICA (Tromsdorff 1914. Sigwart 1915. Warburton 1921. Mitscherlich 1941. Schulze 1941). BECHUANALAND (As “tam. pans: ™J.G.™ 1943). UNION OF SOUTH AFRICA (Lounsbury 1895,1899B, C ,1900A ,B,C ,1903B,1904D. Dédnitz 1907,1910B. Howard 1908 ,190% , 1910. Nuttall et al 1908. Bourlay 1909. Jowett 1910. Neumann Sole 1911. Robinson and Davidson 1913A. Bedford 1920,1926,1927,1932B, 1934. Du Plessis 1932, Robinson and Coles 1932. Bedford and Coles 1933. Bedford and Graf 1934,1939. Monnig and Coles 1934, 1936,1939,1940. Coles 1933,1945. Cooley 1934. des Ligneris 1939. Mitscherlich 1941. R. du Toit 1942B,C,1947A,B. Gericke and Coles 1950. Annecke and Quinn 1952). OUTLYING ISLANDS: MAURITIUS (As A. mauritianus: Guerin. Meneville 1020-165. Neumann 1911. De Charmoy 1914,1915,1925. Moutia and Mamet 1947). MADAGASCAR (Recorded by Blick 1935,1948A, C,1949. Millot 1948 states that A. persicus does not occur on Madagascar but Blick seems to ee evidence tnat it does. Hoogstraal 1953E). REUNION (Gillard 1947,1949). SEYCHELLES (Millot 1948). / ZANZIBAR: ?No records. 7 ARABIA: YEMEN (Hoogstraal, ms.). HOSTS: A. persicus in all stages is chiefly a parasite of chickens. Ducks, geese, turkeys, and infrequently pigeons, are attacked. This parasite often becomes so numerous in fowl houses that the birds die from exsanguination. Canaries are sometimes attacked, and in South Africa young ostrichs have been killed from the vol. ume of blood lost to these ticks. Wild birds may be infested if they construct large, numerous, or fairly permanent nests in the vicinity of human activities. The question of infestation of other wild birds and of mammals is a most uncertain one. Although the fowl argas does parasitize man on occasion, the frequency and fierceness of these attacks have been fancifully exaggerated and enhanced to the point that it might even seem advisable to exterminate Africa's chickens rather than subject mankind on this continent to the scourge of his fowls* argasid parasite. Wild bird hosts Rookeries of the buff—backed heron, Bubulcus ibis ibis, in parks in and near Cairo (Hoogstraal 1952A) and heron rookeries in South Africa are heavily infested (Theiler, correspondence). In the Nile Barrage Park near Cairo, literally tens of thousands of 4 (ee fowl ticks in all stages can be found in crevices and under bark of any large fig tree in which herons roost and nest. In Pakistan, Abdussalam and Sarwar (1953) found frequent para. Sitism of vultures and common herons in sixteen kinds of trees in which these birds nest. Other birds and palm squirrels also perch in. these trees, but only a young kite was found infested. (Whether smaller birds and squirrels were examined for ticks is not clear from the report). On trees with relatively smooth bark and few cracks, ticks extended down the trunk almost to the ground, but on those with cracked bark they concentrated chiefly in the upper branches near the perches of their hosts. (In the Cairo area, rough.barked trees harbor tremendous tick populations from near the roots to the crown). The incidence of ticks in trees harboring vultures and herons was mich higher than it was in nearby chicken houses. Specimens have occasionally been reported from isolated nests of wild birds and on ground birds such as quail. Howard (1908) recorded the secretary bird and Bedford (1934) the guinea fowl as wild hosts, Theiler (unpublished) informs me of the following South African records: wattled crane, hadada ibis, and pelican. King (1926) reported the guinea fowl, buff—backed heron, and crow as wild hosts of the larval stage in the Sudan. Specimens from guinea fowl at Khartoum (SCC) probably came from zoological garden hosts. Identification of larvae from wild birds that construct iso tated nests and that do not live close to human habitations should be regarded with suspicion if these larvae have not been identified by a contemporary expert in argasids. Larvae of related species closely resemble those of A. persicus. Wild bird parasites are so poorly known that the presence of argasid larvae on them should be a hint to consider rare or poorly known tick species before con. cluding definitely that those found are A. persicus. The mouthparts of larvae pulled from birds are Sasills, See oe extreme cau. tion is exercised and the body characters are frequently obscured by engorgement so that it is difficult to identify the material. Wild Mammal Hosts Apparently the only authentic report for the fowl tick from a wild mammal is a note of three adults from a Texas jack rabbit shot in 1906 (Hooker, Bishopp, and Wood 1912). =MOo c= Domestic Mammal Hosts Howell, Stiles, and Moe (1943) believe that A. persicus may feed on cattle more commonly than is generally suspecte ib reasons for this assumption are not presented. This tick has been vaguely reported from Persian animals (Aluimov 1935) and, on the basis of a museum specimen label, from cattle in the Congo (Schwetz 1927B). Hoffman (1930), apparently from personal information, sta. ted that in Mexico A. persicus may bite animals and man in the ab. sence of fowls. In the United Provinces of India, Sen (1938) listed this species "off dog”. Various workers have reported that they were unable to induce the fowl argas to feed on laboratory or domestic animals, or, if some blood was taken, the meal was only a partial one. Human Hosts Authentic records of A. persicus attacking man in almost all instances stress the infrequency of such experiences. Reports in certain textbooks of medical entomology that the fowl tick is an important pest of man or even "a veritable Scourge eeceeees in the Sudan and South Africa (1) are without the slightest foundation (see below). In the Sudan, King (1926) reported, A. persicus rarely bites man, Several nymphs and adults in Sudan Government collections are labelled "from Yemenese man, Suakin, 7-3-09, 0. Atkey". The inference is that the specimens were taken on the person. The numerous Kosti specimens already mentioned in Sudan records arouse suspicion that this species have been a pest in houses there at one time. My own inquiries in many parts of the Sudan and from reading a considerable number of travel, medical, and natural history reports of the Sudan have failed to reveal any indication that A. persicus is known as a human pest anywhere in the Sudan. In South Africa, Bedford (1934) wrote, A. persicus seldom at— tacks man. Lounsbury (19000 ,1903B) recorded a severe bite on a person in Graaff-Reinert, and stated that he had heard of two other persons who were bitten, but, especially in the former paper, he minimized the importance of A. persicus as a pest of people, as did Behr (1899) for California. Howard (190%), however, heard of a South African cart that had been stored in an old infested chicken house; "no one was able to ride in it afterwards", In Southern Rhodesia, A. persicus is pre-eminently a fowl parasite (Jack 1921). = 64. Drake-Brockman (1913B,1920) stated that this tick is found in or near huts in British Somaliland but that it does not bite man there. From the United Provinces of India, Sen (1938) noted A. rsicus "on bed (presumably can infest man)", "This species was reported from Quetta (India) where it was stated to infest houses and to bite human beings” (Warburton 1907). As stated above, Hoffman (1930), remarked that in Mexico A. persicus may bite man and animals in the absence of fowls, but details were not provided, In Korea, Kobayashi (1925) "examined certain specimens of Argas persicus ....... said to have stung men!*, Old Iranian (Persian) reports that A. persicus is such a pest of human beings that whole villages Heverhen to te ee, so wide. ly quoted from Nuttall et al (1908) who reviewed the earlier liter. ature, hardly bear contemporary repetition. The evidence in all cases is circumstantial and based on hearsay. That these fables should have gained the stature of serious fact in mst books of medical entomology is a reflection on methods of textbook fact. finding. One writer has even gone so far as to throw in for good measure a large part of the African continent as a scourged area. Since 1890 there has been hardly a single published eyewitness or corroborated report of Areas persicus biting human beings that has not referred merely to Taclated Tisteases, Though some bites have been described as painful, only one or two have been shown to cause other sequelae. Twentieth century Iran has not provided evidence to corrobo— rate the early apparent misrepresentations concerning the fowl tick. Carre (1909), in reporting on the frequency of larval at— tacks on chickens in Teheran did not mention that man is attacked. Harold (1922) expressed the belief that Ornithodoros lahorensis Neumann is actually responsible for pai ites attributed to A, rsicus in Iran / cf. also Harold (192097. Dr. Baltazard, Direc. = of the Pasteur Institute in Teheran, an outstanding student of argasid ticks and of their disease relations, informs me that he knows of no troubles from A, rsicus in Iran so far as human beings aie ee Delpy T1975) observed large numbers ee persicus O. lahorensis in and around peasant houses and stables near Persepolis In fran but he did not mention bites of either spe. cies. Delpy and Kaweh (1937), however, record an actual observa. tion in Iran of a laboratory person who, when washing his hands, noted a large nymphal fowl tick biting him. The bite was painless but the victim succumbed to a bout of anthrax, demonstrated to have been transmitted by the attack, = (ei In Palestine, Nicholson (1919) and Dunlop (1920) attributed human relapsing fever to bites of A. persicus. Their reports. were based entirely on circumstantial evi ence /cf. also Balfour (1920A,B), Woodcock (1920), MacKenzie (1920), etc.7. Experimental evidence negates this probability. Members of my staff and I on several occasions have questioned people who spend much time in heavily infested parks and houses in and near Cairo without finding anyone who acknowledged being bitten. Apparently reliable accounts of A. persicus infesting human huts in which chickens are also kept, an not infrequently biting persons, are those of Sergent and Foley (1910,1922,1939) from AL geria. Natives there refer to fleas and to the fowl argasid by the same name, Although the ticks are frequently associated with cases of human relapsing fever, they were proven by these observers to have a negative role in the transmission of spirochetes causing the disease. There are a few scattered, apparently authentic reports of A. ersicus biting man outside of Africa. One such, a vivid descrip. tion enhanced by illustrations of the tick and of dark weals where the human victim was bitten, has been reported from Romania by Ciurea and Stephanescou (1929). The attacks occurred inexplicably in the upper stories of a new concrete apartment house and no chickens or pigeons were known to have been associated with the buildings. With regard to the lively account of attacks by ™A. rsicus™ on indigent persons in Chile (Porter 1928), see A. reflexus, p. "ils Reptile and Amphibian Hosts Although A. persicus always shows a predilection for avian blood, it will feed on toads if the skin of these animals is warmed, according to Galli-Valerio (1911B). The blood is probably toxic for the ticks die afterwards. The record of A. persicus from a tortoise in Iran (Michael 1899) is most probably based on misidentification or incorrect or incomplete specimen labelling. = 66— Infestation of Human Habitations (Africa) African records of A. persicus in huts of indigenous people (inferred presence of chickens in some huts) are: Annecke and Quinn (1952) for South Africa, Drake-Brockman (1913) for Somali_ land, Sergent and Foley (1919,1922,1939) for Algeria, and Sudan records above. Lounsbury (1903B) stated that the fowl tick seldom occurs in South African houses unless chickens are kept close by. BIOLOGY Life Cycle Among the many references to some phase or other in the life cycle of A. persicus, some of the mre important are; Lounsbury (1903B) for Satan T Sine Nuttall et al (1908) for laboratory observations, Olenev (19284) for the Saratov area of Russia, Roveda (1940,1950) for Argentina, Bodenheimer (1934) on temperature and humidity tolerance, and Zuelzer (1920A,B,1921) on feeding, excre- tion, and life cycle. Hooker, Bishopp, and Wood (1912) contributed a detailed study of the life cycle in southern United States and reviewed earlier literature. These authors also provided data on growth and size of the fowl tick. The life cycle in general under favorable conditions requires about four months. Larvae attach usually to the base of the host's wing. They feed there for five to ten days before dropping from the host and seeking shelter. Nymphs and adults become satiated in from five minutes to two hours and then seek a sheltered place in the build ing, yard, or tree in which they secrete themselves. Feeding is usually done at night, sometimes in subdued light, seldom if ever in strong light. Coxal fluid is emitted within a few minutes after engorgement is complete and often while the tick is stationary or moving about the host, but only infrequently while the mouthparts are still in. serted in the host's skin. Ove Digestion is extremely slow and fowl blood may be identified by the precipitin test for at least 23 months after ingestion (Gozony, Hindle, and Ross 1914). The following notes are chiefly from Hooker, Bishépp, and Wood (1912). Many more details may be found in their report. Usually females oviposit after each meal, which may number up to six or seven in a lifetime. Under exceptional conditions, a female may require two blood meals before laying eggs. The greatest number of eggs deposited after the first few blood meals increases progressively from 195 to 646, but decreases after sub. sequent feedings to as few as 47 eggs following the seventh or last feeding. The average number of eggs laid after each engorge— ment was: first, 131; second, 159; third, 133; fourth, 110; fifth, 97; sixth, 95; seventh, 47. Eggs are laid in the adult tick's retreat. Oviposition generally commences four to ten days after feeding, in summer sometimes as early as the third day. In winter or in the absence of males, egg laying may be delayed for weeks or months. Oviposition of moderately large batches continues over a six to ten day period but only three days are required for depositing a small number of eggs. In nature it appears that the fowl argas seldom engorges and oviposits more than five times, unless females com. mence feeding early in the spring. Incubation of eggs extends over an eight to eleven day period in warm summer weather, but in cooler climates or seasons this pe— riod is extended to three weeks or even longer. As stated above, larvae generally feed for from five to ten days, but they may complete engorgement in three or four days, and Rohr (1909) recorded two days. There is some indication that quiet, setting hens allow the greatest number of larvae to thrive, and that different breeds of hosts exert no influence on larval devel- opment. In NAMRU_3 laboratories, Dr. Herbert S. Hurlbut (unpub. lished) is finding that only a moderate number of larvae kills chickens used in his experiments, apparently not doing so by trans. mission of pathogenic organisms or by exsanguination. Nymphs and adults resulting from these larvae have no observable deleterious effect on their hosts. Reasons for this exceptional larval toxic ity have not yet been ascertained. =OOr= Larvae survive unfed for over five months in cool weather, but in Texas during midsummer they succumb in about two months. At 30°C. and 70 R.H., unfed Egyptian larvae survive for up to thirty days (H. S. Hurlbut, personal communication). Larvae molt to nymphs in warm summer weather about four days following completion of feeding. Nymphs usually feed twice, in a matter of half an hour (some. times two hours) and mlt a week or two (sometimes longer) after- wards. Some nymphs undergo a third molt before reaching adulthood; this phenomenon cannot be correlated with sex, food supply, or climatic conditions. Unfed second instar nymphs survive up to a year but first instar nymphs are known to live for only up to nine months. Female feeding has been discussed above. Copulation is simi. lar to that described for 0. moubata (page 134). Adults may live as long as three years without food (Laboul- bene 1881) but this appears to be exceptional. Unfed adults gen. erally succumb more rapidly than engorged adults, which normally appear to live from five to thirteen months, but which may on oc~ casion survive longer. Besides being a particularly intriguing study for some workers, the ability of the fowl tick to withstand starvation for long pe— riods no doubt accounts in part for its wide distribution and large numbers. Observations made by Newman (1924) on longevity without food were summarized as follows: Test 1: An isolated female lived two years and three months, (2) it produced fertile eggs four months after isolation, and (3) larvae lived for three months. Test 2: (1) Males died four months after isolation, (2) first female died after two years and four months, (3) two females lived three years, (4) three females lived four years, and (5) the maximum time a fe male lived was four years and five months. Removal of fowls from a house or yard is in itself of little use in ridding the premises of ticks. Larval survival without food for 228 days at 22°C. to 26%. and 906 to 100% relative humidity was reported by Roveda (1940). At temperatures of 37°C. to 38°C. and at relative humidities of 85% to 100% larval survival was reduced to an average of 50 days. ES) = Ecology All stages congregate on walls, in crevices, or between boards of poultry houses. Around Cairo we find literally tens of thousands under loose bark, in crevices, and on the trunks of trees in city- park heron rookeries. Trees in which chickens roost are frequently reported as hiding places for A. persicus. Other habitats have been discussed under HOSTS above. The ability of eggs, larvae, nymphs and adults to withstand a wide range of temperature and humidity conditions has been studied by Bodenheimer (1934). Fifty-nine observations of nymphs and adults in temperature gradients ranging from 29°C. to 47.7. failed to ex hibit a significant response to changing temperature stimuli. While the vital optimum of the egg stage is 20°. and 80% relative humid ity, the tolerance to fluctuating climatic factors is remarkably great. Even at 20 relative humidity, mortality is only slightly greater than at optimum conditions of environmental moisture. The thermal constant for the egg stage is 316 day-degrees. At temper- atures of from 33°C. to 18°C., eggs hatch in from 10.5 days to 33.3 days (from highest to lowest temperature). Temperatures of 15°C. and below inhibit egg hatching. At high temperature (33°C.), a relative humidity of at least 60% is necessary for hatching. At moderate temperature (18°C. to 27°C.), there is little difference in numbers of larvae hatching from eggs maintained at various per. centages of relative humidity ranging from twenty to a hundred. The ability of A. persicus to withstand desiccation and high temperatures has been studi by Lees (1947) in his excellent re- search on transpiration and epicuticle structure in ticks. In Argentina, the optimum temperature for egg hatching is said to be between 22°C. and 38°C. with relative humidity from 9% to 1006. Mortality increased from 4.85% under the above conditions a ki at 37°C. and 8% to 95% relative humidity. / Roveda 1940 It appears, from these observations as well as from the com paratively great adaptability of this species as demonstrated by its wide geographical range, that Argas persicus is less restricted by higher humidity factors than are many other argasids. - 70. In cold climates such as Saratov, Russia, development occurs only at temperatures over 20°C. Exposure to high humidity (pre sumably at cold temperatures) kills the ticks (Olenev 1926,1928A). The presence or absence of A. persicus in coastal areas fre- quently is referred to in literature. Lounsbury (1903B) stated that A. persicus is everywhere common in South Africa, including coastal towns and areas. Howard (1908) reported that in South Africa this tick is common except near the coast and that the same distributional pattern had been reported from Australia. Records from a number of other localities indicate that the fowl tick does indeed inhabit coastal areas. For instance, Theodor (1932) reports this species especially common along the Mediterranean coast and in Jordan Sea areas of Palestine. In Egypt, we find it commonly in coastal villages and cities. We have also found cast nymphal skins at Djibouti, the seaport of French Somaliland. It occurs at Port Sudan on the Red Sea coast of Sudan (Sudan Gov- ernment Collection record) and at Hodeida on the opposite coast in Yemen (Hoogstraal, ms.). In Reunion, Gillard (1949) reported the fowl tick particularly common on the coast, and in Madagascar it occurs chiefly in coastal areas (Buck 1935,1948A,C). Concerning altitudinal range, A. persicus is frequently re- ported as common in lowlands and rare or absent in highlands. In mountains of the Sinai Peninsula of Egypt we find numerous speci. mens at elevations up to 6000 feet. Lewis (1939A) stated that in Kenya A. persicus is present only in European areas. This is certainly not true for the Sudan, where chicken flocks of many remote, indigenous tribes have been known to be infested for half a century. More recently, Wiley (1953) indi- cated that the fowl tick is increasing its range in Kenya. An apparent negative geotropism displayed by unfed larvae reared from adults collected from trees serving as a heron rookery has been observed by Dr. H. S. Hurlbut at NAMRU_3, Cairo. At the same time, larvae from adults collected from chicken houses ap- peared to show a positive geotropism. The F, larvae of adults from heron rookeries were inclined to prefer herons rather than chickens as hosts and the reverse appeared true for larvae from adults from chicken houses. Informal as these observations are, they suggest an interesting research problem. - 71- DISEASE RELATIONS MAN: Reported sequelae of the fowl argas' bite range from itching to death. Actually, there are no trustworthy accounts of severe illness resulting from a fowl argas bite. Anthrax (Bacillus anthracis), however, has been transmitted to man by the bite of this tick 1n the laboratory on one known occasion. Speci. mens have been experimentally infected with plague (Pasteurella stis) and with yellow fever virus. A. persicus has been reported in textbooks and discussion papers to transmit human relapsing fever (Borrelia spp.) but there does not appear to be a shred of conclusive evidence to support this claim. Experimental studies to date negate this possibility. The little work done on A. per sicus in relation to typhus has gone only far enough to eretee the etiologic agent (Rickettsia prowazekii) survives in the tick for ten days. The fowl tick 1s susceptible to parenteral infec— tion with West Nile virus but does not transmit the virus. FOWLS: The fowl argas is frequently so numerous that birds are killed by exsanguination. Spirochetosis / Borrelia anserina (= B. gallinarum) 7 of chickens, ducks, geese, turkeys and canaries is a serious disease transmitted by A. persicus nearly everywhere that it is found, but not everywhere. Fowl piroplasmosis (Ae ianella pullorum) is also transmitted by the fowl argas, which also has been suggested to be a vector of Grahamella gallinarum. A condition called fowl paralysis by some students and tick paralysis by others, due possibly to a toxin from the tick, sometimes occurs after bites. Chicken cholera or fowl plague (Pasteurella avicida). may cause the death of birds that eat infected ticks. Virus in duced fowl tumors are not transmitted by bites of the fowl argas. See next paragraph. WIID BIRDS: Populations of this tick from Egyptian rookeries of the buff—backed heron or cattle egret have been found infected with Salmonella typhimurium, but others from chicken yards were negative for Salmonella spp. CATTIE: Successful experimental transmission of anaplasmosis (Anaplasma marginale ) by A. persicus has been reported. = 12 = MISCELLANEOUS: It has been claimed that West African speci. mens have been found infected with Q fever (Coxiella burnetii). In Egypt, the fowl argas is infected with one or more viruses distinct from West Nile but otherwise unidentified. REMARKS An excellent and detailed study of the internal and external morphology of A. persicus has been presented by Robinson and David. son (1913A,B,19L)- and by Patton and Cragg (1913). An earlier work is that of Heller (1858). Rohr (1909) reported on life cycle and biological studies in Brazil, and included a few photomicro— graphs of internal organs. Regeneration of broken appendages has been reported by Hindle and Cunliffe (1914) and by Nuttall (1920p). Sensory perceptions have been studied by Hindle and Merriman (1912). The coxal cymatium of A. persicus has been discussed by Schulze (19364), who also (1941) described and illustrated the hallerts organ. Micks (1951) gives an account of a convenient rearing technique and life cycle observations and Sapre (1943) described his method for laboratory rearing. Immmity of chickens to bites of the fowl argas has been studied by Trager (1940). Anticoagulin in the salivary glands and gut has been reported by Nuttall and Strickland (1909) and by Cornwall and Patton (1914). The salivary glands have been described and illustrated by Heller (1858) and Elmassian (1910). Larvae of clothes moths, Tineola biselliella, have been ob. served attacking living larvae of E. persicus in laboratory colo- nies (Volimer 1931). According to Zuelzer (1921), A. persicus and A. reflexus mate and produce fertile offspring. We have been unable to duplicate these results in our Cairo laboratories. Observations on the bacteriostatic factors in blood—engorged ticks, including A. persicus (Anigstein, Whitney, and Micks 195QA, B), prompted further studies showing that bacterial growth inhi. bition in vitro is comparable with the phenomenon induced by anti- biotics (Whitney, Anigstein, and Micks 1950) and that a blood hydrolysate called sanguinin is responsible (Micks, Whitney, and “or he 1951). This subject is reviewed under 0. moubata (page 178 = 73)= Symbiots of A. persicus have been described in some detail by Cowdry (1925C ,1920A,1 and by Jaschke (1933). The subgenus Argas is tentatively defined as follows: "Parasites chiefly of fowls. Morphological characters entirely of genus Argas. Sutural line (i.e. lateral groove) encircling body. With a flattened body flange morphologically differentiated dorsal ly and ventrally by a row of quadrate cells or by fine striations or wrinkles; body shape elongate. Integument finely wrinkled; discs conspicuous, radially distributed; lacking ventral “paired organ™, Hood lacking; mouthparts posterior of anterior body margin by a distance about equalling their own length. Legs moderate; tarsal humps lacking", A. reflexus was designated as the type species of the genus Argas by Latreille (1802) and is so considered by Cooley and Kohls TSZZ) and by Pospelova-Shtrom (1946) / Nuttall et al (1908) pre. ferred to use A. persicus 7. A. reflexus would, therefore, also be the type species of the subgenus Argas. The size of each stage and of each sex, engorged and unengorged, has been reported by Hooker, Bishopp, and Wood (1912) and by Campana. Rouget (1954). IDENTIFICATION A. persicus is easily recognized by characters listed above for the subgenus Argas, with the restriction that its dorsal and ventral periphery 1S marked by a row of quadrate "cells™ (fine striations in A. reflexus). The male is seldom over 5.0 mm, long and has a semicircular genital aperture. The female measures from 4.0 m, to 11.0 m, long and has a narrow, transverse genital aperture. Nymphs are similar to adults except that they lack a genital aperture although advanced instars may have a shallow depression in its place. Larvae are nicely illustrated in various editions of Brumpt's Precis. - 74 - ARGAS (ARGAS) REFLEXUS (Fabricius, 1794). (Figures 17,18,29 and 30) THE PIGEON ARGAS ity Nee ot, C! EQUATORIA PROVINCE RECORDS 7b) wel aaa el Juba domestic pigeon cote Nov al! Juba domestic pigeon cote Jan These specimens were collected in 1949 and 1950 but subsequent— ly we have been unable to find the pigeon argas in Juba or elsewhere in the Sudan. These few may have been remnants of stragglers or of a small number of introduced individuals. If it is a normal inhahb itant of the Sudan, the pigeon argas is sporadic and rare here. No other specimens are known from the Ethiopian Faunal Region, except those reported by Rousselot (1951,1953B) from French West Africa and one, possibly this species, from Kenya (Heisch 1954B). DISTRIBUTION Argas reflexus appears to be a Near or Middle Eastern tick that has spread northward through Europe and Southwestern Russia and eastward to India and elsewhere in Asia (the status of related species or subspecies in Asia requires further study). It may have been accidentally introduced into a few localities in the Ethiopian Faunal Region north of the Equator and to parts of the Americas. If so, transportation of infested domestic pigeons undoubtedly has been responsible for this range. NORTH AFRICA: EGYPT (El Dardiry 1935. Hoogstraal 1952A. Taylor, Work, Hurlbut, and Rizk 1956). ALGERIA (Nuttall et al 1908. Neumann 1911. Presence not subsequently verified). ~_ £ Unknown in Tunisia (Colas-Belcour 1929B).7 WEST AFRICA: FRENCH WEST AFRICA (Rousselot 1951,1953B from Bamako, Soudan). = TE EAST AFRICA: SUDAN (Hoogstraal 1952A,1954B). [ ?KENYA: Heischts (1954B) specimen may represent a closely related species. 7 NEAR EAST; A. reflexus has been reported from Palestine (Theodor 1932), Turkey (Vogel 1927, Kurtpinar 1954) and Iran (Delpy 19478). Its occurrence in intervening areas is to be expected. FAR EAST: According to Sharif (1938), this tick, as variety indicus Warburton, is an important pigeon parasite all over India. EUROPE: In Europe, A. reflexus is generally distributed and extends at least as far north as Denmark (Christiansen 1937) 6° ot occurs also in the British Isles. The Russian range of this tick is said to be confined to the Caucasus, Crimea, and areas bordering southern Europe (Pavlovsky 1948, but Olenev (1929A,B,C ,1931A,8,C) also includes Middle Asia and western Siberia. Oswald (1939) did not find the pigeon tick in Yugoslavia. [ AMERICAS: Cooley and Kohls (1944) list western United States and Columbia as collecting localities for ticks that they call A. reflexus but that show morphological differences of yet unknown SSS * : . lmportance as species indicators. 7 HOSTS Domestic pigeons are the chief host of A. reflexus and are mentioned by all authors. Man is frequently attacked, especially in the vicinity of long unoccupied pigeon cotes. Chickens, horses, and (in America, see above) wild birds such as the condor, swallow, and screech owl (Cooley and Kohls 1944) have been listed as hosts. In the laboratory, any usually available mammal may serve as host. The literature contains numerous reports of A. reflexus biting man and the painful sequelae of these attacks. Although the pigeon argas is mostly strictly associated with pigeons, the exigencies of its domestic existence drive it to attack persons, possibly more frequently than does A. persicus. UNS Early literature concerning the pigeon argas as a parasite of man has been reviewed by Nuttall et al (1908). More recently, Kemper (1934) attributed four cases in Germany to the effects of warm weather. Kemper and Reichmuth (1941) reviewed the literature and reported over twenty attacks in Germany. They believe it possible that this tick might not be able to complete its life cycle on hu. man blood. It is now evident that Porter's (1928) spirited account of "A. rsicus" in Celama, Antofagasta Province, Chile, mist be referred to A. reflexus (or to the American variant; see DISTRIBUTION above), as indicated in the following paragraph. Bites of these ticks were sufficiently numerous and painful for attention to be devoted to the matter in the daily press of the region. Specimens furnished parasitologists as the cause of this "grave molestation” were iden tified as A. persicus. Concerning Porter's report, Kohls (correspond. ence) has provided the following note for inclusion here. "Harly in 1950, I received from Dr. Amador Neghme R., Chief Department of Parasitology of the Public Health Service, Chile, four adults and a nymph (said to be) Argas rsicus, collected in the Province of Antofagasta at ths town of Calama. This seems to be the only Chilean place where this tick occurs, and is found in human houses and in dovecotes. In reply to his letter, I said, "Study of this material indicates that the ticks are not Argas persicus but Argas reflexus, ..e.. The only South American specimens of ae that 1. have seen previously were collected in chicken coops at Bogota, Colombia..... The Calama speci_ mens appear to agree in all particulars with reflexus of the Old World and from Bogota except for the presence of a few quadrangular plates interspersed with the striae on the flattened margins. This difference could well be due to variation and for this reason I would like to have more specimens from Calama for study™. In response to this I received eleven adults from Calama and twenty adults and fifteen nymphs from Chuquicamata, a town about twenty miles away. The source was not mentioned in either case, but all the specimens proved to be the same as those sent previously. In brief, these specimens from Chile that I have seen ere not Argas persicus but are probably local variants of A. reflexus”. The subject of A. reflexus as a human parasite will be treated more fully in a subsequent volume of this work. = = BIOLOGY The pigeon tick may remain unfed in or near pigeon houses for many months, or even for several years (Nuttall et al 1908, Mayer and Madel 1950). Feeding is much like that of A. persicus, which attacks poultry, and is accomplished at night. “Domestic chickens are apparently considerably less liable to attack by A. reflexus than are pigeons. Hiding places of these ticks are easily found in the cracks and crevices of pigeon cotes. The life cycle ap pears to be much like that of A. persicus. Restrictive and opti. mum biological and climatic factors have not yet been reported in literature. Females feed prior to oviposition, but according to Schulze (1943B), males require only a single blood meal annually. During the larval stage there is no urinary or fecal excretion (Enigk and Grittner 1952). Nymphs and adults immediately after feeding discharge a mixture of urine and feces, followed by further excretion the following day. The simultaneous deposition of urine and feces causes the rapid formation of a "guanocrystal™ in the viscous mass, thus frequently leaving a white center of urine sur- rounded by a dark fecal ring on the surfaces on which the substance has been deposited. (Note: Compare this type of excretion with that of Ornithodoros moubata)., Adults deposit only urine for some two weeks after feeding, then at long intervals a mixture of feces and urine. Four weeks after feeding, females begin oviposition, during which time no excretion is seen. Coxal fluid is seldom voided during feeding, but usually begins only following complete engorgement (Zuelzer 1920B, and our own observations). DISEASE RELATIONS MAN: Human beings who venture near occupied or long abandoned pigeon houses are readily attacked, and the ticks may invade nearby human habitations after pigeons have left their usual resting places. Pain or irritation may be felt for years after the pigeon argas has bitten. This species is incapable of transmitting spirochetes of African tick-borne relapsing fever (Borrelia duttonii). PIGEONS: Squabs are especially susceptible to bites of this tick and adults too may suffer to the point of death by exsanguina. tion when their houses are heavily infested. The pigeon argas is of negligible importance in the transmission of Salmonella bacteria among pigeons but does transmit fowl spirochetosis, B. anserina (= B. gallinarum). It is said to be probably capable of transmitting fowl piroplasmosis (Aegyptianella pullorum). Eis = MISCELLANEOUS: In Egypt, the pigeon argas is infected with one or more viruses distinct from West Nile but otherwise uniden. tified. REMARKS Egypt is the only territory on continental Africa where the pigeon tick is known to be of some economic importance. The re— view of this species for the present work has not been as inten. sive as for most other -species. The temperature preferences of unfed and engorged pigeon argas have been described by Herter (1942). The anatomy has been described by Pagenstecher (1862). According to Zuelzer (1921), A. persicus and A. reflexus mate and produce fertile offspring. We have unsuccessfully attempted to duplicate this phenomenon in our Cairo laboratories. With reference to remarks on coxal fluid by Remy (1921,1922B), see O. moubata section, page 173. See also Lavoipierre and Riek (Ss) a am Senevet (1920A) discussed the relationship of the size of the pads of the first pair of legs in larvae in relation to overall body size. Schulze (1943B) figured the midgut, as A. columbarun, to il lustrate his observation that in the argasids, and particularly in this species and in bat-parasitizing species, there is little basal branching of the diverticula but considerable distal branching. Schulze (1941) also noted and illustrated the haller's organ of each stage of the pigeon argas. K. W. Neumann (1942), a student of Schulze, discussed the morphology and function of the dorsal plate of the larva, also under the name A. columbarun. There is some chance that the name A. reflexus refers to a European parasite of wild birds and the correct name of the pigeon parasite should be A. columbae (Hermann, 1804) (cf. DuBuysson 1924). - 19 - Schulze (1943B), referring to this species as Argas columbarum Shaw, 1793, a name usually considered as a nomen nudum, cited as his authority an apparently unpublished thesis on biology of the pigeon argas by one K. H. Muller (1939, Berlin), whose report I have not seen. Note, under remarks for A. persicus, the characters of the sub. genus Argas and that A. reflexus is considered to be the type spe~ cies of the genus and of the subgenus (page 74 ). A tick of questionable systematic status, A. hermanni (Audouin, 1827) is closely related to or identical with A. reflexus. When commencing the study of this group (Hoogstraal 1952A) it appeared that the two were valid species but subsequent investigation has left me with some doubts. Further observations are at present under way. IDENTIF ICAT ION Remarks under identification of A. persicus apply. Note that the dorsal and ventral body periphery of A. retlexus is composed of irregular striations. OO = POP MEME A OT Figures 31 and 32, 9, dorsal and ventral views ARGAS (OGADENUS) BRUMPTI Sudan specimen PLATE X oles ARGAS (OGADENUS) BRUMPTI Neumann, 1907(B). (Figures 19,20,31 and 32) BRUMPT'S ARGAS LONE On, EQUATORIA PROVINCE RECORDS 1) Imurok Heterohyrax brucei hoogstraali Feb al Imirok Heterohyrax brucei ogstra Feb Cin aon il! Imatong Heterohyrax brucei hoogstraali Feb i Nimule Acomys eeteotls Apr DISTRIBUTION IN THE SUDAN Blue Nile: Gebelein (El Jebelein) (King 1911,1915), "Blue Nile districts" (Archibald 1923). Kassala: Erkowit (King 1926). Kordofan: Nuba Mountains (Ruttledge 1930). Sudan Government collections contain King's Gebelein collections, numerous laboratory reared progeny, and Ruttledge's specimens from Delami in the Nuba Mountains. Note the Egyptian records below, most of which are from the Southeastern Desert near the Sudan frontier in that part of Egypt administered by the Sudan Government. DISTRIBUTION Argas brumpti is a tick of drier East African areas that has spread into South and Southwest Africa, into that part of south. eastern Egypt that is included in the Ethiopian Faunal Region, and some distance into the Western Desert of Egypt (Palearctic Faunal ee The distribution has been mapped by Hoogstraal and Kaiser 1956). NORTH AFRICA: EGYPT (Hoogstraal 1952A. Garnham 1954,1955. Hoogstraal and Kaiser 1956. Davis and Mavros 1956B. Schmidt and Marx 1956). EAST AFRICA: SUDAN (King 1911,1915,1926. Archibald 1923. Beery te 1930. Hoogstraal 1952A,1954B. Hoogstraal and Kaiser 1956). ETHIOPIA (as Somaliland) (Neumann 1907B,1911,1922. Nuttall et al 1908. Stella 1938A,1939A,1940). KENYA (Neave 1912. Cunliffe 1914B. Anderson 1924A. Warbur- ton 1933. Walton 1950B. Garnham 1954,1955. Heisch 1954F). UGANDA (Hoogstraal and Kaiser 1956). SOUTHERN AFRICA: OVAMBOLAND, SOUTHWEST AFRICA (Theiler, un published). UNION OF SOUTH AFRICA and BECHUANALAND (Bedford 1936). HOSTS Available records indicate that in nature larvae feed on liz ards and on a number of mammals inhabiting dry caves, lairs, and rock ledges. Nymphs and adults also attack lizards and almost any mammal that happens to stop near their retreat. Certain birds are acceptable as larval hosts in the laboratory but birds have not yet been found infested in nature. Larvae have been reared on guinea pigs and nymphs and adults on white mice. Larvae Animals on which larval A. brumpti have actually been found in nature are the following: Lizards: ama colonorum in the Sudan (Ruttledge 1930). Uro mastix ocellatus an ama a. spinosa in Southeastern Egypt (Hoog— Straal and Kaiser 1956. Schmidt and Marx 1956). Gerrhosaurus validus in Transvaal (Bedford 1936). The gecko Tarentola 4. anni aris in the Western Desert and in the Southeastern desert of Upper Egypt. Rock hyraxes: Heterohyrax brucei hoo straali (Equatoria Prov— ince record above), Procavia sp. in Egypt Weccete ea 1952B), and Procavia capensis burtoni in Southeastern Egypt (Hoogstraal and Kaiser icety 33 Rodents: Spiny mice, Acomys hystrella (Equatoria Province record above) and Aco oehieteas atus in Southeastern Egypt (Hoogstraal aiser 1950). Nymphs and Adults Specimens from the Kitui District of Kenya (Heisch 1954F) were determined by precipitin tests to have fed on porcupines and not on hyraxes, rats, or gerbils. Lizards and baboons were suspected as possible hosts. Subsequently, Garnham (1954) work. ing in the same area found blood corpuscles of lizards in re. cently fed ticks and noted that undigested corpuscles could be identified in the ticks at least a month after feeding. Garnham fed captive nymphs and adults on geckos and agamid lizards. Inasmuch as nymphs and adults feed rapidly, they are seldom found when the vertebrate host is examined. It may be assumed, however, that these stages probably feed on most of the larval hosts noted above. Walton (1950B) reported that Brumpt's argas attacks hyraxes and people who take refuge near hyrax dens in Kenya. Theiler's five female specimens from Ovamboland are from a mierkat, Cynictis nicillata cinderella. Africans of the Yatta Plains say that fers tick (kitunu) feeds on human beings, elephants, buffalo, elands, and giraffes, and that specimens may be found in dust where big game animals roll (Cunliffe 1914B). I have seen three adult specimens from a lion's lair near Pusa, Kenya (EMNH collec_ tions). In Ethiopia, Brumpt found A. brumpti near porcupine bur_ rows, and reported its bite on himself (Nuttall et al 1908. Brumpt's Precis). King's (1925) Sudan records are from sparsely vegetated areas containing caves and crevices in which many kinds of animals rest. Uganda hosts are the African porcupine and the rock hyrax, Procavia capensis meneliki (Hoogstraal and Kaiser 1956). Experimental Hosts King (1926) reared larvae on the bare skin of the head of guinea fowl. Larvae failed to engorge on man, dogs, cats, goats, pigeons, doves, sparrows, or bats, though some attached to spar. rows and pigeons. Nymphs and adults fed on rabbits and man. Ruttledge (1930) found no larvae on guinea fowl in the Nuba Mourn tains and believed that lizards are the favorite larval host there. = One Davis and Mavros (1956B) successfully reared larvae on guinea pigs and nymphs and adults on white mice. That the various stages of Brumpt's argas feed to an extreme degree of repletion on white mice and guinea pigs has been noted in Dr. G. E. Davis* and in ow laboratories. This phenomenon is exceptional in the genus Argas. BIOLOGY Life Cycle Ruttledge (1930) found larvae on lizards only in March, at the end of the dry season. Larvae taken in Egypt were attached to hosts in the winter and spring months of February, March, and April; but it should be noted that these are the only months during which we have extensively explored Upper Egypt, to which area A. brumpti is probably restricted here. At Khartoum, eggs were laid in the laboratory in March, April, and October; some of these hatched about a month later (King 1915). Females brood over their eggs until larvae emerge (confirmed by Davis and Mavros 1956), as has been observed for many argasid spe— cies (Hoogstraal 19528). Hosts on which King fed immature stages have been listed above. Cunliffe (1914B) reported egg laying from a single female as follows: 53 eggs between 99 and 106 days after emergence, 66 eggs between 118 and 125 days, 21 eggs between 152 and 156 days, and 18 eggs between 161 and 166 days; total 158 eggs. The female had fed on a fowl on the 12th, 17th, and 143rd day and had been fer. tilized on the 13th, 17th, 142nd, 158th, and 168th day. Larvae hatched from two egg batches 24 to 27 days after the eggs had been laid but refused to attach to a fowl and died. King (1915) ob served that larvae do not feed readily until they are about ten days old. Subsequently, eggs were laid in the laboratory in batches of about seventy eggs, about a monthapart, after each feeding, through the cooler months of the year in Khartoum (King 1926). One male fertilized at least three females. Four nymphal stages were ob. served. A certain mature female collected in July, 1918 was still alive in December, 1926. 5 re Recently, Davis and Mavros (1956B) have been most successful in rearing larvae on guinea pigs and nymphs and adults on white mice. Between feedings, ticks were held at 30°C. in a humidified cabinet. The life cycle under these conditions required from four to eight months. Larvae fed from six to fifteen days, most— ly for eleven to thirteen days, and molted to nymphs ten to twelve days later. This lot all reached the adult stage after three nymphal molts, as follows: first to second instar, fourteen to twenty days; second to third instar, seventeen to 32 days; third instar to adults, 21 to 37 days. One third nymphal instar specimen fed twice before molting to an adult male. This individ ual required 140 days postlarval feeding to reach adulthood while the most rapid time for this interval, also for a male, was ninety days. In an additional lot, six males and fourteen females required three nymphal instars and two females underwent a fourth nymphal molt. Three pairs fed twice in the last nymphal instar. The last nymphal instar lasted from 26 to 65 days, but most required only 26 to 33 days. Females fed for eight to 21 minutes and mated for ten to 27 minutes. Oviposition commenced eight to 58 days after feeding and larvae hatched after eight to 24 days. The number of eggs per batch ranged from 24 to 96, and while generally smaller batches showed a higher rate of fertility than larger batches this was by no means consistant. For instance, in one batch of 24 eggs 22 larvae hatched, one batch of 72 was entirely fertile, a batch of 62 was about fifty percent fertile, and the batch of 96 eggs hatched 81 larvae. The least fertile female oviposited a second time without a second feeding or mating. The most fertile fe male fed a second time and then oviposited though without having mated since the first feeding three months previously. Further observations showed that while larvae take a small amount of blood within an hour of attaching to the host, the bulk of the blood is ingested shortly before completing feeding. Molt— ing ticks free themselves from the exuvia within a few minutes after the skin splits. Mating and feeding takes place readily in daylight. Feeding adults stand perpendicularly, bracing themselves against the tube; when partially distended they fall over on their dorsum to complete engorgement. = 16% Ecology A. brumpti lives in areas of low rainfall (Ovamboland, Kala. hari, Nuba Mountains, Egypt) or in dry niches within areas of comparatively high rainfall (Kenya, southern Sudan, Transvaal). It rests in caves, lairs, or dens, where it feeds on either permanent mammal residents or on visiting mammals or lizards, or it may hide under rocks from whence it emerges to feed on passing lizards or mammals. The original Ethiopian specimens were found in dust under rock ledges in a dry streambed visited by porcupines (Nuttall et al 1908. Brumpt'!s Precis). It was said that these adults wandered about at night but not during the day. King (1915) found specimens in caves and crevices where they were living among debris of soil and rotting leaves and twigs. One of Bedford's (1936) individuals was hiding under a stone. In Southeastern Egypt we have found from one to 23 nymphs and adults together a millimeter or two below the surface of fine, dry sand in small caves and in holes in rocky hillsides. These areas range from barren desert bordering wadis with a little vegetation to arid parkland at the base of mountains facing the Red Sea. In the lowlying country north of Kitui, Kenya, Heisch (1954F) observed immobile specimens on fine, brown earth under large boul. ders on a peneplane of red, sandy soil covered by thornbush. Dr. Heisch has informed me (conversation) that the loose soil below these rocks is probably too sparse for burrowing by the ticks. He and Dr. Garnham, who accompanied him on a collecting trip to this area, have also told me that the ticks rest in rock crevices or between rock layers where there is little or no accumulation of soil. It is of interest to learn from information furnished Cun. liffe (19148) for other parts of East Africa that Brumpt's argas burrows in the dust of termite mounds where large game animals moll Biologically, therefore, in its feeding and resting habits, - brumpti appears to show tendencies towards certain intermediate ans between typical Argas ticks and typical Ornithodoros ticks. anaes ticks in all Stages are considered typically as fairly host-specific parasites of birds or of bats and as ticks that nor- Ola mally hide in crevices but do not burrow in sand or soil. Orni. thodoros ticks, on the other hand, usually parasitize mammals, although exceptions are known, and unless the soil is too hard, they normally burrow below the surface or at least hide in cracks in soil. It is, therefore, of interest that larval A. brumpti feed, apparently more or less indiscriminately, on reptiles, birds, and mammals, and that nymphs and adults attack either liz. ards or mammals. A further interesting observation, from both Dr. G. E. Davis® and our laboratories, is that the various stages of this tick feed fully on white mice, whereas most other Argas species feed only partially on mice. When resting, A. brumpti apparently prefers to burrow in soil or sand, but if this is not possible it hides among surface debris or between or under rocks. King (1926) mentioned the possibility that Brumpt!s argas might infest human dwellings although actual records had not been obtained. Archibald (1923) reported Brumpt's argas among a col. lection from in and around human dwellings, but he did not spec. ify exactly in which situation specimens were taken. Archibald's records and specimens cannot now be located. Walton (1950B) states that in Kenya this species does not occur in human huts. DISEASE RELAT IONS MAN: Bites of A. brumpti may be quite painful to human beings and cause itching lasting for years. Kenya natives claim that this tick causes pain and sickness when it bites man. LIZARDS: In Kenya and in Egypt, A. brumpti and lizards have been found infested with a hemogregarine, Hepatozoon argantis Garnham, 1954. HYRAXES:; This tick should be considered as a potential vector of the piroplasm Echinozoon hoogstraali Garnham, 1951, a parasite of Heterohyrax brucei hoogstraali. REMARKS The diagnostic characters listed below comprise the criteria for Pospelova.Shtrom!s (1946) genus Ogadenus, herein considered as a subgenus of Argas. Warburton's (153) views on the generic position of A. brumpti will be reviewed in subsequent studies of the genus. Sis) Schulze (1936G) compared the body outline of A. brumpti with that of certain fossil trilobites that it happens to resemble. IDENTIFICATION Neumann's original adult specimens from Ethiopia measured from 15 mm. to 20 mm. long and from 10 mm. to 13 mm. wide. The size range among available Sudan and Egyptian females is from 10 m. to 12 m. long and from 7 m. to 8 mm. wide. The dorsal integument is marked most distinctively by large, symmetrical polygonal depressed areas bounded by rugose ridges; laterally the dorsal integument is evenly striated. A continuous, clear, and distinct sutural line divides the dorsal and ventral sur. faces. Discs, which are small and inapparent, lie in clumps or lines in the integumental depressions, and short hairs are scat— tered anteriorly and posteriorly on the periphery of the body. The outline of the body is subquadrangular with parallel lateral margins, a bluntly rounded posterior margin, and a pointed pro- jection of the anterior margin over the mouthparts. Tarsus I has two dorsal humps and the other tarsi each have a prominent sub apical dorsal protuberance. Males have a narrow, rounded genital aperture; females have a transversally elongate, narrow genital orifice. The nymph closely resembles the adult, except for size and absence of genital aperture. The larva has been described by Cunliffe (19148) and by Hoogstraal and Kaiser (1956). = SO9ke Figures 33 and 34, 9, dorsal and ventral views ARGAS (CHIROPTERARGAS) BOUETI Egyptian specimen PLATE XI |= SO= ARGAS (CHIROPTERARGAS) BOUETI Roubaud and Colas—Belcour, 1933. (Figures PBN tp SINe and 34) THE LONG.LEGGED BAT-ARGAS. i ee rwte : EQUATORIA PROVINCE RECORDS jf eal Sunat Taphozous rforatus haedinus Feb Ti Torit Rhino lophus Tobatus Jan These are the only records of this species from the Sudan. DISTRIBUTION Argas boueti is known from scattered localities in Africa as far a as Transvaal and is also present in the Near East. A tick apparently of drier areas, which has been widely spread by its chiropteran hosts, the long-lepeed bat-argas is obviously more frequent than present meagre records indicate (Hoogstraal 1955B). Whether North Africa, the Near East, or the Ethiopian Faunal Region is its origin is difficult to determine from evidence at hand. NORTH AFRICA: EGYPT (Hoogstraal 1952A,1954A,1955B). WEST AFRICA; FRENCH WEST AFRICA (Roubaud and Colas—Belcour 1933. Hoogstraal 1955B). CENTRAL AFRICA: FRENCH EQUATORIAL AFRICA (Hoogstraal 1955B). EAST AFRICA: SUDAN (Hoogstraal 1952A,1954B,1955B). KENYA (Heisch 1951B. Hoogstraal 1955B). SOUTHERN AFRICA: ANGOLA (Hoogstraal 1955B). UNION OF SOUTH AFRICA (Hoogstraal 1955B. Subsequently, Dr. Zumpt has sent me additional specimens from Tzaneen, Transvaal). NEAR EAST: Palestine (Hoogstraal 1954A,1955B). o Blea HOSTS Bats Asellia (= Hi siderus) tridens and Taphozous perforatus (Roubaud and Colas-Belcour 1933). Taphozous p. perforatus, T. (Liponycteris) nudiventris, Hee hardwickei cystops, and Otonycteris h. hemprichi are mos eavily infested in Egypt. Less frequent hosts here are Rhinopoma pero Asellia t. tridens, Rhinolophus clivosus brachygnathus, Re. me elyi, Roussetus ae aegypticus, Nycteris t. the ca, arida a. ae Tacasme Teniotis ruppelli, Pipistrellus k. kuhli and Plecotus auritus christei (Hoogstraal 19528,1955B). Megaderma cor (heisch 19513). Rhinolophus lobatus (Sudan record above). Eptesicus tenuipinnis (Hoogstraal 19555). Otonycteris h. hemprichi, a bat which roosts in small caves, crevices a niches, usually singly or with very few other of the same species, is most heavily infested in Egypt. Rhinopoma h. cystops, one of the most common cave-inhabiting bats near Cairo, is frequently heavily infested and probably represents the most important host in this area owing to its great abundance. All stages appear to feed on the same kinds of bats. Man Nymphs and adults have bitten us in caves on a few occasions. They readily do so when allowed to in the laboratory (Hoogstraal 1952A,1954A,1955B). BIOLOGY Life Cycle Rearing of A. boueti has been accomplished in our laboratories at temperatures of from SO°F. to 90°F. with relative humidity ranging from 406 to 508. Exceptionally large females may lay single egg batches of almost two hundred eggs over a two or three day period. An average size female deposits from 35 to 40 eggs in a single batch usually on a vertical surface. Afterwards, she ieee: stands motionless over or next to the eggs for fifteen to 22 days until the larvae hatch. (Earlier, at unrecorded high summer room temperatures, we obtained hatching in eleven to fourteen days). Larvae attach to a host after twelve to fifteen days (earliest host offered), and feed from eight to 42 days, but mostly from sixteen to 25 days. Afterwards, larvae may require from four to thirteen days before molting to first instar nymphs, but they usually do so after four to seven days. Nymphs molt two or, uncommonly, three times before reaching adulthood. In our laboratory, those nymphs that reach adulthood after two molts have never fed in the nymphal stage, even ug bats were frequently offered. When a rhea instar nymph, which has not molted to an adult, feeds it does so for half an hour to an hour from seven to 26 days after the previous molt and becomes an adult from twenty to 32 days after feeding. The duration of each nymphal instar is seven to seventeen days for the first in. star, with eleven to fourteen days the most common, The duration of the second instar is longer, from sixteen to 43 days, with 22 to 29 days average. The duration of the third stage, when it oc- curs, is erratic and lasts from 27 to 58 days. No significant data on sexual differentiation from the unusual third nymphs have been obtained. Adults feed for thirty to 35 minutes beginning some five days after molting. Further studies on the life cycle of progeny are under way. A biting tick remains motionless during feeding. It often stands the full length of its anterior legs away from the point of insertion of mouthparts that are extended by a pendulous tube from the basis capituli. Once the hypostome is inserted, the host's hand or arm (or the bat) may be moved freely till the tick is satiated without causing it to remove its mouthparts. When blood is rapidly engorged a large drop of clear coxal fluid ap. pears beneath the body, but none is emitted during slow feeding. Repletion from the human hand or arm requires from 25 to 35 minutes but full engorgement from the membrane of a bat's wing may require three or four hours. Individuals that feed slowly become very lethargic and one may remove them, even though fully fed, after seven to 24 hours with the mouthparts still inserted in the wing membrane. Itching at the site of the bite on man may persist for several weeks. = Bic When disturbed the long-legged bat-argas is much more active and moves with greater speed that either A. confusus or A. vesper- tilionis. The long anterior legs wave up and down While walking ap objects antenna.fashion. This tapping is especially active just before the mouthparts are inserted. During feeding the anterio: legs may or may not touch the host skin but they seldom function as a support for the tick. These ticks will feed in light or in dark ness but prefer darkness. Ecology The original authors of A. boueti described this species from material collected about 1910 in hollow trees inhabited by the two species of bats listed above. In the generally arid Northern Prov. ince of Kenya, Heisch found adults on walls of an underground con. crete shelter and larvae on Megaderma cor in the same structure. Those South African specimens with collecting data are from houses. In Egypt we commonly find the long-legged bat-argas thriving under the most severe desert conditions. It is less common in more humid buildings inhabited by bats in Cairo. A. boueti is the most numerous bat-parasitizing argasid in Egypt and the same eco. logical observations noted for Egypt under A. vespertilionis apply to this species. ad The frequency with which these long-legged ticks fall from rough surfaces in the laboratory is surprising in view of their usual habitat on walls and on ceilings of caves and chambers. A number of specimens exhibit a body tremor that causes them to flip over on their dorsal surface with every few steps. Righting move- ments require considerable effort. DISEASE RELATIONS MAN, Attacks on our laboratory personnel have caused mild itching persisting for several weeks in warm weather. BATS. We have been unable to find spirochetes in Egyptian material. Collections of local specimens injected into laboratory animals have resulted in negative findings for viruses and rickett. siae. Other material from the Cairo area has not yielded Shigella organisms. The hosts from which the type series was ee iietted in French West Africa showed a trypanosome infection but research to ascertain the relationship of ticks and trypanosomes was not under- taken. Ss REMARKS For a discussion of long-legged parasites of bats, see REMARKS under Ixodes vespertilionis,. A. boueti populations consist of two size groups, the larger about 6.8 mm. long and 6.9 mm. wide and the smaller about 5.0 m. long and 4.5 mm. wide (Hoogstraal 1955B, figure 125). The signif— icance of these "races™ is at present being studied. The subgenus Chiropterargas Hoogstraal, 1955(B), of which A. boueti is the type species, contains one other species, A. confusus. This subgenus is defined as follows: “Parasites of which bats are hosts of predilection. Morpho— logical characters intermediate between those of typical Argas and typical Ornithodoros; with a general Argas facies but lacking a sutural line; with a flattened body flange but lacking "cells™ and with exceedingly slight integumental differentiation at periphery; body shape circular to transversely elliptical. Integument with fine, close granular projections; discs mostly small, conspicuous, radially distributed, ventral "paired organ™ present. Definite hood over Wouthparts; mouthparts about level with anterior body margin. Legs of variable length, arising from anterior half of body; tarsal humps lacking.™ IDENT IFICATION The remarks above include the outstanding characters for iden tifying this species and A. confusus. The male of A. boueti is somewhat smaller than the female and has a semicircular genital aperture while the female has a narrow, transversely elongate genital aperture bounded by two rugose lips. In both sexes, the body outline is subcircular to pearshaped (definitely wider than long in A. confusus), leg IV extends far beyond posterior margin of body (only slightly beyond in A. confusus), basis capituli and palpal segments comparatively narrow and elon gate (globose in A. confusus), minute integumental protuberances are mostly flat (mostly tapered in A. confusus), etc. The dental forma of both species is 1/1, the apex of the hypostome is slight— ly indented, and a corona is fen: > 85S The larva and nymph have been described by Roubaud and Colas. Belcour (1933) and in more detail by Hoogstraal (1955B). = 965 Figures 35 and 36, 9, dorsal and ventral views ARGAS (CHIROPTERARGAS) CONFUSUS Egyptian specimen PLATE XII ARGAS (CHIROPTERARGAS ) CONFUSUS Hoogstraal, 1955(B). (Figures 25,26,35, and 36) THE WIDE BAT-ARGAS Ne OC EQUATORIA PROVINCE RECORDS 2 Torit Chaerephon major Dec a) Torit *Eptesicus pusillus - (svs) 1 Latome *Pachyotus sp. Mar (SVS) 3 Sunat Taphozous perforatus haedinus Feb DISTRIBUTION IN THE SUDAN As A, vespertilionis (in part): Khartoum and Northern Prov. inces (King i 5 The following lots are in Sudan Government collections: Khartoum: Khartoum. Larvae from unidentified bat. Northern: Dongola. Larvae from unidentified bat. DISTRIBUTION A. confusus is recorded from scattered localities from Egypt to the Cape of South Africa. Additional collecting will undoubtedly reveal its occurrence elsewhere on the continent. This species is thus far not known outside of Africa. NORTH AFRICA: EGYPT (Hoogstraal 1955B). EAST AFRICA: SUDAN (In part as A. vespertilionis: King 1911, 1926. As Le vespertilionis group: Hoogstraal 19543. As A. confusus: Hoogstraal 1955B). KENYA (Hoogstraal 1955B). *Host name on labels; identity not checked by authority in host group. = 9S = SOUTHERN AFRICA: SOUTHERN RHODESIA, BASUTOLAND, and BECHUANA~ LAND (Hoogstraal 1955B). UNION OF SOUTH AFRICA [For details con cerning A. confusus reported as A. vespertilionis by Nuttall et al (1908), Howard (1908), and Bedford (1 os 7), and for more recent rec- ords, see Hoogstraal (1955B). Subsequently, Dr. Zumpt has sent additional specimens from Bloemfontein, Orange Free State, and from Lady Frere, Cape Province./ HOSTS At the present time we have definite evidence of larval A. confusus from only a few species of insectivorous bats; Chaerephon major and Taphozous perforatus haedinus (Equatoria records above); questionably (host field determinations not checked by a specialist) from Eptesicus pusillus and Pachyotus sp. (Equatoria records above); Pa and from Eptesicus capensis, Pachyotus sp., Miniopterus natalensis arenarius, Taphozous p. perforatus, ae Gs nudiventris, Otonycteris h. hemprichi, Nycteris t. thebaica, and Tadarida a. aegyptiaca THoogstraal igSBD- Nymphs and adults found in bat-infested caves and buildings probably feed on the same species of hosts as do larvae. The record of A. confusus (= A. vespertilionis) attacking pen. guins in Queenstown, Cape Colony (hatter et al 1908) mst be re garded as questionable (Hoogstraal 1955B). BIOLOGY Life Cycle A. confusus has been reared in our laboratory at temperatures between SOF. and 90°F. and at relative humidities between 40% and 506. A single egg batch consists of from forty to seventy eggs With fifty to sixty the most common quantity. Eggs hatch from 21 to 25 days after being laid. Larvae have commenced feeding five to 26 days after hatching. The duration of larval feeding varies from five to fifty days but most larvae drop from the host in two or three weeks. Afterwards, larvae remain quiet for seven to twelve days before the nymphal molt. - 99- Four nymphal instars are invariable in our numerous observa. tions. Nymphs, like adults, feed for from 25 to fifty minutes. The first nymphal instar, however, never feeds; it molts to the second instar usually in ten to thirteen day Ss (range eight to 21 days). One to three weeks later the second ice nymph feeds and molts to the third instar some two weeks later (range eight to 23 days). The third instar nymph takes food between three and 35 days afterwards and molts three or four weeks later (range fifteen to 59 days). The fourth instar nymph feeds between five and 36 days afterwards and molts to the adult stage three or four weeks later (range seventeen to forty days). Note that althoush there are many morphological similarities between A. confusus and A. boueti, the biology of the nymphal stage of each is distinct. In this species, the observed period between nymphal molting and feeding is quite variable. Some accept food within five to seven days although two to three weeks, or longer, is more common for this and for other bat-parasitizing argasids. Also noteworthy is (1) the average two week postfeeding period of second instar nymphs in contrast to the three or four week average postfeeding period of the two subsequent instars, and (2) the nonfeeding first instar. Adults placed together shortly after molting have been observed to mate only after feeding which may occur from eight to sixteen days after molting. The feeding period varies from forty minutes to two hours. No egg batches have been deposited before three and a half to four months after molting. Further studies on the Fy, generation are in progress. Ecology We have collected, with considerable and strenuous effort,-. several hundred specimens of A. confusus in Fgypt. This species, together with A. transgariepinus, is the most secretive of bat im festing argasids. It wedges itself deeply into the narrowest crevices of caves and of hillside crannies in which bats rest. It is never found easily or in groups of more than two to a max imum of twelve specimens. We know of only a single exceedingly small population in Cairo, where bats roost in buildings that are more humid than desert caves. In the environs of Cairo, A. 1KG0) = confusus is scattered throughout desert and desert-edge retreats, all arid, such as antiquities structures, caves, and hillside crannies. Records from northern Sudan and from the Protectorates of South Africa indicate a more or less similar tolerance of aridity, but those from various regions of South Africa, Torit, and the crater of Mt. Menengai in Kenya indicate also that certain popu. lations exist in markedly humid environments where they tolerate lower temperatures and higher relative humidity than they do in Egypt. DISEASE RELATIONS BATS: A small number of specimens thus far studied in NAMRIL3 laboratories have been negative for blood protozoa, spirochetes, viruses and rickettsiae, and Shigella organisms, REMAR KS Further studies: on the habits and ecology of this species are presently under way and will be reported when completed. For a definition of the subgenus Chiropterargas and for criteria to dis. tinguish this species from A. boueti, see page 95. IDENT IFICATION A. confusus adults have an extremely wide body outline, and, in common with A, boueti, are characterized by the absence of a sutural line dividing dorsal and ventral surfaces, and by the pres— ence of a conspicuous hood over the mouthparts. In A. confusus the dorsal integumental protuberances are fine, shiny-tipped, tapering points which on the lateral margin are more closely spaced and more regular. The posterior discs are arranged radially; the legs are shorter than the body length; and the hypostome has only a single pair of denticle files. The tarsi have no dorsal protw. berances. A pair of grooved organs of unknown function is present just posterior of the anus on the ventral surface. = Ol Except when greatly engorged, the peripheral flange of the body remains partly unfilled. In partially engorged individuals this flange is flat, and in dry specimens it may be turned up like a rim. The body color is reddish yellow with a central, darker area of varying extent. Males measure from 5.9 mm to 6.4 mm long, and from 7.4 mm. to 7.8 mm. wide (average 6.1 mm. long and 7.5 mm, wide). The genital aperture forms a wide arc. Females are larger, and measure up to 8.0 mn. long and 9.5 mm. wide. The female genital aperture is a transverse groove with thick, rugose lips. The nymph and larva have been described by Hoogstraal (1955B). The larva first instar nymph are quite similar to those of A. boueti but the successive instars of each resemble the associated adULtS . 5 10k S a f. Pn LO) es "Ning Figures 37 and 38, o&, dorsal and ventral views ARGAS (CARIOS) VESPERTILIONIS Egyptian specimen PLATE VIIT = 1037— ARGAS (CARIOS) VESPERTILIONIS (Latreille, 1802). (Figures 21,22,37, and 38) THE ROUND BATARGAS N @Q ron EQUATORIA PROVINCE RECORDS Lokwi Rousettus aegyptiacus June (CNHM) Katire *Mimetellus ?moloneyi Sep Sunat Taphozous perforatus haedinus Feb orn DISTRIBUTION IN THE SUDAN Northern; One larva from an undetermined species of bat at Dongola, L- April 1917, Bedford legit; Sudan Government collections. Khartoum: Several larvae from an undetermined species of bat(s?) at Khartoum, 20 September 1914, R. Cottam legit; in Sudan Government collections, one retained in Hoogstraal collection. DISTRIBUTION The Argas vespertilionis group, consisting of A. vespertilionis (Latreille, 18 in Europe and Africa, A. pusillus Kohls, 1950, on Palwan Island in the Philippines, and of numerous closely related forms of yet uncertain species status, ranges throughout the conti. nents and island groups of the world, except in the Americas. It is possible that certain African populations presently identified as A. vespertilionis will prove to be separate, closely related species. i fuller study of this group is under way. African Records Only Eventually, the round bat-argas most likely will be found in many more territories of Africa. “Field identification of host; specimen not seen by a specialist in bat taxonomy. =) INO = NORTH AFRICA: EGYPT (As A. fischeri: Audouin 1826, Savigny 1827, and Lavolpierre and Riek 1955. Hoogstraal 1952A). TUNISIA (Colas-Belcour 1933B). WEST AFRICA: FRENCH WEST AFRICA: Although reported as A. vespertilionis by Marchoux and Couvy (1912A,B,1913A,B), there is Some likelihood that some or all of these specimens may have been those subsequently used as the types of A. boueti. Rousselot (1953B). GOID COAST (Simpson 1914). ~ CENTRAL AFRICA: BELGIAN CONGO (Schoenaers 1951A). EAST AFRICA: SUDAN (King 1911,1926; in part. Hoogstraal 1954B). KENYA (HH collecting in crater of Mt. Menengai). SOUTHERN AFRICA: ANGOLA (Larvae from Dundo, Lunda, north eastern Angola, CNHIM). MOZAMBIQUE ("Brumpt's Precis”). SOUTHERN RHODESIA (Jack 1932. Bedford 1934). UNION OF SOUTH AFRICA: South African adults described and illustrated by Nuttall et al (1908) as A. vespertilionis represent A. confusus. Howard (1908), D8nitz (STOsy hedtord (15 Bedford (1932B,1934), Also confused these two species as probably also did Cooley (1934); ef. Hoogstraal (1955B, p. 586) for details. Dr. G. Theiler has sent a female and nymph of A. vespertilionis from Pretoria and Grahamstown. These were among Larger numbers of A. confusus and A. boueti. No specimens of A. vespertilionis were included with Nraterial of A. confusus and A. boueti from collections of the South African Institute for Medical Research, recently sent for identification by Dr. F. Zumpt. These observations lead one to suspect that A. vespertilionis may be less common in South Africa than A. confusus. OTHER AREAS: Available material referable to this group is from England, Netherlands, Sweden, Spain, Germany, Korea, China, Philippines, and Ceylon. The group is also known to occur in southern India, Cambodia, Australia, France, Italy, and Russia. Differences between African and European specimens and those from Australia and Asian areas are very slight indeed. — 105) HOSTS (Africa) Bats Almost any bat, whether it lives in large colonies or in small groups, may be parasitized by A. vespertilionis. All stages prob. ably infest the same kinds of hosts. Three species of chiropteran hosts are thus far known from the Sudan (records above). The Angolan larva is from Pipistrellus nanus. In Egypt, we find larvae chiefly on Otonycteris f. hemprichi, Rhinopoma hardwickei cystops and R. microphyllun, Taphozous p. perforatus and ais nudiventris, Nycteris t. thebaica, Tadarida a. aegyptiaca and T. teniotis ruppelli. They are less numerous on Rhinolophus clivosus brachygnatus, R. mehelyi, Asellia t. triedens, Plecotus auritus christiei, Pipistrellus k. kuhli, and Rousettus ra aepyptiacus. /Egyptian bats reviewed by Sanborn and Hoogstraal igh ic Man Nymphs and adults on several occasions have attacked us in caves and we easily induce it to bite ourselves in the laboratory (Hoogstraal 1952A,1955B). BIOLOGY Life Cycle In our laboratory, Mr. Sobhy Gaber successfully rears spec— imens of A. vespertilionis at 80°F. to 90°F. and 40% R.H. to 506 R.H. Egg batches consist of thirty to fifty eggs, one-fifth or one-sixth of which usually do not hatch. Larvae emerge from sixteen to twenty days after the eggs are laid and some will feed as quickly as four days afterwards. The duration of larval feed ing varies from fourteen to 31 days, but is usually seventeen to nineteen days. Five to ten days later larvae molt to nymphs, which are capable of feeding three or four days after this and after subsequent molts. Usually two feedings are indulged by nymphs, followed by a molt eight or nine days after the first meal and twelve to fourteen days after the second meal. Nymphs = 106 = become replete in from twenty to fifty minutes, usually in thirty to forty minutes. Males may emerge from the first nymphal molt, but usually nymphs molt twice before becoming adults. Males and females may feed within seven days after molting. Duration of adult feeding is thirty or forty minutes. No female has ovi- posited within six months after the nymphal-adult molt, even though she has been with a male continuously and both have had two to six blood meals. The first egg batch follows a blood meal by about a week. The first oviposition appears to trigger a physiological release mechanism for, in several instances, three months after- wards females have deposited a fertile egg batch with or without a meal. We are at present attempting to ascertain whether the long interval between molting and oviposition is peculiar to these laboratory observations or whether it is a usual feature in our local populations. A. vespertilionis is more lethargic than A. boueti. Adults, if undisturbed while imbibing from a vein in the wing membrane of a bat, may remain attached for as long as five hours after engorgement is apparently completed. The feeding tick remains motionless with all legs down but, when fully distended without release of mouthparts from the host skin, it usually raises the fore legs to an antennalike position. During engorgement the beak is disengaged from the host skin only after considerable disturbance. Large blood clots form at the site of the bite, both on the bat's body and in the wing membrane. This phenomerion, on bats, is in marked contrast to that observed by Lavoipierre and Riek (1955), using ticks from our collections, and laboratory rodents. The greater avidity with which these ticks attack bats probably accounts for the more conspicuous sequelae in these animals. Larvae may be found anywhere on the body or wing membranes, but most commonly at the edge of the hairy parts, seldom on the head, feet, tail, or trailing edge of the wings. Ecology European and African populations of this tick, which thus far cannot be morphologically differentiated, withstand a wide - 107 - range of temperature and humidity conditions. Host flight habits account for the wide distribution of A. vespertilionis, but we are not aware that host migration is a factor 1n mixing populations from widely differing ecological situations. European and South African populations exist under temperate climatic conditions with pronounced seasonal changes and with moderate to heavy rainfall. Those of Egypt and of northern Sudan normally tolerate the most extreme arid niches in which any arthro. pod is known to survive. Their engorged larvae, however, are found usually among moist dung or in dung between crevices of bats' retreats. Just where females commonly oviposit in nature and where unengorged larvae rest before seeking a host has not yet been satisfactorily determined Throughout Europe and Africa interstices in the walls of bat— infested caves and buildings are the most common habitats of A. vespertilionis. They may also be found in tree holes and in other Situations frequented by certain bats. In Cairo a specimen, re- calling Robert Burns* wee louse, has been taken from a worshipper during church service by an observant but distracted friend sit— ting behind. In Iraq, Patton (1920) reported the same or a closely related species in Bedouin tents in which bats presumably rested by day. Egyptian specimens hide alone or clustered in large or small groups usually well concealed between shale or in crevices of walls. Some individuals are observed wandering openly on the walls. Unconcealed individuals are noted much more frequently in those caves or niches that only erratically harbor a few bats than in large caves where many bats usually roost. Possibly our entry into caves infrequently visited by any animals induces these ticks to investigate the possibility of a meal. Small numbers of the round bat-argas frequently are found in niches in the most unexpected cliffsides where a few old pellets of dung indicate that hermit bats such as Otonycteris h. hemprichi occasionally spend the day. These ticks lead a most uncertain existence and often wait months on end for a host, as revealed by their compressed bodies and by the age and scarcity of hosts! dung in these places. = il In Egypt, A. vespertilionis occurs with the more common A. boueti and with the less common A. confusus and A. trans ariepinus. &. vespertilionis and the other two Species are rare, however, ‘in the comparatively humid situations in Cairo favored by the fruit-bat parasite, Ornithodoros salahi. DISEASE RELATIONS MAN: Mild itching resulting from a bite may persist for several weeks. BATS; Large blood clots form at the site of the bite, both on the host's body and on the wing membrane. It has been stated that this tick is a vector of a spiro- chete of bats but reports of conclusive supporting evidence have thus far not been located. In the Cairo area blood of a few of these ticks has been found to contain a most interesting organism resembling, accord— ing to Dr. P. C. C. Garnham (correspondence): "the sporozoites of a Haemoproteid; they are not unlike the sporozoites of Leuco— cytozoon ...e. described ..... from the abdominal cavity o imulium flies.” Unfortunately, it has thus far been impossible to undertake further study of this phenomenon. Egyptian specimens examined in NAMRU_3 laboratories have been negative for spirochetes, viruses, rickettsiae, and Shigella organisms. REMARKS The taxonomy and biology of bat-infesting Argas ticks is presently being studied and the first report, on k. boueti and A. confusus has been presented (Hoogstraal 1955B)e The second Section will deal with the confounded status of Argas (Carios) rtilionis. Some workers have considered Carios as a full ee indiscriminately including in it features of a variety of species based on vague and ambiguous remarks in the literature. koe The presence of the ventral paired organ related the subgenus Carios to the subgenus Chiropterargas and separates Carios from the subgenus Argas and from other subgenera. The presence of a lateral suture = of peripherally differentiated integument, and the absence of an appendagelike hood clearly separates the subgenus Carios from the subgenus Chiropterargas. The name vespertilionis was assigned by Latreille in 1802, not in 1796 as stated by most authors. The reference to this name as of 1796 (Hoogstraal 1955B) derives from an editorial change. Schulze (1943B) noted that the immature stages of species of this group (A, uel as have especially highly devel- oped terminal branching of midgut diverticula but little basal branching. IDENTIFICATION Males and females are alike except that males average some what smaller in size and their genital aperture is semicircular in outline, rather than narrowly ovoid, and is not bounded by thickened rugose lips as in females. In both sexes and in nymphs, a definite lateral suture en circles the body, a dorsal and ventral row of rectangular "cells" marks the body periphery, no appendagelike hood is present over the mouthparts, but ventral paired organs are present just pos terior of the anus. The body outline is generally circular or subcircular, but may be somewhat longer or wider in some specimens. Few individuals reach six millimeters in length or breadth. The integument is smooth, marked by a fine network of small, irregular cells among which regular, subparallel rows of larger discs radiate. Legs arise from the anterior half of the body and are short— er than the body; coxae are contiguous; and tarsi are tapered and lack dorsal humps. LO Mouthparts are situated close to the anterior margin of the body. The hypostome formula is 2/2 to 2.5/2.5, the apex is notched and bears a corona of three or four rows of small denticles in four to six files. The larval and nymphal illustrations and descriptions of A. vespertilionis iy ame 1896) are quite good, though they shall have to be expanded for present day purposes. These have been employed subsequently by the same author, Nuttall et al (1908), Bedford (1934), and others. > al = Jit ARGAS SP. (Not illustrated) A single damaged larva, mounted on a slide, sent by Mr. E. T. M. Reid of the Sudan Veterinary Service, represents an unm known species of tick. Although many of the characters of this specimen are obliterated, enough are preserved to indicate dis. tinct differences from A. vespertilionis, to which it is probably more or less closely allied, and from all other described species. This larva was collected from a Pachyotus bat at Latome, Equatoria Province, on 16 March 1951 by Mr. J. Owen. eee = ORN | THODOROS* INTRODUCTION Both important African species of Ornithodoros, 0. moubata and 0. savignyi, occur in certain areas of the Sudan, where they are known as haim ( -—~4 ). From two to five other species indubitably exist in the Sudan but have not yet been found there. Approximately fifty species comprise the genus throughout the world. O. moubata is the most important tick vector of relapsing fever in Africa and its bite is often painful. 0. savignyi has been sus— pected to be a relapsing fever vector, although incriminating evi dence in nature is negative or unconvincing. Ornithodoros ticks are thick, leathery, and podlike. They may be more or Less difficult to find but pain when they bite signifies their presence. In contrast to Argas ticks, which usually parasitize birds and bats, most Ornithodoros ticks parasitize mammals, including bats, and only exceptionally attack birds, reptiles and ampnibians. The two species under consideration represent a somewhat more advanced stage in evolution of parasitism than do Argas and other Ornithodoros species in that their larvae remain in the large, leathery egg until ready to make the larval_-nymphal\molt. The safety of the tough egg capsule affords delicate larvae considerable protection from the elements. 0. moubata and O. savignyi are un. usual in this respect; larvae of most other Ornithodoros species are active and feed from animal hosts. Shortly after hatching larvae molt to nymphs that soon set out to find a host. Nymphs and adults feed rapidly, in a matter of a few minutes to an hour or two, and are seldom transported while feeding on the host. They are, there— fore, usually found only in their resting places. Ornithodoros *Some writers replace the os ending used here by us. The original name used by Koch (1844) was spelled with an os ending, and this is generally though not universally conceded to conform to the rules of nomenclature and of philology. This question has been reviewed by Najera (1951). - 113 ticks are able to withstand long periods of starvation and are very resistant to aridity. The two species discussed in the following pages have been widely spread along man's trade routes. Subgenera of Ornithodoros are scarcely better established than those of Argas. 0. moubata and 0. savignyi are closely re lated species with only one other kindred kind, 0. eremicus Cooley and Kohls, 1941, of Utah, western North America. Some workers would include only these three species in the genus Ornithodoros and all others in several different genera. This position seems an unnatural approach and a useless complexity. For present pur poses, 0. moubata and 0. savignyi are treated as in the subgenus Ornithodoros, the only one presently known to be represented in the ane In addition to these two tampans, it is certain that several other species of Ornithodoros occur in the Sudan and await dis_ covery. Among these should be 0. foleyi, 0. delano#i subsp., and possibly some member of?the 0. tholozani group. When more inten sive search is undertaken in the Sudan it will probably be found that 0. erraticus is localized but widely distributed here. 0O. erraticus 1S broadly characterized by small size (maximum length 7.5 mm™., usual length 3.0 to 5.0 mm), oval shape, closely crowded, hemispherical granulations interspersed by large discs, and absence of tarsal armature and of cheeks surrounding the mouthparts. Spec- imens will probably be found in small mammal burrows. The following are the other known Ornithodoros species that occur in Africa, with selected references concerning them: 0. arenicolous Hoogstraal, 1953(C); description of all stages; biology; from rodent, hedgehog, and Varanus lizard burrows in Egyptian deserts. Absence of spirochetes (Davis and Hoogstraal 1954). 0. capensis Neumann, 1901; described from penguin-inhabited islands oT Cape Province. Found on Cargados Carajos Island in Indian Ocean (Neumann 1907E). Challenger Expedition specimens from St. Paul's Rocks (Nuttall et al 1908). Present on islands off Southwest Africa (Tromsdorff 1914). Biology of hosts and description of habitat, St. Croix Island off Cape Province (Hewitt 1920). Other South African records (Bedford 1934). 5 1s = Records from western Australia (Taylor and Murray 1946). Present on marine birds, Guam (Kohls 1953). Specimen from leg of soldier on island in Lake Nyasa (Hoogstraal 19510 ). O. coniceps (Canestrini, 1890); described from Venice, Italy. — pecimens, from near Aral Sea, in St. Petersburg Museum (Birula 1895); these quoted by Yakimov and Kohl_-Yakimov (1911) and Yakimov (1917,1922). Present in France (Guitel 1918, Theodor 1932, Roman and Nalin 1948). As 0. talaje from Fezzan, Morocco; introduced with pigeons; Severe se quelae in human victims (Martial and Senevet 1921). From bats and pigeons, life cycle, Tunis (Colas-Belcour 1929). Description of all stages, Palestine (Theodor 1932). Bit ology, Palestine (Bodenheimer 1934). Present in Spain (Gil Collado 1947,1948A,B). Present in Morocco (Blanc and Maurice 1950). Present in Nablus area of Jordan; parasite of chickens and persons; infected with spirochetes (Badu. dieri 1954,1955). Transmits fowl spirochetes (Brumpt's Precis"), Life cycle (Davis and Mavros 1956). 0. delano#i delanoei Roubaud and ColasBelcour, 1931; described a from porcupine burrow, Morocco. Biology (Roubaud and Colas. Belcour 1936). Life cycle and larval feeding (ColasBelcour 1941). Non-transmission of spirochetes (Colas—Belcour and Vervent 1949). Present in Egypt (Hoogstraal 19530). Ab sence of spirochetes (Davis and Hoogstraal 1954). Life cycle (Davis and Mavros 1956A). Biological observations and dis— tribution in Egypt; descriptions of immature stages (Hoog- straal 19555). O. delanoéi acinus Whittick, 1938; described from cave in British 7 Somaliland. Haemoglobin (Wigglesworth 1943). Biology (Ro- binson 1946). Coxal organs (glands) (Lees 1946B). Trans piration from cuticle (Lees 1948). Egg waxing organ (Lees and Beament 1948). Weight of tick and of its cuticle, fed and unfed (Lees 1952). Larval and nymphal measurements, and the increase in size following each molt and male meas— urements have been noted by Campana-Rouget (1954). - 115 . O. erraticus Lucas, 1849; described from Algeria and now known in Iran, Turkey, and throughout mich of the European and African Mediterranean area as well as in French West Africa, Kenya, and Uganda. This tick chiefly inhabits rodent burrows, some. times lairs and dens of other animals, and pigsties. It also may parasitize man, reptiles, toads, and birds. The very extensive literature on 0. erraticus will be reviewed in Volume II of this work. O. foleyi Parrot, 1928; single female described from the Algerian a ara. Description repeated by Foley (1929). A synonym of O. foleyi is 0. franchinii Tonelli-Rondelli, 1930(B) from Libya; . Roubaud and Colas-Belcour (1931). As 0. lahorensis, Q. lahorensis group, or O. franchinii from Libya by Franchini T1927, 1928B, 1932A,,B,1933A,D,1054A,1955A,1937) and Franchini and Taddia (1930); in these reports, the general remarks and those concerning fever in man as a result of bites do not ap. pear to be based on sound evidence; the "biological differ— ences™ (1934A) are pointless. Morphology and generic dis. cussion (Warburton 1933); cf. remarks herein under Argas brumpti (page 88). Presence in Southeastern Egypt (Hoogstraal and Retook 1956). Life cycle (Davis and Mavros 1954). O. graingeri Heisch and Guggisberg, 1953(A); all stages described from coral cave near sea, Mombasa. Life history (Heisch and Harvey 1953). Infected with spirochetes (Heisch 1953). Parasitizing bats (Garnham and Heisch 1953). Parasitizing porcupines and man (Heisch 1954A). Note: The actual date of publication of this species is 8 January 1953 although the volume number is that of 1952. 0. normandi Larrousse, 1923; all stages described, life cycle, from rodent burrows in Tunisia. Morphologic characters and biology (Colas-Belcour 1928). Egg laying and hatching (Colas_Belcour 1929A). Spirochete studies by Nicolle, Anderson, and Colas— Belcour (1927A,B,1928A,B,C ,D,1930). VO: joe nnonus Neumann, 1901; reported from Southwest Africa. Cos under 0. savignyi by Theiler and Hoogstraal 1955). ¥ leone O. peri Bedford and Hewitt, 1925; scanty descriptions and ot illustrations of male, female, and nymph from South Africa. Cliff swallow as host (Bedford 1929A,1932A). Failure to transmit Aegyptianella pullorum (Bedford and Coles 1933). All stages redescribed and reillustrated (Bedford 1934). O. salahi Hoogstraal, 1953(B); a parasite of fruit bats in the is Nile Valley and Wadi Natroun (Western Desert) of Egypt; also known from Palestine; all stages described; life history. Absence of spirochetes (Davis and Hoogstraal 1954). 0. tholozani tholozani Laboulbene and Megnin, 1882(A); first des cribed from Iran. An important Asiatic vector of spirochetes of relapsing fever; the tholozani group consists of several subspecies and related species; reviewed by Desportes and Campana (1946). Rare in western Egypt and eastern Libya (Coghill, Lawrence, and Ballentine 1947; Hoogstraal 1953C) but accused of transmitting spirochetes causing disease in troopse Now known from several restricted, but large, spirochete-infected populations in Egypt (Davis and Hoog- straal 1956) and from Jordan (Babudieri 1954,1955). 0. zumpti Heisch and Guggisberg, 1953(B); female and nymph des. ceribed from burrow of rodent (Rhabdomys ilio) in Cape Province, South Africa. Onderstepoort collection material recently sent by Dr. Theiler for identification includes males, females, and nymphs from the nests of Aethomys and ?Tatera in Cape Province. £0. lahorensis Neumann, 1908, an Asiatic-Near Eastern species, ~ Said by Franchini (1929B,1932B,1935) and Garibaldi (1935) to occur in Libya; most probably does not extend its range into North Africa, 7 O. sp. nov.; an undescribed species closely related to 0. ot foleyi has recently been found in porcupine burrows near Pretoria in the Union of South Africa (Theiler, corres. pondence). = 117 — KEY TO SUDAN SPECIES OF ORNITHODOROS MALES AND FEMALES With two pairs of eyes in lateral fold. (Northern and Central SUMAN es venice cie maine 6 cicisisinanisie si5-Os SAVIGNYI Figures 6 to le Without eyese (Southern SUAGH) cje sencciccin se sesaininiciaisieOs MOUBATA Figures 39 to 62 = 118 - . LEWS dorsal and ventral v Q> NITHODOROS MOUBATA > Figures 39 and 40 ed Laboratory rear OR PLATE XIV = DLO KNOWN APPROXIMATE DISTRIBUTION OF QO. MOUBATA RECORDS TO 195! (After Leeson 1952) 20 Present in town or small area Present in larger orea Recorded absence in small area Recorded absence In large area SUBSEQUENT RECORDS + Present in smoll area (HH) Absent in large area (HH) x Tick-borne relapsing fever present (Ordman 1955) Figure 41 KNOWN DISTRIBUTION OF ORNITHODOROS MOUBATA PLATE XV 5 IAG ORNITHODOROS (ORNITHODOROS) MOUBATA (Murray, 1877). (Figures 39 to 62) THE EYELESS TAMPAN* JES oti Sheet EQUATORIA PROVINCE RECORDS 2s Val: Liria African hut Nov Sy Wands Malice: 1 Nov Without locality data (King 1911,1926. Maurice 1932. Kirk 1939). Maurice (1932) stated that in connection with the 1925 relapsing fever outbreak, 0. moubata had been found in "four Equatoria Province rest houses north of Minule". The only eyeless tampans in Sudan Government collections are a few specimens from Wani Mika, collected by J. Dervish, 3 Novem. ber 1925. The three specimens collected from a hut at Liria in Novem ber of 1949 are the only ones that I have been able to find in the Sudan. A search of this and other Liria huts during January of 1952 failed to reveal additional material. During the same trip, preserved specimens were extensively exhibited to residents of many villages on the Juba.Torit-Tarangore track, and at Katire, Gilo, Yei, and Kajo Kaji. Although some persons who saw them professed to know the whereabouts of similar "dood", no tampans could be found. DISTRIBUTION IN THE SUDAN Bahr El Ghazal: From warthog burrows on Guar-Galual road near Guar, Gogrial Subdistrict, Tonj District, April, 1953, col lected by E. T. M. Reid and P. Blasdale. Three burrows examined yielded one nymph in the first, two males and nine nymphs in the second, and two males and thirteen nymphs in the third. Mr. Thomas W. Chorley and the collectors are to be congratulated for *In South Africa, called "The Hut Tampan" (Theiler 1952A,B). = WA these fine records of 0. moubata. Guar is situated at approximately 8°50!N. and 28°30'E, The specimens were collected during the dry season; but for several months of the year this "Nile sponge" area is a vast flooded plain. It would be interesting to know the ac. tivity of the ticks at that time. No evidence of 0. moubata has been discovered from buildings in Bahr El Ghazal Province, but search in these places has not been extensive enough to draw conclusions from this negative data. DISTRIBUT ION O. moubata is widely distributed locally throughout East and northern South Africa, and extends into the drier part of Central Africa. A few specimens indicate its presence in West Africa. In South Africa and elsewhere, human activities have carried the eyeless tampan far from its former habitats. Assuming, for lack of evidence to the contrary, that burrow-inhabiting populations and hut-inhabiting populations are identical species, it would appear that the distributional picture of the former populations, when it becomes better known, will be true indication of the prim itive range of 0. moubata. Early collecting records were summarized by Neumann (1901) and by Merriman (1911). Some early records are under 0. savignyi caecus Neumann, 1901, a synonym of 0. moubata, and for a time 0. moubata and O. savignyi were confused by Neumann and other writers. More recently, Leeson (1952) published a distribution map and ieee additional notes that have been brought up to date herein Figure 41). The mapped distribution of tick-borne, human relapsing fever in the world (American Geographical Society 1955) contains numer- ous errors in the area devoted to the Ethiopian Faunal Region, where 0. moubata is the only known vector, except rarely the human louse. A large section of the Sudan is shaded to indicate the presence of both tick-borne and louse-borne spirochetes — which would infer the widespread range of 0. moubata in the Sudan. This is contrary to the available data, presented above. The same ine dications for the Yemen are based on uncritical repetition of See ie earlier errors in literature, even though subsequent references disclaim earlier assertions — a frequent evil of uncritical fact-gathering from literature. Few outlines on the African map bear close relation to available facts. WEST AFRICA: Not recorded by Simpson (1912A,B) from the northern or southern parts of Nigeria. Absent from northern Ni. geria (McCullough 1925). No evidence to associate relapsing fever in northern Nigeria with 0. moubata (Caffrey 1926). Not known from Liberia (Bequaert 1930A). Absent from thirteen French West Africa villages on the Niger (Kerrest, Gambier, and Bourowm1922A,B). Recorded “south of Lake Chad" (Neumann 1901) but not found in Chad localities by Le Gac (1931); O. savignyi has been found in this general area (Alcock 1915) and cone me have misidentified his material. Apparent. ly absent in French West Africa (Gouzien 1923). Absent around Dakar (Mathis 1928A,B. Mathis and Durieux 1932A,B. Mathis, Durieux, and Advier 1933,1934). GOID COAST: Appears to be absent according to Selwyn-Clarke, Le Fanu, and Ingram (1923) and Ingram (1924); nevertheless a single nymph from Ashanti has been seen (Hoogstraal 1954C). SIERRA LEONE: Specimens seen by Hoogstraal (195/C). It is difficult to reconcile the GOID COAST and SIERRA LEONE records from British Museum (Natural History) collections with previous distributional concepts of 0. moubata, which have held that this species is absent from West Africa. Yet there is little reason to believe that the specimen labels are incorrect. A re— newed search in these areas is indicated; otherwise one hesitates to ae conclusions from these unexpected records (Hoogstraal 19540 ). CENTRAL AFRICA: FRENCH EQUATORIAL AFRICA (Blanchard 1913, 1914. Le Boeuf and Gambier 1918A,B. Rodhain 1919A. Guillet 1924. Blanchard and Laigret 1924. de Buen 1926. Rousselot 1951,1953B). BELGIAN CONGO and RUANDA_URUNDI (Livingston 1874, vol. 2, pp. 33 and 115. Neumann 1901,1911. Ross and Milne 1904. Dutton and Todd 1905A,B. Newstead 1905A,B,C,1906A,B. Massey 1908. Rodhain, = 123) = Pons, van den Branden, and Bequaert 1913. Roubaud and Van Saceghem 1916. Van Hoof 1917,1924. De Ruddere 1917. Rodhain 1919A,B,1920, 1922A,B,C. Todd 1919. Bequaert 1919,1930A,B,1931. van den Branden and Van Hoof 1922. Ghesquiere 1922. Van Saceghem 1923. van den Branden 1924. Olivier 1924. Seydel 1925. Schwetz 1927A,B,1932, 1933A,1942,1943. Van Hoof and Duke 1928. Flamand 1928. Schouteden 1928. Dubois 1931,1949A,B. Trolli 1931. Seraglia 1932. Pierquin 1950. Jadin and Giroud 1950,1951. Giroud and Jadin 1950,1954,1955. Jadin 1951A,B. Himpe and Pierquin 1951. Pierquin and Niemegeers 1953. Jadin and Panier 1953. Davis and Burgdorfer 1954. Theiler and Robinson 1954. Burgdorfer and Davis 1954. Babudieri 1955*). {NOTE: Rageau (1953B) suggests the strong possibility of the tampan's presence in Cameroons. He states, on epidemiological grounds, that relapsing fever there is presumed to be tick-borne and that the most likely areas in which the tick may occur have not been well studied. EAST AFRICA: SUDAN (Balfour 1906; tampan then not yet seen in the Sudan. King 1911,1926. Maurice 1932. Kirk 1939. Hoog- straal 1954C. See DISEASE RELATIONS and introductory paragraphs of DISTRIBUTION section). ETHIOPIA (Brumpt 1908A. Bergsma 1928,1929. Giordano 1936. Absence of tampans at 2,200 meters altitude; Scaffidi 1937. Bruns 1937. Franchini 1937. Mennonna and Modugno 1937. Manson.Bahr 1941,1942. Bertazzi 1952). ERITREA (Franchini 1929D,E. Niro 1935. Stella 1938A,1939A,1940). / FRENCH SOMALILAND (No known records).7 BRITISH SOMALILAND (Brumpt 1901. Drake-Brockman 1913 ,1915A,B,1920, identity in part confused with 0. sevigny Donaldson 1926. Clark 1937, remarks questionable. Stella 1938, 1940. Cullinan 1946. Anderson 1947. Heisch 1950A. Heisch and Furlong 1954. Davis and Burgdorfer 1954). ITALIAN SCMALILAND (Brumpt 1901,1908A. Paoli 1916. Rodino 1922. Reitani and Parisi 1923, Franchini 1925,1929C,E,1937. Bartolucci 1933. Mattei 1933. Niro 1935. Massa 1936A. Moise 1938,1950. Stella 1938A 1939A,1940. Lipparoni 1951,1954. Falcone 1952. Giordano 1953). *Professor Babudieri (1955) states that he utilized the Itete strain procured from the Congo by Professor Geigy. This strain was col. lected by Geigy and Mooser (1955) in Tanganyika. eat eres KENYA (As 0. savignyi: Karsch 1878. Ross 1912. Neave 1912. Todd 1913. Anderson »B. Garnham 1926,1947. Mackie 1927. "Kenya 1928". Lewis 1931A,€,1939A. Hynd 1945. Quin and Perkins 1946. Jepson 1947. Absence of Ornithodoros ticks: Garnham, Davies, Heisch, and Timms 1947. Meisch and Grainger 1950. Heisch 1950A,B,1954C,E. Walton 1950A,1953. Teesdale 1952. Bell 1953. Heisch and Furlong 1954. Geigy and Mooser 1955). UGANDA (Christy 1903A,B,1904. Pocock 1903. Johnston 1903. Sambon 1903. Ross and Milne 1904. Cook 1904. Ross 1906,1912. Hirst 1909,1917. Bruce et al 1911. Ross 1912. Neave 1912. Todd 1913. Neumann 1922. Mettam 1932. Hargraves 1935. Hopkins and Chorley 1940. Chorley 1943). TANGANYIKA (Neumann 1901. Sambon 1903. Christy 1904. Ross and Milne 1904. Dutton and Todd 1905A,B. Koch 1905,1906. Werner 1906. Mollers 1907. Todd 1913. Morstatt 1913,1914. Manson and Thornton 1919. (?Lester 1928). Loveridge 1928. Bequaert 1930A. Knuth 1938. Hawking 1941. Schulze 1941. Knowles and Terry 1950. Phipps 1950. Geigy 1951. Geigy and Burgdorfer 1951. Walton 1953. Davis and Burgdorfer 1954. Mooser and Weyer 1954. Smith 1955. Geigy and Mooser 1955. Babudieri 1955: see footnote under Belgian Congo). SOUTHERN AFRICA: ANGOLA (Livingstone 1857, pp. 382.383. Murray Lov”. As 0. savi i: Neumann 1896. Neumann 1901. Pocock 1903. Calman 1905. antes 1905A,B,€,1906A,B,C,D,1907A,B. Dutton and Todd 1905h,B. Nuttall et al 1908. Gamble 1914. Rodhain 1919A. De Almeida and Rebelo 1928. Sousa Dias 1950. Santos Dias 1950. Theiler and Robinson 1954). MO4AMBIQUE (Livingstone 1857, pp. 383, 628,629. Dowson 1895. Pocock 1903. Johnston 1903. Howard 1908, 1911. Amaral Leal and Sant'Anna 1909. Absent from Alto Molocue: McFarlane 1916. Neumann 1922. Marques 1943,1944. Santos Dias 1952H,1953B,1954H,K). NORTHERN RHODESIA (Neave 1911,1912. Wallace 1913. Lloyd 1913, 1915. Holmes 1953. Hoogstraal 1954C). SOUTHERN RHODESIA (Jack 1921 ,1928,1931,1937,1938,1942. Leeson 1952). NYASALAND (01d 1909. Neave 1912. De Meza 1918A. Lamborn 1924,1927,1939. Wilson 1943, 1950B. Hardman 1951). Sl) = BECHUANALAND (Specimens from Ngamiland in Theiler collection). SOUTHWEST AFRICA (Neumann 1901. Trommsdorff 1914. Sigwart 1915. Mitscherlich 1941. Hoogstraal 195/C). UNION OF SOUTH AFRICA (Lounsbury 19000, in part confused with O. savignyi. Pocock 1903. D8nitz 1906,1907A,C,1910B. Greenway 1907. Howard 1907,1908. Cowdry 1925C, 1926A, 1927. Curson 1928. Bedford 1920,1926,1932B,1934,1936. peotes 1934. Bedford and Graf 1934,1939. Cluver 1939,1947. Ordman 1939,1941,1943,1944A, B,1955. De Meillon 1940. Mitscherlich 1941. Collen 1943. Polakow 1944. _R. du Toit 1942B,C,1947A,B. Thorp, De Meillon and Hardy 1948. Monnig 1949 statements refer largely to O. savignyi. Annecke and Quin 1952. Annecke 1952. Davis and Burgdorfer 74 s ISLAND GROUPS: MADAGASCAR (Lamoureux 1913A,B. Suldey 1916. Poisson 1927. poisson and Decary 1930. Buck 1935,1948A,C,1949. Le Gall 1943. Millot 1948. Neel, Payet, and Gonnet 1949. Decary 1950. Colas-Belcour, Neel, and Vervent 1952. Hoogstraal 1953E). Although 0. moubata has not been reported from the COMORES group, Neel, Payet, and Gonnet (1949) state that relapsing fever of the apparent type borne by 0. moubata exists at Dzaoudzi. Records from Zanzibar (Brumpt 1901,1908A) probably should be disregarded. They may refer to “East Africa" in general. Accord ing to Dr. S. D. Robertson, Pathologist, Medical Department, in personal correspondence, 0. moubata does not occur in Zanzibar. It has been stated to be absen ere (Aders 1913) and was not listed by Aders (1917B) in his report of insects injurious to man and stock in Zanzibar. Odd individuals of the tick may be im ported in dhows from Tanganyika but these do not appear to become established on the island (Leeson 1953). ZNORTH AFRICA AND ARABIA: Records from Libya are difficult to accept. Franchini (1932A,B,1933A,B,C ,D,1934B,1935A) listed Tripolitania as the source of specimens. He was probably dealing with 0. savignyi, but Garibaldi (1935) accepted these reports. Zavattar1 =1933, 1934) stated that 0. moubata is absent in Libya. Though rranchind (loc. cit.) and Gaspare (1933,1934) as— serted differently, their remarks are so confused as to negate their argument unless fresh specimens can be procured. One of NAMRU.3's well trained assistants has been unable to find 0. moubata in the Libyan localities from which it was reported. = 126"= Records from Egypt (Neumann 1896,1901,1911), without question erroneous, are probably based on mistaken identity of 0. savignyi, or possibly on mixed locality labels (Hoogstraal 1954A)« awani (1946) stated that although 0. moubata was supposed to be common in Egypt, he could not find Specimens in houses. Yakoub (1945) also noted its absence here. According to Petrie (1939), the eyeless tampan is widespread in the Yemen (Arabia) (copied by American Geographical Society 1954,1955). From experience in the Yemen (Hoogstraal 19520 and ms., Girolami 1952, Mount 1953) it is questionable whether it is present there at all, to say nothing of being widely distributed. 7 HOSTS Introduction Man is frequently attacked and is probably the chief host of O. moubata. Warthogs and a few other wild animals that inhabit large burrows, and domestic pigs appear to be the only other fair— ly common hosts of this tick. Incidentally, it should be noted that frequent textbook assertions that larvae feed are incorrect (see Life Cycle below). Most laboratory animals including chickens serve as experi- mental hosts. Different “strains” may have different laboratory feeding habits, “burrow-haunting” populations being more difficult to induce to feed in the laboratory than those from domestic habitations (Heisch 195/C). Human Hosts The major portion of the literature concerning the eyeless tampan refers to its parasitism of human beings. Indication of this may be found in the section on wind below; specialized features of this problem will be considered in the forthcoming volume on tick.borne diseases. Domestic Mammal Hosts From the prevalence of records of attacks on domestic pigs in South Africa (Bedford 1936), Nyasaland (Wilson 1943,1950B), = 127 eS Belgian Congo (Roubaud 1916, Roubaud and Van Saceghem 1916, Schwetz 1927A), Southern Rhodesia (Jack 1921,1931,1942), and Angola (Wellman 1906) ,1907A,B), it appears that this animal often is an important host. In Southern Rhodesia, 0. moubata sometimes increases prodi- ziously in pigsties (Jack 1921,1931,1922). In the Zambi area of Belgian Congo, 0. moubata was abundant in pigsties and in huts of pig keepers, but relapsing fever was absent, and the tick was un known in local huts where no pigs were kept (Roubaud and Van Saceghem 1916). In Angola, Wellman (190@,1907A) found "as many in pigsties as in any other situation, In Nyasaland, Wilson (1943) stated, O. moubata is suspected of causing mortality in pigs. Jadin (1951A) found specimens from pigsties in Ruanda-Urundi infected with the causative organisms of food poisoning, Salmonella enteritidis; these ticks were able to transmit the bacteria to experimental animals, by biting, over a year later. O. moubata has been said to cause much trouble in Southwest Africa by feeding on sheep in resting places and pastures (MBnnig 1949), but Theiler states (correspondence) that the ticks actually involved in this situation are quite likely 0. savignyi. Domestic animal corrals are frequently cited in review papers as important habitats of the eyeless tampan. I can find little con clusive substantiation for this assertion, except for domestic pigs. Wellman (1904),1907A) was possibly the first person from the field to state that 0. moubata bites all domestic animals, but he did not mention that he had made personal observations, Careful search of corrals and comparison of incidence in these and in human habita. tions should make an interesting and simple research project in infested areas. One would expect that if domestic animals are attacked, it is chiefly in circunstances in which they are housed more or less like human beings in the same area. (See also Ecology below). Domestic Fowl Hosts Domestic fowls in human habitations usually are considered to be important in maintaining the nymphal stage, but there is some controversy on this point (Knowles and Terry 1950, Phipps 1950). Rodhain (1919A) found avian blood in specimens from a vacant house in the Congo inhabited by chickens. Geigy and Mooser (1955) failed ies) = to find evidence of spirochetes in domestic fowls from tampan in fested dwellings in Tanganyika, a suggestion that these ticks do not feed on fowls or do not transmit these organisms to fowls, or else that Borrelia duttonii does not survive in fowls in nature. Along with A. persicus, Mitscherlich (1941) discussed the ravages of 0. moubata in chicken houses in the Union of South Africa and in Southwest Africa (= Deutsch Stidwest Afrika). It is not, however, clearly stated that this writer actually saw eyeless tampans in these situations. His remarks give the im pression of being based on the assumption that 0. moubata is an important parasite of domestic chickens. aataltvadaail (See also Ecology below). Wild Mammal Hosts See also "“Wild™ Habitats under Ecology below. A wild relative of the domestic pig, the warthog, Phacohoerus aethiopicus subspp., is a normal host of QO. moubata under condi. tions not influenced by man. An African boy in Northern Rhodesia has been observed emerging from a warthog burrow with about thirty nymphs biting him (Lloyd 1915). During a survey of the plains south of Lake Edward in the Belgian Congo, Schwetz (1933A) dis covered that O. moubata was abundant in warthog burrows but rare in native huts. Chorley (1943) found over forty specimens crawling on a warthog shot in Uganda. He stated (personal conversation) that all these specimens were nymphs. Heisch and Grainger (1950) found numerous specimens in widely scattered warthog burrows in Kenya and presented a theory on the relationship of wild and dow mestic populations, discussed below in the section on "Wild™ Habitats, under Ecology. A single specimen from a Northern Rho desian warthog and a large lot of nymphs from a Nyasaland warthog burrow have been reported from material in the Nuttall collection (Hoogstraal 1954C). Warthogs are also hosts in the Suden, as noted above and reported earlier (Hoogstraal 1954B). These mammals also neve been noted as hosts in Mozambique (Santos Dias 1952H, 1953B). = 20 = In certain areas of Tanganyika, infestation of warthors and other large mammals is well known in some quarters (Walton 1953). Walton described a warthog burrow in which 41 hungry later—stage nymphs and adults were found; stomach blood smears from these gave a positive reaction to pig antisera. QO. moubata was also discovered in three other warthog and porcupine burrows in foot— hills of the Usambara Mountains. Jiterally hundreds of nymphs and adults emerged from the floor and ceiling to attack Walton and a friend when they entered some of these burrows. Subsequent— ly, specimens were found in six other burrows and in two hollow baobab trees that were used from time to time as retreats by var. ious kinds of animals. Smaller burrows in the Usambara Mountains area, presumably belonging to the giant forest rat, Cricetonys Spe, were uninfested. More recently in Tanganyika, Geigy and Mooser (1955) examined 55 burrows of warthogs, originally dug by antbears (Orycteropus afer), and found eyeless tampans in eighteen of them. ‘More than 1,200 tick specimens were collected from these retreats and an additional one was taken on the body of a freshly shot warthos. They also found the burrows of other kinds of mammals infested in Kenya. In connection with Sudan specimens from warthog burrows (Hoog- straal 1954B) (see also DISTRIBUTION IN SUDAN above), it is of interest to note that these are from the "Nile sponge” region that becomes a vast lake during the rains. Just what the ticks do during these floods should be worthy of investigation. Walton's (1953) records for porcupine (Hystrix sp.) burrows are noted above. Heisch (1954E) noted nymphs adults in por- cupine burrows in Kenya and found that they had fed on porcupine blood. Geigy and Mooser (1955), also working in Kenya, did not find ticks in a porcupine burrow that they examined but a nearby hyena shelter was heavily infested. In South Africa (Theiler, unpublished), specimens of 0. moubata have recently been taken from burrows of aardvarks or entbears, Orycteropus afer, near Stockpoort in the Potgietersrust area. Search for ticks in the retreats of these large, almost hairless animals will undoubtedly provide further interesting data. As noted elsewhere, other workers have found eyeless tampans in bur. rows originally dug by antbears but later occupied by warthogs. = WS{0) = Loveridge (1928) ambiguously associated GC. moubata with gi- raffes in Tanganyika, and Santos Dias (1952H,1953B,1954K) men tioned small nunbers of nymphal and adult specimens from lion, Lichtenstein's hartebeest, waterbuck, and scaley anteater. Fur ther data for these exceptional records are desirable. Heisch (1950A) obtained negative results when he attempted to induce 0. moubata in Kenya to bite house rats, Rattus rattus, placed in huts for experimental purposes. Wild rodents fron tickinfested Tanganyika dwellings gave no evidence of spirochetes when tested in the laboratory (Geigy and Mooser 1955). van den Branden and Van Hoof (1922) fed laboratory specimens on the fruit bat, Eidolon helvun. No other wild mammals have been reported actually to have been observed as hosts of 0. moubata in nature. The fact that the burrow-inhabiting warthog and the domestic pig each serve as a host of this tick is of special interest. NHeisch and Grainger (1950) have concluded that before 0. moubata became “domesticated it inhabited large burrows of wild aninals. FRoubaud (1916) conjectured that sone of the several external parasites of warthog and nan alike may be attracted to these hosts because of their hairless skin. This interesting theory is prob ably not now tenable for 0. moubata in the light of present know. ledge. In review, it appears that large burrows of wild aninels, among which those of the warthog are the most common, are the favorite and quite possibly the original habitat of O. moubate. It should be borne in mind, however, that those populations of this tick inhabiting wild animal burrows may possibly represent a different physiological or biological race, or a distinct sub species. It would be of value to determine the domesticability of “wild populations. Recently, Heisch (1954C) has noted that ticks from burrows are more difficult to feed on laboratory animals than are those from domestic habitations. Geigy and Mooser (1955) observe that bush ticks are more blue gray in color, move more quickly, attach to the host and suck blood more quickly, and are hardier in cap. tivity than specimens from domestic populations of 0. moubata. ee Contrary to Heisch's experience, they state that wild specimens “adapt themselves to feeding on mice and guineapigs easier than house ticks*, Wild Reptile Hosts Bedford (1934) listed several collections from South African tortoises. Theiler (unpublished) has records of nymphs and adults from four species of South African tortoises, Testudo oculifera, T. verreauxii, T. sch§nlandi, and Homopus femoralis from Kimberley and Wodehouse Districts and fron Namaqualand. Theiler considers tortoises to be exceptional hosts. Rodhain (1920,1922B,C) found that blood of lizards, geckos, and snakes is easily digested by 0. moubata. Although nymphs that had fed on snakes died in larger nunbers than those that had fed on mammals, survivors reached the same size as mammal-fed individ— uals. Chameleon blood is initially very toxic, and digestion is slow and difficult. Though many ticks die after feeding on cha meleons, a few do become adapted to it. Individuals that had fed exclusively on chameleons for sixteen months subsequently fed on mice when allowed to do so. Van Hoof (1924) reported similar findings. As already stated, tortoises sometimes are infested in South Africa, but no other collections from cold—blooded verte. brates in nature have been reported. BIOLOGY Life Cycle Life history details have been studied and reported by Dutton and Todd (1905A), Newstead (1905A,B,C,1906A,B) and Wellman (1906, D,1907A). These were reviewed by Nuttall et al (1908). Subsequent observations were reported by Cunliffe (1921), Jobling (1925), and Pierquin and Niemegeers (1953)*. Other contributions on special ized phases are noted below. Some discrepancies in observations exist, but the broad outlines of the life cycle are well established. Critical and restrictive factors are poorly known and no observa- tions on the life cycle under natural conditions have been under taken. The natural history of 0. moubata is gradually being elu. *The dates of publication of these reports will not be repeated in the life cycle section. = 132 = cidated, but each new observation suggests how many other details are yet to be known. A summary of the life cycle is as follows: Copulation is effected by transfer of a male spermatophore to the female, after which the female indulges in a rapid blood meal and subsequently deposits a small batch of unusually large eggs in or on the soil. After the larva emerges it remains nonmotile and nonfeeding till the nymphal stage some hours or days Later. The active nymph, after a short rest, feeds on an available host for about half an hour, then retreats to the soil or a crevice to digest its meal. Subsequently, the nymph molts, usually four or five times, with a similar pattern of resting, feeding, and resting between each ecdysis. Sexually mature adults emerge from the last molt and normally mate shortly afterwards. The female feeds two days later and several days afterwards deposits a batch of eggs. Adult hid ing and feeding habits are like those of nymphs. Seven feedings and egg batches appear to be maximum in one female's lifetime. A minimum of about two and a half months is necessary to complete the life cycle, which normally is probably considerably more ex. tended than this. Apparently these ticks do not voluntarily wander far in search of food and considerable numbers may develop in a single building or large animal burrow. The mating behavior of oO. moubata was described by Nuttall and Merriman (1911) but the account of mechanism of insemination has been augmented by Robinson (1942B). The development of the sperm has been described by Samson (1909). In the male the spermatids travel down the vas deferens either in a continuous stream or are aggregated in rounded pellets, each containing a few hundred male elements. As stated by Robinson and Davidson (1914) (for Argas persicus), it is probable that the male accessory glands secrete the spermatophore case into which these elements pass. According to Robinson, the spermatophore is not chitinous. It completely dissolves in strong KOH solution at 150°C., and becomes red in Millon's reagent; therefore it is probably largely protein Ad - 133 - In order to mate, the male crawls beneath the female and clings to her so that the two ventral surfaces are in apposition. After dilation and stimulation of the female orifice by insertion and movement of male mouthparts, a spermatophore issuing from the male genital aperture is grasped by the male's mouthparts and transferred to the female genital aperture. /Coxal fluid is emitted by the male during the course of these activities ac. cording to Nuttall and Merriman, but Dr. G. E. Davis and Dr. W. Burgdorfer state (conversation) that they have not observed this. It is possible that coxal fluid may or may not be emitted at this time, due either to copiousness of supply or to degree of excite- ment. 7 The spermatophore is bulb shaped (Figure 42) as it issues. After the male applies it to the female aperture, contraction and evagination force out the long neck with the capsules (Figure 43) that are inserted into the aperture. Most of the spermatids are forced into the capsules but the bulb remains outside the female aperture and drops off sooner or later. As many as ten bulbs have been seen in situ. The neck dries and twists, making an effective seal at the capsule closure. After five days at 30°C., the now mature sperms escape into the uterus by rupture of the capsule wall. / For further details, see Robinson (1942B)./ As stated below, the initial fertilization usually occurs shortly following molting to the adult stage, and females first feed about two days afterwards. They may feed before mating, presumably chiefly when males are not readily available. How. ever, according to Jobling, the period of time between fertiliza. tion and feeding has no effect upon the period between feeding and oviposition. - 134- ‘Www CAPSULE CLOSURE ~~ CAPSULE SS LAINE CK =a e BULB 43 42 Figure 42. Invaginated, bulb.shaped spermatophore as it issues from male genital aperture. Figure 43. Evaginated spermatophore, with neck and capsules, after having been applied to female uterus. ORNITHODOROS MOUBATA SPERMATOPHORE {After Robinson (1942) 7 PLATE XVI = 13> Generally, oviposition only follows a blood meal. For various reasons the interval between feeding and egg laying varies from five to 25 days, although (in tubes in jars with slightly moist cotton, maintained at 29.5°C. to 30°C.) the average is six to nine days (Jobling, and others). / Specialists will be interested in Wigglesworth's (1943) brief account of the fate of haemoglobin in ovipositing females. As each egg is emitted from the female genital aperture it , normally comes in contact with a peculiar glandular organ, gene's organ, that lies dorsally at the base of the capitulum. Gene's organ, which is everted only during oviposition, envelops each egg and provides it with a waxy, waterproof coating. Should this organ fail to evert or if any eggs are missed, these shrivel and fail to hatch, even in a humid atmosphere. The waxy coating is soft and viscuous (melting point 50°C. to 54°C., in contrast to cuticular wax, which is hard and crystalline with a melting point of 65°C.). The critical temperature of 0. moubata eggs well covered by this wax is 45°C. Lees and Beament (1948) have made a detailed study of gene's organ and its secretion, temperature and water loss of eggs, morphology of the female genital tract, structure and chemistry of the egg shell, and permeability of the egg shell. Eggs are deposited in masses on the soil or in hollows bur- rowed out by the female. It has been stated that the masses are agglutinated. Actually, individual eggs have a somewhat adhesive coating. When a container in which they are kept is jostled they roll about like globules of mercury. This is true also for eggs of numerous other argasids that have been observed. After oviposition, the female “broods™ over the eggs for some days (Wellman), a phenomenon of unknown function common among argasids. Jobling observed that this “brooding” sometimes con. tinues till the nymphal molt, after which the female may walk about for a time with several nymphs clinging to her. Dutton and Todd recorded individual batches of ten to twenty eges, with the greatest total of several batches from a single female numbering 139 eggs. Millers (1907) observed a single batch of eighty eggs. Wellman mentioned a lifetime total of 88 eggs and Newstead reported a total of 94 eggs. Records obtained under opti. mum laboratory conditions have been higher than those secured by = 136— these early field and laboratory workers. Cunliffe observed a female that produced a lifetime total of 535 eggs of which over ninety percent were fertile. In Jobling's tests, one female deposited several batches totalling 1,217 eggs and eight other females laid totals of from almost 700 to over 1000 each. Dr. G. E. Davis and Dr. W. Burgdorfer report (conversation) that the largest number of eggs they have observed in a single oviposition has been 233 and 327, respectively. Most eggs are laid at night and sometimes more than one day is necessary before a full batch is deposited. Six or seven batches, gradually diminishing in numbers, ap. pear to be usual in one female's lifetime. The amount of the previous blood meal influences the number of eggs subsequently produced. Jobling noted that the fertility of later batches de creases. In a laboratory study of 0. moubata fertility, Robinson (1942C ) found that three egg batches may be laid after one mating but that egg fertility is considerably increased if mating occurs before each oviposition. Fertility decreases when the interval between mating and oviposition is extended. Oviposition occurs almost without exception only after a blood meal. Eggs show no alteration in fertility when maintained between 22°C. and 32°., but at 34°C. no larvae emerge. As already stated, Lees and Bea ment (1948) have stated that 45°C. is the critical temperature for normal eggs. Robinson recommenced a temperature of 30°C. and a relative humidity of 50 in the breeding chamber for safe and speedy production. He found that a female might deposit a few eggs without a blood meal and that large females produce more than do small ones. The range in number of eggs per female per batch in these experiments varied from fifty to 250, with an av. erage of 170. Many females died shortly after their first blood meal; others after depositing their first egg batch. According to Robinson, females lay over twice as many eggs when sand rather than a flat surface such as filter paper is pro~ vided for this purpose, but Dr. G. E. Davis and Dr. W. Burgdorfer report (conversation) that in their experience the opposite is true. = Ug = Figure 44, Ege just deposited Figure 49, Rupture of egg shell Figure 45, Embryo, fourth day Figure 50, Larva hatching Figure 46, Embryo, sixth-seventh day Figure 51, Larva with shell, ventral view Figure 47, Hatching egg, lateral view Figure 52, The same, dorsal view Figure 48, The same, alternate contraction Figure 53, Larva, without shell, dorsal view and expansion Figure 54, The same, ventral view ORNITHODOROS MOUBATA EGG AND LARVA [After Jobling (1925) 7 PLATE XVII = 136— The egg of 0. moubata is among the largest known from ticks. A newly laid egg (Figure 44) is slightly ovoid, glistening golden yellow, and measures approximately 0.9 x 0.8 m. Later it becomes reddish brown. Eggs from older females are light to dark brown in color. An irregular, faint, whitish, polygonal reticula tion and interrupted radiating streaks may be seen through the cuticle. The internal larva becomes discernable four days after the egg is deposited and occupies the whole egg by the sixth or seventh day (Figures 45 and 46). [An alkaline haematin product originating from haemoglobin in the maternal blood meal has been demonstrated in eggs (Wigglesworth 1943). 7 Eight days after the egg has been laid (temperature 30°C.), the larva emerges by alternate contractions of the anterior and posterior ends of the body (Figures 47 and 48) that rupture the shell (Figure 49) and expose the larval dorsal surface. The shell may be completely detached in this manner, but usually remains on the ventral surface covering the mouthparts and legs (Figures 50 to 54). / Jobling 7 When movements necessary for emergence are completed, the larva becomes Lescent till the nymphal molt. That larvae are nonmotile after farce and do not Feol has been conclusively established for over a century, though several recent textbooks on medical entomology report differently. All observers have noted the quiescent stage between hatching and molting, and have differed only in the time required for a larva to molt to a nymph. Davis (1947) found that this molt occurred only a few hours after emer- gence from the egg. Robinson (1942) and Jobling stated that larvae molt four days after emerging from the egg (minimum, three days; maximum, five days). The various early observers reported periods of from three to 23 days from hatching till the nymphal mlt. The sacculated gut of a newly hatched larva is filled with a reddish brown fluid (Wigglesworth 1943). The inference is that this is an alkaline haematin resulting from the ingestion of hae— moglobin by the mother tick. / Before molting, the larva pales in color; its legs and mouth. parts shrink. Its skin becomes detached from that of the internal = Jg8).= 55 Figure 55. Nymph emerging from larval skin. ORNITHODOROS MOUBATA LARVAL.NYMPHAL MOLT [After Jobling (1925) 7 PLATE XVIII nymph; surface grooves disappear and the contour becomes more con vex. The internal nymphal outline and limbs are now visible. The two fore pairs of legs move to cause pressure on the larval skin resulting in a transverse rupture from which the anterior part of the body and the anterior legs emerge (Figure 55). After all legs are free, the larval skin is abandoned. / Jobling 7 The nymphal stage, in contrast to the quiescent larval stage, is very active. Cunliffe observed four to eight nymphal instars - 140. before ddulthood. He noted that most males appear at the fifth molt, most females at the fifth or sixth molt. Jobling (1925) and Pierquin and Niemegeers, however, observed no more than five molts and found the majority of male emergence at the fourth mlt and the majority of female emergence at the fifth molt. Dr. G. E. Davis* unpublished records show that in his laboratory most females reach the adult stage at the fourth molt.* Discrepancies in findings among various careful observers of this subject sug— gest an interesting field for research. / See also the section on symbiotes and growth-promoting substances, page igen), The interval between successive nymphal molts depends on the time of the preceding blood meal not on the time of the last molt. This is agreed by all workers. First instar nymphs require a longer period before they are capable of feeding (three to twelve *It should be noted that while argasids have several nymphal in. stars, ixodids molt directly to adults from the nymphal stage. Ixodids remain on the host for several days in each stage and ac- commodate the huge volume of ingested blood by slow cuticular growth (whether this is true for all ixodids, as for instance males of several Madagascan haemaphysalids, should be investigated; cf. Hoogstraal 1953E). Argasids, on the other hand, feed much more rapidly and accommodate the volume of fluid ingested by stretching the skin. Lees (1952) believes that this feature necessitates the several nymphal instars of argasids. The rapid feeding of argasids on animals that are usually resting reduces the danger of their transportation under inclement conditions and to unfavorable envi. ronments; they normally remain in handy feeding range in the host's house, burrow, den, or lair. Lees cites the several blood meals that a female argasid may take to nourish several egg batches as an adaptation to maintain her fecundity. Female ixodids, which take only a single, extended meal as adults, oviposit only once over a period of several days. It is well known that ixodids de. posit huge numbers of eggs but argasid eggs are relatively few in number. This discrepancy, however, is overcome by the more favor. able environment for obtaining a host in which argasid larvae and nymphs usually find themselves. Survival of argasid populations does not depend on large numbers of eggs but it does in ixodids. = IA days, mean five days; at 30°C.) than do later instars that feed on an average of two days (minimum one day, maximum five days) after molting (Jobling). Dr. G. E. Davis reports (conversation) that nymphs kept at normal room temperature require eight days before molting to the second nymphal instar and longer for suc. cessive instars. Jobling noted that first instar nymphs feed on an average of 25 minutes (minimim thirteen and maximum 87 minutes). Second and third instars average about four minutes less (minimum eleven and maximum 54 minutes), while fourth instar feeding is the longest (average 26, minimum 17, maximum 53 minutes). Jobling believes that the longer final nymphal feeding may possibly be necessary due to the requirements for metamorphosis to the sexually mature adult stage, which demands more nourishment than simple nymphal instar-to-instar development. These figures are in essential agreement with those of other students of the life cycle, mentioned in preceding paragraphs. Shortly before feeding is completed, a clear fluid begins to emerge from the coxal organs of all nymphal stages (as it also does from both adult sexes during feedings) and continues to issue until after feeding is completed and the tick has left its nost. (See REMARKS below). Nymphs are more resistant to adverse temperature and humidity factors than egg and larval stages, during which there is a much higher mortality than among nymphs. This is also agreed by all workers. The ratio of males to females is practically equal (Jobling). Males emerge from the last nymphal molt with a strong sexual urge and may fertilize several females before feeding. The aver. age male feeding time is sixteen minutes (maximum 42, minimm nine). After feeding they are less active and less eager for females and bury themselves in soil. Three or four dayslater they again be. come active and seek females. / Jobling 7 Females can be fertilized immediately after molting and several males may engage a single female before she seeks a blood meal. A female feeds for an averaze of 35 minutes (minimum 21, maxima 92). 6 WAe = This feeding period is longer than those of nymphs and twice as long as that of the male. Females commence feeding about two days after molting. / Jobling 7 Frequently repeated remarks by workers of the 1905 to 1907 period that 0. moubata may molt after reaching adulthood un. questionably were based on erroneous identification of advanced nymphal stages as adults. The minimum time necessary for 0. moubata to complete its life cycle is 62 days for males and 73 days for females, but in practice in the laboratory there seems to be some advantage to lengthening the periods of rest after molting and before feeding (Pierquin and Niemegeers), The life cycle can be enormously lengthened by delaying feeding and mating; and, for laboratory rearing, nymphs can be produced to meet any desired schedule, within certain limits, by selective timing. The longevity of O. moubata has excited much interest since it may be an important factor in allowing new populations to develop from a few imported specimens in areas where hosts are searce. Hirst (1917) maintained unfed specimens alive for four- teen months and Mayer (1918) kept others alive as long as five years. Cunliffe (1921) recorded female longevity averaging 715 days under ideal conditions of temperature and humidity with food available, and 441 days when food was unavailable. Nymphs (re ported as larvae) have been kept alive without food for over 710 days in the Nairobi medical laboratories ("Kenya 1928"). These figures are representative of numerous other records. The prac— tical importance of the long life of this species needs to be determined inasmuch as the fertility of long unfed females is much less than that of individuals that are permitted to feed at will. It has also been shown that female fertility decreases sharply five or six months following the nymphal-adult molt. No reports have been encountered that indicate a difference be— tween male and female life expectancy. Parthenogenesis of 0. moubata may have been observed by Cun. liffe, although he hesitated to be assured that the female had not been fertilized when unobserved. Parthenogenesis definitely has been established by Davis (1951), who reared 38 of 48 indi- - 143 - viduals hatched from unfertilized females. Oviposition in un fertilized females was mich delayed and the interval between hatching and molting of their progeny much prolonged. All prog- eny were females, but when these were mated with normal males both sexes were represented in the subsequent generation. According to Cunliffe, 0. moubata and 0. savignyi may copu. late but the resulting eggs are umertile. This is contradicted by recent, unpublished findings of Dr. G. E. Davis who writes (correspondence) as follows; “I have found that the interbreeding of these two species not only results in progeny but in fertile progeny when the products of the first interbreeding are allowed to interbreed among themselves™. The foregoing is a reasonably complete though brief summary of what is known about the life cycle of 0. moubata. Before leav— ing this subject, attention should be called to the additional temperature and humidity studies discussed under Environmental adaptability below for these factors exert considerable influence on the life cycle. Ecology Environment and Domestic Habitats The ecology and distribution of 0. moubata, as summarized in the paragraphs below, has always been considered in the light of domestic populations. The significance of the increasingly more numerous reports of the eyeless tampan in large animal burrows from the Sudan to South Africa awaits to be determined. Should it eventually be found that these two populations are a single biological entity that has happened by chance to occupy one or the other habitat, the conclusions of early workers, who believed that man has been wholly responsible for carrying this tick out— ward from its primitive range in the East African lowlands, will have to be modified. The arid environment preferred by domestic populations of 0. moubata restricts their presence to dry, permanent huts and structures where people gather. In its probable original area, the Somali Arid District and possibly the East African Lowland = 1A District, this tampan appears to be more uniformly distributed than elsewhere. These details, however, await confirmation. Outward from the Somali District the eyeless tampan normally inhabits dry structures in savannah areas, especially those with sandy or sandy clay soils with light woods. Riparian forests through grasslands, dense forests, and areas of heavy rainfall are usually free of the tick, although exceptional human culture patterns sometimes allow important foci to develop in dry habi. tats in these situations. Such details have been described most vividly by workers in the Belgian Congo (Bequaert 1919,1930A; Rodhain 1919A,B,1922A,€; Ghesquiere 1922; Schwetz 1932,1933A,1942, 1943; and others). O. moubata appears to have spread gradually outward from somewhat dry areas of East Africa along main paths of human trav- el. Old Arabic slave routes are considered to have been largely responsible for its initial distribution by man (Dutton and Todd 1905A; Bequaert 1919,1930A). Although especially common along important old and new travel arteries, the tampan is often mark edly absent a few miles distant. Exceptions do occur. For instance, Koch (1905) reported 0. moubata from the Rubafu Moun. tains and elsewhere in villages away from trade routes in Tanga. nyika. More and more exceptions should occur as travel becomes easier and quicker, tribal customs disintegrate, and labor de— mands call numerous individuals, with possibly tick-infested personal effects, far from their usual range of activities. 0. moubata is said to be frequently concealed in sleeping mats, spare clothing, or baskets and thus may be transferred easily from one area to another. South African authorities blame the tampan's increasing spread in the Union on migratory laborers from Nyasaland and Portugese territories. In the Belgian Congo it has been found in potato baskets sent to distant markets (Ghesquiere 1922) and is frequently introduced in goods sent from the lowlands to villages at high elevations (Schouteden 1928). This tampan is common in fish baskets of vendors bicycling from Lake Nyasa and Lake Shirwa to villages in other parts of Nyasa. land (Hardman 1951). Christy (1903A,B) collected specimens in salt bags being transported between Lake Albert and Tete. - 145- Where soil consistency permits, the eyeless tampan usually burrows to a depth of approximately an inch; but in soil cracks it burrows deeper. On the ground surface it may rest under any object that offers shelter. If soil is too wet or too hard for burrowing the tick is induced to crawl up walls and seek con. cealment behind hanging objects, in cracks or in ceilings. The tampan's presence is often indicated by spots or streaks from its excretory products left on walls. In Kenya, 0. moubata ranges from sea level to an elevation of 8000 feet (Lewis 19398) or of 9000 (Heisch 1950A). In Ethiopia, Manson.Bahr (1941) stated it is absent above 6000 feet elevation and Scaffidi (1937) reported that it is not present above 7150 feet. This tampan survives in the Transvaal Highveld in spite of “bitterly cold winters” there (De Meillon 1940, Ordman 1941). Theiler reports (correspondence) that winter day.temperatures in the Transvaal highlands are “high enough” and that residents bring fires into their huts at night. In certain Congo areas, Flamand (1928) found 0. moubata thriving at about 10,000 feet elevation. Schouteden (1923) replied that these populations are the result of repeated introductions in coods from the lowlands. While tampans survive at these altitudes, they do not reproduce there. Incidental to a disease transmission study, Van Oye (1943) reported that O. moubata dies in less than 24 hours at tempera. tures of 0°C. to 50. (419F.). However, Burgdorfer reports (con versation) that he has maintained tampans at 3°C. for at least ten days without death of the specimens. Feng and Chung (1938) maintained these ticks alive for months at 59°C. to S0C. It is obvious that the critical temperature range for the survival of 0. moubata requires further study. The absence of 0. moubata from certain volcanic areas in the Congo was thought by Van Saceghem (1923) to be owing to an un favorable chemical action produced by contact of lava with oxygen of the air. Dr. J. Bequaert reports (conversation) that these areas are all at high elevation and that he believes altitude to be the important limiting factor in these volcanic areas. = lio Wallace (1913) noted that in Northern Rhodesia 0. moubata abounded in hilly country and on the Mpika plateau, but was ab sent in the hot Luangwa Valley. The combinations of temperature and relative humidity factors that restrict 0. moubata in nature are still poorly known. te is was Dutton and Todd (1905A,B) and Bequaert (1919) mentioned an exception to the general rule that 0. moubata does not occur in deeply forested, humid areas. The explanation was that infesta. tions in the Upper Ituri Forest are in wooded areas where arab isized Negro inhabitants have cut over extensive forests and constructed villages of dry, permanent buildings. In these, ticks brought along the Arab trade route have been able to survive in spite of inclement conditions outside. Another notable exception to the usual finding that 0. moubata is absent from high rainfall areas is cited by Walton (1950A). In Meru District, Kenya, even under unfavorable high rainfall and humidity conditions, large populations of this tick survive in some huts. The predisposing factor is that the local tribes sleep on dry, raised mud beds. Hosts are readily accessible to ticks and fires near beds keep an area of ground dry enough to meet the tampan's requirements. Agricultural implements in these huts provide additional shelters behind and under which the ticks also hide. Contrary to usual advice to remove domestic animals from human habitations, Walton believes that under Meru District conditions animals provide enough extra humidity and pound floors hard enough to reduce tick populations. Under more usual conditions, however, this suggestion would probably not be an effective one. In some Meru District huts, where sticky soil has a humidity of about 9%, it is difficult for ticks to burrow and few specimens are found. Whether this investigator searched for ticks climbing walls or pillars in these huts is not stated. At Kisum, Heisch (19504) found that the size of tampan popu lations in huts is uninfluenced by seasonal variation in rainfall. Knowles and Terry (1950) collected hundreds of nymphal tampans on fowls kept in human habitations in Tanganyika, but Phipps (1950) asserted that chickens are seldom infested there. Careful research into the highly practical problem of relationship of chickens to = WAT tampan infestation is indicated. Rodhain (1919A) reported finding avian blood in specimens taken from an empty outhouse inhabited by chickens. It has been suggested that periodic forays by chickens into infested huts may partially reduce the tick population in these places. Aside from usual indigenous dwellings, the eyeless tampan is frequently encountered where people congregate. In Uganda, rest camps often have been burned because of heavy infestation (Bruce et al 1911); jails and semipermanent buildings used by itinerant Hfricans are frequently infested (Hopkins and Chorley 1940). In Kenya, 0. moubata is “alarmingly abundant" in labor camps (Jepson 1947) and Military barracks are specially constructed to resist infestation (Hynd 1945). The tampan is a coffeehouse inhabitant in British Somaliland (Anderson 1947). In a Somaliland focus of relapsing fever, all patients were found to be members of a polit- ical party the headquarters building of which was infested with OQ. moubata and had escaped insecticiding when other structures were dusted (Lipparoni 1951). In South Africa, it is an important pest in “lesser mine™ labor camps but in larger mines, such as those at Johannesburg where sanitary measures are practiced, the tick is absent (Ordman 1941,1943). In contrast, the closely-related eyed tampan, 0. savignyi, usually lives away from habitations, under trees, in village squares, near wells, in stockades, or in shaded spots along trails where men and animals rest. There are but few reported observations of 0. moubata living under outdoor conditions approaching those favored by 0. savignyi. In 1916, Belgian colonial troups operating in Urundi, while ten ing under a row of mango trees that had bordered buildings des— troyed some six years earlier, recovered several specimens from the soil around the roots of these trees. The assumption was that these ticks had survived since the destruction of the near. by buildings some years earlier (Rodhain 19198). Ordman (1941) listed two cases of the eyeless tampan in South Africa living “in and under trees", but further corclusive evidence is not presented. - 148. In various editions of Brumpt's "Precis,™ 0. moubata is con sidered as an outdoor as well as an indoor species. There is, however, no published evidence to support the supposition that this species normally lives away from human structures, except in large animal burrows and in pigsties. Rare exceptions, such as ticks remaining in the area after a building is destroyed, or dropping from a bedding roll during transit, must be expected. Further search may, of course, show that the tampan has a broader range of habitats than present evidence indicates. “Wild™ Habitats A gradually increasing body of information indicates the not uncommon occurrence of 0. moubata in large animal burrows through. out tropical and southern Africa (see HOSTS above). The relation. ship of these populations to those of human habitations awaits determination. The environment of infested burrows has been only briefly described and it is not known whether wild populations have the same temperature and humidity requirements as domestic populations. In Tanganyika burrows, Walton (1953) observed tampans clinging to the roof close to the entrance as though waiting for some animal to squeeze past. Ticks were found among the hair of the back of warthogs shot in the early morning. In the burrows, temperature was 75°F. and relative humidity of the soil 77%. Other infested Tanga. nyika burrows examined by Geigy and Mooser (1955) with thermohygro- meters showed that the microclimate of these holes corresponded closely to that observed by them in infested native huts (details not stated). Discovery of numerous specimens in large burrows in several widely scattered parts of Kenya has led Heisch and Grainger (1950) to speculate on the relationships between wild and domestic popu. lations of eyeless tampans. The ticks were obviously breeding in these burrows that originally had been dug by antbears and later were inhabited by porcupines or warthogs. Other specimens were found in large burrows on a long-isolated Lake Naivasha island seldom visited by man. Heisch and Grainger conjecture that large burrows were the original or primitive home of the eyeless tampan and that it later became adapted to human habitations. The several reports of 0. moubata from burrow-inhabiting warthogs, porcupines. =- 149 and antbears, already mentioned in the section on HOSTS, bolster this theory. Further indirect support is gained from the preva. lence of the warthog's relative, the domestic pig, as a host. The tampan of human habitations may have evolved from populations formerly parasitizing burrowing, wild pigs, and they may still retain some predilection for pigs. As already noted, it is also possible that “wild™ and “domestic™ populations represent separate biological or physiological or even unrecognized morphological entities. Predators and Enemies Chickens, rats, and mice are said to feed on the eyeless tam pan, and ants carry off eggs and nymphs. An Angolan Reduviid bug, Phonergates bicolor Stal. sucks the blood of both man and 0. mow bata / Wellman (1906B,D,1907B). Austen (1906,1907) reported on the nomenclature of this bug. The actual specimens involved may still be seen in British Museum (Natural History) collections 7. Ant lions (Neuroptera, Myrmelionidae) have been observed feeding on nymphs (Ghesquiere 1922). In the laboratory, larvae of clothes moths, Tineola biselliella, are said to feed on eggs and on living larvae of 0. moubata (Volimer 1931). What was once described as a fungus disease beginning as an opaque white spot at one edge of the body and spreading out to stupify and destroy the tick (Wellman 1906A,D,1907B) is now be. lieved by experienced workers to be a normal phenomenon of aging in engorged ticks. Christophers (1906) suggested that this "fun cus* is actually a white rectal secretion of aged ticks. Burg- dorfer (conversation) is of the opinion that this “white fungus" is nothing more than crystallized fluid in the malpighian tubules. Often this crystallization produces a complete, hard blockage. The lumen of such tubules fills with white crystals so that nor- mal activity can no longer occur and soon the tick dies. (See Internal Anatomy below). Numerous factors affecting the ecology of the eyeless tampan are discussed below. el Ole REMARKS Environmental Adaptability The xeric environment in which 0. moubata is capable of sur. vival is best explained by two physiological studies by Lees (1946A, 1947). In his research on water balance in ticks, Lees (1946A) found that among the species studied, 0. moubata shows the greatest ability in limiting evaporation from its own body. In this species, the critical temperature at which water loss increases through the superficial waxy epicuticular layer is also high (Lees 1947). This resistance to desiccation at temperatures within its biological range may be correlated broadly with the argasids' choice of dry, dusty ecological niches. Lees summarized his 1946A studies, in which Ixodes ricinus was the principal species for research and 0. moubata was one of eight other species used for comparative purposes, as follows: “The unfed tick gains water from humid air or from water in contact with the cuticle, and loses water by evaporation. Whilst attached to the host the tick is gaining water from the ingested blood and losing water in the excrement. The engorged tick usually lacks the ability to take up water from humid air. "The exchange of water takes place mainly through the cuticle. Regulation of the water balance is there— fore brought about by the activity of the epidermal cells. "The cuticle comprises two principal layers, the epicuticle and endocuticle. The epicuticle is overlaid by a lipoid possessing important waterproofing proper. ties. The pore canals, which traverse the endocuticle, are occupied by cytoplasm, and may in consequence play an important role in the active transfer of water through the cuticle; they do not penetrate the epicuticle. “Water loss from the unfed tick is not closely re lated to saturation deficiency, particularly at high a lophe humidities. This departure is due to a physiological cause, namely, to the ability to secrete water. The effects of this activity are such that a state of equilibrium is attained at a relative humidity of about 926; at lower relative humidities it takes up water. The retention of water at humidities below the point of equilibrium is due not only to the physical properties of the epicuticle but also to this secretory activity, for water loss increases when the tick is temporarily asphyxiated, poisoned with cyanide, or injured through excessive desicca. tion. Near the point of equilibrium the loss or gain of water over a wide range of temperature is determined by the relative humidity. "The uptake of water from humid air occurs when the tick is in a desiccated condition but ceases as the normal water content is restored. After previous exposure to saturated air the adapted tick at first loses water at relative humidities above the point of equilibrium, but later comes to retain water com pletely. “Both unfed and engorged ticks possess the abil. ity to prevent or to limit temporarily the entry of water in contact with the cuticle. "The engorging female, originally weighing about 2mg., ingests about 600 mg. of blood. About 300 mg. or two-thirds of the contained water are usually eliminated before the end of engorgement. Evapora. tion from the cuticle may account for a considerable fraction of this, for the temperature to which the attached tick is exposed (about 37°C.) is, in Ixodes ricinus, above that temperature at which a marke Increase in the permeability of the epicuticular lipoid takes place. "The nine species of ticks examined differ con. siderably in their powers of limiting evaporation. This may reflect specific differences in the nature we FLO os of the epicuticular lipoid. The order of their resistance is as follows: Ornithodoros moubata; Dermacentor ander— soni; D. reticulatus; Rhipicepnalus sanguineus ; Amblyomma cajennense and A. maculatum; Ixodes cane us I. hexagonus; e .) > e Le Ticinus. In dry air, water loss oug the oe : is ten to fifteen times more rapid in Ixodes ricinus than in Dermacentor andersoni. The more resistant spe- cies also take up water through the cuticle after desic— cation; indeed, the rate of uptake over a unit area of cuticle is approximately the same in all species of Ixodidae. Uptake thus appears to be limited by the ability of the epidermal cells to secrete water." As already stated, Lees has shown that 0. moubata is more re. sistant to desiccation than most ixodid ticks. Nymphs exposed to dry (G5 R.He) @ir at 25°C. survived for 35 days and lost only from one to three percent of their original weight daily. This survival period is strikingly longer than that of several ixodid tick spe- cies used in the experiments. After a period of desiccation (five days at @& R.H.), O. moubata regains most of its original body weight when placed in 95% R.H. for five days. Water is taken up through the spiracles, for no increase occurred when these open ings were blocked. loss of water occurs through the cuticle and spiracles (see Spiracular Morphology and Function below). In order to carry Leest work one step further, Browning (1954B) conducted a study on the exchangesof water between the atmosphere and O. moubata. Unfed nymphs were able to abstract water from moist air (95% RH.) and to restrict their rate of water loss in dry air. This ability was lost (a) in atmospheres containing 3% to 45% C02; (b) in atmospheres containing more than 9% N2; (e) immediately after the tick fed; and (a) gradually after the tick has been starved for some five months. It was shown that the action of high (30% to 1.5%) concentrations of C02 is mainly upon the activity of the epi- dermal cells, possibly mediated through the central nervous system. The concentration required to cause opening of the spiracles is only about five percent. These findings are of considerable interest in relation to Lees! (1947) basic work. By way of introduction to his 1947 study, Lees stated: = 153u— "In considering the mechanisms involved in the ex. change of water through the cuticle the assumption was made that, in addition to active secretion, the passage of water, and particularly its retention, is also im fluenced by the presence of lipoid material in the cuticle. Ticks show great diversity in their powers of resisting desiccation, anc this was thought to be accounted for by the specific nature of the waterproofing lipoid. Never theless, no direct evidence of such a component was ad. vanced in this paper (i.e., Lees 1946A). "Ramsay (1935B), and mre recently Wigglesworth (1945) and Beament (1945), have shown that the imper- meability of insects is entirely due to a thin, discrete layer of wax or oil in the outermost part of the epi. cuticle. Any agents such as abrasive dusts, wax sol. vents, or detergents, which interrupt the continuity of this layer, at the same time greatly increase transpira. tion. Water loss through the wax layer is also enor— mously increased if the temperature is raised above a certain critical value. ...... methods devised by Wiggles. worth for demonstrating the properties of the waterproofing layers in insects have been applied to a number of species of ticks. .eecce Observations on the structure and depo. sition of the epicuticle, and on the functions of the dermal glands (are provided). The outermost layer of the tick cuticle visible in ordinary sections has hither. to been referred to as the "tectostracum" (Ruser 1933) eseeee (but) the similarity of this layer with the insect epicuticle is so marked that the abandonment of this term seems fully justified." The results and conclusions of this work, Lees summarized as follows: “], Ticks owe their impermeability primarily to a superficial layer of wax in the epicuticle. After expo. sure to increasing temperatures, water loss increases abruptly at a certain critical temperature. The critical temperature varies widely in different species, in Ixodidae ranging from 32°C. (Ixodes ricinus) to 45°. / Hyalomma marginatum (= savignyi) 7; and In Argasidae Som OI 5 bys = (Ornithodoros moubata) to 75°C. (0. savignyi). Species having higher critical temperatures are more resistant to desiccation at temperatures within the biological range. A broad correlation is possible between these powers of resistance and the natural choice of habitat. Argasidae infest dry, dusty situations whereas Ixodidae occupy a much wider variety of ecological niches. m2. If the tick cuticle is rubbed with abrasive dust, evaporation is enormously increased. Living ticks partially restore their impermeability in mist air by secreting wax from the pore canals on to the surface of the damaged cuticle. "3, Unfed ticks are able to take up water rapidly through the wax layer when exposed to high humidities. Water uptake, which is dependent on the secretory activities of the epidermal cells, is completely in. hibited by the abrasion of only part of the total cuticle surface ~ a fact which suggests that the cells are func- tionally interconnected. Resistance to desiccation at low humidities is achieved by a dual mechanism: active secretion and the physical retention of water by the wax layer. "4. In Argasidae the epicuticle consists of four layers: the cuticulin, polyphenol, wax, and outer cement layers. Only the three inner layers are present in Ixo- didae. Since the wax layer is freely exposed in the latter group, chloroform and detergents have a marked action in increasing transpiration, particularly in those species with low critical temperatures. In Ar. gasidae the cement layer is very resistant to extrac- tion but is broken down by boiling chloroform. "5. The cuticulin, polyphenol, and wax layers are all secreted by the epidermal cells. The water— proofing layer, which is deposited on the completed polyphenol layer, is secreted by the molting tick relatively early in development and may be nearly complete by the time molting fluid is abundant. In 0. moubata the cement is poured out by the dermal - 155 - glands shortly after emergence. In Ixodidae the dermal glands undergo a complex cycle of growth and degeneration, but their products appear to add nothing of functional significance to the substance of the cuticle." lees! important contributions indicate why 0. moubata is capable of surviving in the dry niches in which domestic popu lations occur. However, we still lack data on the actual rel ative humidity of these niches in nature. We know only that the tampan can withstand these conditions in laboratory investiga. tions. And it should be stressed that we still know nothing about preferences and critical levels of temperature and humidity among burrow-haunting populations. The Bahr El Ghazal collections, from warthog burrows in the "Nile sponge area, especially ex- cites curiosity in this respect. Laboratory studies on the optimum temperature and humidity conditions under which 0. moubata survives have resulted in wide ly differing data and conclusions. The reports in question are those of Cunliffe (1921) and Brett (1939) together with those of Robinson (1942C ) and others already reviewed in the section on the life cycle of 0. moubata. Cunliffe found that a saturated atmosphere has no inhibitory influence on molting but is decidedly unfavorable for vitality (only one specimen passed the third nymphal stage under these conditions). Even under “medium conditions of humidity", mor- tality is high, but under "dry conditions”, 64 of the nymphs complete metamorphosis and the rate of development is increased. High temperature increases the number of eggs laid but decreases fertility, longevity, and time required for metamorphosis. Brett, on the other hand, found that (at 25°C.) higher rel. ative humidity (up to 804) was more favorable for survival of eggs, larvae, and first instar nymphs (the only stages and im stars tested) though a proportion of all eggs were able to dev— elop at any "low humidity normally met with in nature, He also found that the first nymphal instar is much more resistant to desiccation than larval and egg stages. The apparent inconsist— ency between Brett's findings and the known fact that domesticated populations of 0. moubata are chiefly inhabitants of drier areas = 1565 AT Gene> organ+s~< Oesophagus Central ganglion. cee iy Coxal organ — Accessory €—) gland Salivary Filter Waco gland chamber <~ 4 a x Oviduct — < Stomach and Or diverticula Oe Malpighian ~ ag Rectal ampulla 56 Figure 56 Q ORNITHODOROS MOUBATA INTERNAL ORGANS / After Burgdorfer (1951), with permission of the editor of ACTA TROPICA 7 PLATE XIX = 157 = Salivary gland Central ganglion Pherynx \ esophagus mK =i 0 0 OO ~< : OCP Se 006 | Spirochete 0 °cSHost blood cells 58 Figures 57 and 58, 9, Unengorged and Engorging Spirochetes of African tick-borne relapsing fever, Borrelia duttonii, are illustrated, as short wavy lines, in the positions they occupy in the tick's body. Note their escape routes from the tick's body and into the host's body while the tick is feeding. ORNITHODOROS MOUBATA DIAGRAMMATIC SAGITTAL VIEW {After Burgdorfer (1951), with permission of the editor of ACTA TROPICA_7 PLATE XX Sls) is explained on the basis of Williams! (1923,1924A,B) and Buxton's (1932,1933) exposition of the comparatively high humidity in sand, cracks of walls, and soil in areas that are otherwise dry. Brett's discussion and the comparison of his findings with those of Cun liffe and of other workers, especially those of Robinson (19420) discussed on p. 137, which corroborate those of Brett, should be studied for their practical importance by anyone concerned with O. moubata. Since only careful and thorough research in the field ‘as Well aS in the laboratory can conclusively settle the matter, a more complete discussion of this question is hardly in order here. Structure and Function Introduction No thorough studies of the internal anatomy and histology of ©. moubata have been undertaken, What has been done on certain aspects of these subjects is reviewed in the following paragraphs. On the whole, workers have been content to accept Christophers! (1906) careful though still somewhat general description of the internal anatomy of 0. savignyi as also applicable to 0. moubata. Recently, Burgdorfer (1951) = provided a short account of the internal anatomy of 0. moubata and some of his excellent illustra. tions are reproduced (Figures 56 to 58). However, 0. moubata deserves more specialized attention than it has thus far been accorded. These two species differ in habits, habitats, distri- bution, and receptivity to pathogenic organisms. It may be ex. pected, therefore, that under their leathery shells, which also differ, significant anatomical and physiological differences re— main to be demonstrated. Internal Anatomy The general features of the internal anatomy of these two species are similar and Christophers! (loc. cit.) description of a dissection of 0. savignyi, as presented below, applies equally well to 0. moubata Garth oon differences noted): “Over the whole dorsum lies a fine membranous ex- pansion of tracheae and trabeculae of the fat body. Lying in this, in the median line, is the delicate tubular heart. Posteriorly, at about the junction of - 159 - the middle with the posterior third of the body, this is considerably dilated. Stripping off the expansion, the main mass of the viscera, consisting largely of the large dark red blood sacs of the alimentary canal, are exposed. By carefully unravelling these, the ar. rangement of long diverticula, described later, can be made out. Lying upon the diverticula in the poste— rior portion of the body is the ovary, studded with developing ova. Upon either side of the ovary are the coiled oviducts, and in the middle line is the large conspicuous bilobed spermatheca (uterus). In almost every region of the body a portion of the thin coiled malpighian tubules will be found. Behind the sperma theca 1S an opaque white organ, having very thin sac- cular walls and filled with characteristic white secre- tion from the malpighian tubules. This is the rectum (rectal ampulla), which in ticks serves as an excretory bladder. = displacing the diverticula from the extreme anterior portion of the body a bilobed glandular organ, the cephalic gland (gene's organ) is displayed. Further back, the bulbous ane of the cheliceres with radiating muscular fibres are seen. Around them will be noticed the ringlike chitinous fold at the base of the rostrum. By displacing to one side the whole of the anterior and lateral diverticula, a member of further structures are apparent. Passing in from the stigmatic (spiracular ) openings is a leash of tracheal branches, of which the large anterior ventral trachea is the most conspicuous. Lying upon the origin of the first and second legs is the large racemose gland which functions as the salivar land in ticks. Lifting this gland by its posterior extremity, which lies on the anterior ventral trachea, and tracing it forward, the short salivary duct will be apparent entering the ringlike fold of chitin, already mentioned, immediately beneath the cheliceres. Lying partly under the salivary gland, and partly internal to this structure is (the large, saccular coxal organ) conspicuous from the number of tracheae which Anas it. “By careful examination, the delicate, colorless esophagus can be made out entering the lower surface of the large median blood sac of the alimentary canal, whilst lying behind the spermatheca is the fine hair— like termination of the sac in the rectum, To the rec- = 160 tum can be traced the attached end of the two extremely long malpighian tubules. To display the esophagus in its passage from the pumping organ to the alimentary sac it is necessary to tear away the dense mass of muscle from which it will be seen to emerge. By seizing the muscular mass boldly in the forceps, the unattached entosternum surrounded with muscle will come away, exposing the central lion, perforated by the eso- phagus. By seizing aie ends of the cheliceres they may be drawn from their sheaths. Lying beneath them is the horizontal entosclerite of the head. Beneath this, again, is a dense mass of muscle within which lies the chitinous pumping pharynx. “Tn the male, in the position of the ovary in the female, there is a delicate tube abundantly supplied with trachea. On either side this is continuous with a coiled duct much resembling the oviduct in the female. In the middle line, much in the position of the sperma— theca in the female, is a curious lobular organ, the white gland ......" (that) is probably concerned in the elaboration of spermatophores. Following this, Christophers (loc. cit.) presented a more complete account of each structure and a generalized description of the digestive process in QO. savignyi. This should be consulted by anyone interested in the ee anatomy and function of either species. Sections of Christophers! study dealing with the digestive system are abstracted below because of their re— lation to the ingestion, development, and passage of pathogenic spirochetes and other organisms, but it is advisable first to mention more recent studies of feeding organs and mechanism. Feeding and Digestive Organs The capitulum and related organs of 0. moubata have been studied in considerable detail by Bertram (1939) and reviewed in relation to these organs throughout the Arachnida by Snodgrass (1948). Both papers, which also review previous studies and concepts, deserve careful study. Because of their specialized nature, a short abstract of either of these two studies hardly meloiees does it justice. Snodgrass observes that "the exact method by which a tick bites perhaps needs more study than has been given to it", According to Bertram, the capitulum of 0. moubata is es. sentially similar to that of other argasid ticks (See Christophers 1906 for 0. savignyi, Robinson and Davidson 1913A,B,1914 for A. rsicus, True ieee for 0. coriaceus, and Sen 1934,1935 for O. ranieran although certain modifications in the eyeless tampan are either absent in other species or have not been adequately described. Bertram also differs widely from Sen in explanation of specific structures and fundamental interpretations. The capitulum (Figures 59 and 60), situated in a depression (camerostome) of the anteroventral body surface, consists of a median hypostome flanked by a pair of four-segmented palpi and a pair of long, shaftlike chelicerae arising from a conical pro- longation of the basis capituli. Each of these hollow appendages contains haemocoele. The hypostome is concave dorsally; ventrally it bears rows of distinctive retrograde denticles. The chelicerae distally each bear a small, triangular, articulated digit, at- tached by flexor and extensor muscles, with laterally directed denticles. These digits make the initial incision in the skin. A triple sheath arrangement of no little complexity encases the chelicerae proximally. The buccal canal (i.e. "mouth" ) lies between the dorsal chelicerae and the ventral hypostome; proximal- ly it is much compressed. The size of this canal is somewhat in creased by the medial emargination of the closely appressed che— liceral sheaths and by the dorsal groove (“gutter”) of the hypo. stome which forms a food conduit. Extending into the center of the buccal canal is a hollow, “tonguelike process”, the basal fusion of which with the hypostome forms a dorsal, blindly-ending pouch, the buccal cavity, into which a salivary duct issues at each posterolateral angle. The buccal canal opens directly into the pharynx, as one might logically assume it should, except that previous workers have found tnat in other ticks the basal fusion of the hypostome, palpi, and dorsal conical prolongation of the basis capituli causes the pharynx to open into the floor of the buccal cavity. Slee = TH BCH , 7. --BUCCAV. ee ema Oph f SCH) SUB.CHP HD.. CH : = -M D.PH Z =. ry QCH---~ hint ai BUC.CAN. ~ r < Ue SAN . ->>M.CPH H.-- = / M.D.HG. tubes. Lat a O.PH. lies ‘CAM. M.D.PH. | : basis capituli M.D.HG. dilator of hypostomal gutter expanded base of chelicera M.D.PH. dilator mscles of pharynx buccal canal O.PH. pharyngeal orifice buccal cavity PSGi.Gi. posterior part of closed camerostome chamber chelicera Teds pharynx cone sheath S.CH. outer sheath digit of chelicera S.CH. inner sheath dorsal conical prolongation SUB.CH.P. subcheliceral plate of basis capituli TG. tongue.like process salivary duct THe transverse bar formed by fusion flange of chelicera of lateral thickenings of hypostome tongue—-like process hood V.R. ventral rod of tongue-like constrictor muscles of process pharynx V.S. ventral scutum in cavity of closed chamber Figure 59. Longitudinal vertical section, diagrammatic f After Bertram (1939)17 ORNITHODOROS MOUBATA CAPITULUM PLATE XXT Silsee = Figures 60 and 61, dorsal and ventral views ORNITHODOROS MOUBATA CAPITULIM PLATE XXII When preparing to engorge, the tick inserts the chelicerae and hypostome (not the palpi 3 into the host skin as far as the dorsal conical prolongation of the basis capituli. During feeding (according to Bertram), the dilatation and constriction of the pharynx by certain muscles cause the fluid contents of a closed chamber just posterior of the tonguelike process to be forced into and sucked out of this process through a vertical septum. Furthermore, relaxation of hypostomal muscles obliterates the hypostomal gutter as the dilated pharynx constricts to force ingested blood into the esophagus. The effect of this swelling of the tonguelike process and closure of the hypostomal = 1047 = furrow is to prevent the spilling back into the host's wound of any blood already in the pharynx. The tonguelike process also appears to play an essential part in the mechanism of ejection of salivary fluid into the blood as it is being ingested. Since the salivary fluid is discharged into the distal region of the buccal canal, it is assumed to reach the wound in the host (and thus might transmit disease-causing organisms contained in it). As stated above, Bertram's study has considerable practical value, but must be read in its entirety to be fully appreciated. It should be noted that Snodgrass (1948) refers to the tongue— like process as the labrum in his noncommittal review of Bertram's findings and conclusions. Alimentary canal: We now return to Christophers! (1906) study o : Savignyi, and it is interesting to note that he found the pharynx to open into the floor of the "mouth" (buccal canal) in contrast to Bertram's observation on 0. moubata, mentioned above. At any rate, the pharynx leads to a narrow, straight esophagus. The latter, after perforating the central ganglion, enters the enormous saccular midgut, which, with its diverticula, forms the great bulk of body contents. Posteriorly, an extremely fine canal, which appears to be a functionless rudiment, joins the midgut with the rectal ampulla (but in 0. moubata even this is absent and the alimentary canal ends in a completely closed sac separated from the rectal ampulla) (see below). Esophagus and proventricular fold. The esophagus, a short, straight aS perforating the central ganglion in its course from pharynx to alimentary sac, is lined with a layer of clear columnar cells with small nuclei. The irregular outlines of these cells are mutually adapted to one another in a dovetailing arrange ment. At the juncture of the esophagus and large blood sac there is a small solid organ. This organ, in section, consists of a thick fold of epithelium of the same general character as that in the esophagus but of more columnar and less irregular cells. In the fold are some thick circular bands of muscular tissue and outside are longitudinal fibers passing from the esophagus to the gut. The epithelium of the fold passes imperceptibly into that of the esophagus, but ends abruptly on reaching the wall of the alimentary sac. The organ is very similar to, though = lo57= still more rudimentary than, the proventricular fold in the ms. quito and probably has a similar function. The importance of this fold in the trypanosome infection of Culex makes its pres— ence in ticks of concern in connection with spirochete infection. Aliment sac and its diverticula. These organs, when fresh. ly distended with blood, form smooth, dark red, lobulated masses. As the amount of blood diminishes, the diverticula become almost black in color and exhibit innumerable small lobulations. In the young, unengorged tick, the long, narrow diverticula show active pulsatile movements that probably have no effect in drawing blood from the host but serve to distribute fluid to dif. ferent parts of the sac. From the entrance of the esophagus and extending posteriad to the neighborhood of the rectum there is a large central reser- voir. This reservoir extends anteriorly a little beyond the entrance of the esophagus so that this latter is situated upon the ventral surface of the sac. From the ventral surface poste. riorly a conical tag passes ventrally behind the spermatheca to- wards the rectum. The basal part of this tag contains blood, but as it narrows it becomes a clear tube of capillary character. This portion of the canal appears functionally inactive and can play no part in the passage of matter from the sac to the rectum, In 0. savignyi, therefore, the alimentary system is practically a closed one ‘£ but in 0. moubata it is entirely closed; there is no passage between the Small intestine and the rectal ampulla (Enigk and Grittner 1952) 7. From the central reservoir a number of blind diverticula originate, the disposition and extent of which are constant in all ticks. There is an anterior, a lateral, and a posterior series. The anterior series, found only in Ornithodoros, is absent in Rhipi- cephalus and Hyalomma. It consists of a single, small median dit verticulum extending anteriorly so as to lie over the gene's organ. The three pairs of lateral diverticula arise together at about the level of the entrance of the esophagus. The anterior lateral di- verticulum is short, the median one is somewhat longer, and the posterior one is large and long. The anterior and middle branches divide into two or usually three blind pouches. The posterior = 1607— branch divides into two branches that curve to the ventral surface. The more anterior of the branches ends beside the common genital duct. The most posterior surrounds the anus and ends a short dis— tance anterior of this structure. Irregularities in diverticula arrangement are sometimes seen. The median lateral diverticulum may be large and give rise to the anterior of the two ventral branches. The various sulci and prominences on the surface of Ornithodoros have relation to these alimentary diverticula. On the dorsum, the transverse sulcus limits posteriorly the central alimentary sac. Ventrally the region between the coxae supports upon its inner sur. face, with which the viscus is in actual contact, the caecal ends of the posterior lateral diverticula. The lesser prominences cor-— respond in nearly every case with a particular diverticulum and the sulci with the intervals between two diverticula. Structure of the aliment sac. The structure of the sac and its divert Se SS Se entical. The cavity is lined by a single layer of large cells resting upon a thin basement membrane. Ex— ternally, very large single muscular fibres, arranged circularly and longitudinally, form an open meshwork with square meshes as in the mosquito. The lining epithelial cells are large with reticular protoplasm and large vesicular nuclei, some of which project freely into the lumen. Such cells are especially large and have their inner portions much swollen and vacuolated; they may contain dense black globules as well as red cells in various stages of intracellular digestion. In addition to large pro jecting cells, smaller cells, whose nuclei are situated nearer to the basement membrane, are present. Practically all cells of the sac contain small black granules, evidently derived from the digestion of the blood in the lumen, In undistended diver- ticula, the epithelium may form a mre or less continuous lining of the tube, but in the distended tube the cells become very un. evenly distributed, being almost absent in some places and in others forming very striking projecting masses. Rectum and malpighian tubules. The rectum, which lies im mediately behind the spermatheca, or the white gland in the male, is an irregular sac having several capacious but short saccular dilatations. (In 0. savignyi) it receives the rudiment of the dls) intestine and the two malpighian tubules (but in 0. moubata the rectum receives only the malpighian tubules and there 1S no com nection between intestine and rectal ampulla). Its walls are extremely thin and consist of a single layer of flattened cells. It contains a white fluid identical with the secretion of mal pighian tubules. The rectum therefore does not serve as an ad junct to the alimentary canal, but functions as an excretory receptable. The white matter passed ine anum by ticks also can not, strictly speaking, be regarded as feces. The malpighian tubules are important because of their great length and functional activity and because of the frequency with which such organs are utilized by parasites in other animals. They consist of two long, fine white or transparent tubules arising on either side from the rectun, and after a complicated course among the viscera, ending blindly in the anterior portion of the body. These tubules come in relation with almost every important organ in the body and drain every quarter of the body cavity. The tubes in young ticks are of an even calibre through. out and contain small quantities of secretion only. They are often swollen in aged ticks to form sacs similar to, but smaller than, the rectum. After oviposition the tubes and rectum may be greatly distended with characteristic white fluid that is evident externally as patches of lighter color. It is probable that the appearance is that described as a “fungus” in these ticks by Well- man (19064 ,D ,1907B). Feeding habits of 0, moubata have been discussed in the sec- tion on the life cycle of this tick. Certain aspects of digestion and excretion of fluids are of considerable practical importance for they concern, at least, the fate of ingested spirochete para sites of relapsing fever (Borrelia spp.) and the discharge of these organisms onto or into a new vertebrate host when the tick subsequently feeds. Digestion The volume of a blood meal is from two to six times the tick's original body weight (Lees 1946B). During ingestion, the cuticle stretches to accomodate this huge amount of fluid. Engorgement is completed in about half an hour. In order to reduce this tremendous volume ingested so rapidly, fluid is discharged from the coxal or- = 16S; = gans while the tick is yet feeding. Rectal discharge is very slight. These two methods of excretion are discussed in separate sections below. Digestion in O. moubata apparently is much like that in 0. savignyi, as described by Christophers, abstracted below. Twenty-four hours after a meal the greatly distended di- verticula contain a soft coagulum from which a considerable amount of fluid blood may drain. Blood corpuscles are apparently un. changed. Scattered through the fluid are numbers of intensely black, globular granules measuring from 5u to .5u or less in diameter. In sections these granules are collected especially at the periphery of the (fresh) blood, but they are also present in large numbers scattered throughout the mass. The black gran. ules are derived from a previous meal, and there is therefore a considerable degree of mixture between the new blood and the contents of the diverticula prior to the meal. Diverticula examined at some considerable time after diges. tion show a number of reddish granules lying in the still partial. ly fluid blood. These are free from attachments and when washed out fall to the bottom of the dish or among the viscera. Each is an entire cell containing a well-marked nucleus. Films of the sac contents made twenty-four hours after a blood meal show cells derived from the epithelium of the sac in addition to the host's leucocytes. Many of these are evidently the smaller un distended cells, previously noted as lying near the basement mem brane, now detached in preparation of the specimen. They contain a large circular or oval nucleus and finely reticular or partial. ly vacuolated protoplasm. Similar cells, but larger and with portions of the vacuolated protoplasm stored with black granules, are also seen. In addition to these cells of the sac epithelium, there are other large, dark staining, circular cells with rather small nucleus. Their substance is markedly vacuolated and crowded with matter that they evidently have engulfed, blood corpuscles, black granules, chromatin fragments, etc. In section specimens made even six hours after the ingestion of blood, they appear lying apparently in isolated positions far removed from the sac walls. These probably function as wandering digestive cells. Their rela. tion to the epithelium of the sac is not clear. As digestion ad SyiliserS vances they become more and more replete with material and in. crease in size until readily visible to the naked eye as red granules already noted. In early stages of digestion, cells packed with chromatin bodies and superficially resembling macro. phages, the nature of which is not clear, may be seen. Although a prominent part in digestion is taken by the free cells just alluded to, epithelium lining the diverticula also takes an active part in the process. The swollen and vacuolated portion of the large projecting cells is crowded with products of digestion very much as is that of the free cells. Smaller cells lying nearer the basement membrane are also, as a rule, packed with fine black granules, though they rarely contain the large granules seen in the other cells. The intensely black and opaque globules are highly charac— teristic of digestion in the tick and undoubtedly represent the ultimate condition to which blood remaining in the gut is reduced by the digestive process. These globules probably represent only the portion of food not assimilable, for in Ornithodoros ticks, which may be kept alive for long periods without food, the diver- ticula contain, after some weeks, an inky black material consist— ing entirely of these granules. As diverticula contents are digested, the muscle fibres, which in the fully distended organ slightly indent the surface, sink more and more into the body of the viscus. The wall between the fibres becomes ballooned and eventually forms flasklike pock ets with only a narrow opening connecting with the lumen. The epithelium is, as a rule, present in the pockets, though Leas generally more noticeable on the ridges formed by the contracted muscular fibres. Remains of ingested blood, in the form of black granules, are present both in the pockets and in the lumen. Ticks examined months after a meal still have the diverticula loaded with the black material. Waste matter is not passed into the rectum and any remnant of food not absorbed must remain in the diverticula until death of the tick. The method by which absorption takes place has not een ascertained. Black pigment is not detected in the tissue cells or in the body cavity. Note that excess fluid in the blood is excreted by the coxal organ during and following feeding so that a large amount of blood can be rapidly ingested; this is elucidated in the section on the coxal organ below. = 170 = Wigglesworth (1943) confirmed that in 0. moubata, blood (haemoglobin) is absorbed by swollen epithelial cells of the wall of the large stomach and its diverticula. In these cells, which detach and remain free in the lumen, blood pigment is converted into black globules that are ultimately discharged into the gut cavity. Similar dark granules are dispersed through smaller cells of the gut wall, but no black pigment can be seen in other tissues or in the body cavity. The haemoglobin is digested more or less to protohaematin and is demonstrable in the tick's haemo_ lymph probably as alkaline haematin. The gut contents are reddish brown with black haematin deposits. No free iron can be detected in the gut lumen or cells, or in other tissues, and no nephrocytes containing haemoglobin derivatives can be found. The type of host from which 0. moubata draws a blood meal may be identified by the precipitin test more than six months follow. ing feeding (laboratory studies at 20°C. and eighty percent rela tive humidity) (Weitz and Buxton 1953), or even for twelve months (fowl blood meal, kept at 30°C., ticks also fed on mouse) (Gozony, Hindle, and Ross 1914). Rectal Excretion As stated above, 0. moubata has no passage between the small intestine and the rectal ampulla, and defecation does not occur. Excretion of water ("urination ) from the malpighian tubules takes place only after the first nymphal stage has been reached and a blood meal has been absorbed; this excretion is viscous and dries within a few hours. In the weeks following the first excretion only a slight amount of water is irregularly excreted (but can be produced through various stimuli). This pattern is similar in each developmental stage after the larva. Variations in tick excretion and a comparison of this function according to species, morphology, number of hosts, size, duration of development, quantity of blood ingested, and transmission of disease organisms to vertebrate hosts have been analyzed by Enigk and Grittner (1952). -171- Coxal Organ Morphology and Function* Inasmuch as the volume of the blood meal is from two to six times the tick's original body weight and engorgement is usually completed in about half an hour, the tick must have a means of reducing the total intake volume and of preserving the internal medium while feeding. For this purpose coxal organs function as ionic (chloride) regulators and for ultrarapid excretion of a large volume of water during ingestion of blood. Coxal dis. charge, which commences about fifteen minutes after the tick has begun to feed, continues till completion of the meal and inter mittently for about an hour afterwards. (See also Lavoipierre and Riek 1955). Malpighian tubules do not function until about an hour after feeding is completed, and the amount of water they excrete is limited. Chloride regulation. About half the ingested water is ex ereted in coxal fluid. The mean haemolymph chloride concentration before feeding is 1.00% and after feeding 0.96 NaCl; that of coxal fluid is 0.806 NaCl. These values are similar to those determined by Boné (1943). Morphology of coxal organs. The flaskshaped coxal organs, which elaborate the bulk of fluid, consist of an outer filtration chamber with an inner tubule system leading to the external open. ing and of a small organ with glandular structure, the so-called accessory gland. The filtration chamber, which communicates with the tubules of only one point, is highly folded into an elaborate series of pockets and fingers that closely invest the tubules; oe eS ee *Chiefly from Lees (1946B). See other remarks in section on life cycle. It should also be mentioned, for practical significance in relation to disease, that Lees found that 0. delanoeéi acinus and 0. parkeri have coxal organs differing from those of 0. moubata, and that in these species coxal fluid is liberated only after cessation of ingestion. It should also be noted that what Lees and others have called "coxal gland" is rather a coxal organ (Burgdorfer 1951) because it excretes fluid rather than secreting fluid, and the filter chamber histologically has no glandular structure. We = numerous small muscle fibres inserted in these pockets pass out— wards from the organ to attachments on the body wall. The histol. ogy of the two regions is entirely different. The filtration membrane is only one or two microns thick and its cellular origin is much obscured. The tubule walls, from five to thirty microns thick, are composed of cells with a dense, deeply-staining cyto- plasm and are richly supplied with tracheae. Function. The production of coxal fluid is under muscular control. .t is believed that contraction of coxal organ muscles enlarges the filtration chamber and sets up a sufficient pressure difference across the membrane to initiate filtration into the organ. In subsequent passage of fluid down the tubules, threshold substances such as chloride are reabsorbed. That the coxal fluid is primarily an ultrafiltrate of the haemolymph is suggested by (a) the rapid passage of dyes and even haemoglobin into coxal fluid after injection into the haemolymph, and (b) the very high rate of fluid liberation. Serum albumin sometimes passes into coxal fluid after injection, but casein (and normal haemolymph proteins) are fully retained (Lees! summary). Boné (1943) proposed somewhat different explanations concerning coxal organ function. Lees further indicates that Patton and Evans! (1929) opinions regarding the functions of the coxal organs are im correct. An earlier work on the same subject is that of von Kunssberg (1911). The small accessory coxal glands have an unknown function. Rapid engorgement in argasid ticks is allowed by passive cuticular stretching. In ixodid ticks new cuticle is produced to allow for volume of intake and engorgement is much more slowly accomplished. When specimens of 0. moubata and other argasid ticks that possess coxal organs are warmed or irritated they exude from these organs a clear fluid. This may possibly serve in part as a defen. sive mechanism although the actual reason remains to be determined. Coxal organ discharge has been observed and reported, highly in. accurately, by Remy (1922A, and for Argas reflexus, 1921 and 1922B), who believed the exudate to be haemolymph containing haemocytes. Lees (1946B) has shown that these structures are actually small, globular clusters of refractive granules, possibly derived from - 173 = partial regression of the salivary glands during molting or from granule.bearing cells in the accessory organs. The density of these granules in the coxal fluid of newly molted but unfed ticks is much greater than in the fluid of engorged ticks, in which the granules are more widely dispersed in the greater amount of fluid. Spiracular Morphology and Function Argasid spiracles have been described by Robinson and Davidson (1913), Cunliffe (1921), Mellanby (1935) and Browning (1954A). The last two workers paid particular attention to the spiracular struc— ture and function of 0. moubata. As described by Mellanby, the spiracle consists externally of a semicircular cribiform plate inserted into a smooth macula of thickened skin, with a slitlike ostium between these. The thin external layer of the plate is supported by rodlike pedicles. The external layer was stated to be pierced by minute pores opening into the tracheal atrium, which is a tube connected to the ostiun. Muscular attachments of the maculaallow opening and closing of the ostium. If it were true that the external layer is pierced by pores, it would appear that there is no way for the tampan to close off the direct connection between the external air and the internal body tracheae. Since tampans show remarkable ability to withstand desiccation in the laboratory and in nature, Browning (19544) was led to investigate the spiracle anew. He found that a surface view of the spiracular plate gives the impression of being porous. On examination of transverse sections these "pores™ are shown to be expanded distal junctures of branching pillars (pedicels) arising from a basal, underlying layer of sclerotized endocuticle. These pillars support the very thin outer membrane, which is, however, not porous but continuous. The cavity between the basal cuticle and outer membrane and ramifying between the pillars is continuous between the atrium and the spiracle. From surface view the hard maculum can be seen between the inner curves of the crescent of the spiracular plate. The macula encloses a slitlike aperture, or ostium, connecting the atrium of the trachea with the outside air. The argasid spiracular plate functions to provide a pad against which the macula can impinge when depressed = 174 = IITIXX GLVId Cwrs6r)_Suqunosg s0a5v_J ATOVULdS VIVENOW SOMOCOHLINYO *1JOT Sud SPTeMOL TIOTIeqUB S7T pue OMsTF ouy Jo doy oy4 spzemoy qoodse Tesiop SqT YYIM pequetso eToeitds yyTM ‘MeTA eoejams f¢9 emsTy *€9 OMIBTY UT A-K eueTd ut ynoqe uoTYoes esseAsuety ‘79g OITA \ 3, ©) ie SSMS fb \ ee) zi . do J - 175 — and thus to form a very efficient seal of the ostium, a necessary condition for animals so likely to be exposed to desiccation. It is of interest to compare Browning's study with that of Arthur (in press) on the spiracle of Ixodes 7. According to Mellanby (1935), the physiological reaction that governs the opening and closing of the spiracular ostium is similar to that of insects. Browning (1954B) appears to accept this conclusion. The physiology of spiracular action has been discussed under Environmental Adaptability above. Haller's Organ The structure of haller's organ and its pot value as a phylogenetic indicator has been discussed by K. W. Neumann (1943). Schulze (1941) also described and illustrated haller's organ, which functions as an organ of smell. Incidentally, Zumpt (1949) summarized his studies on the systematic importance of this structure as follows: Research up to now indicates that haller's organ will have to be considered in the future as having a role in tick systematics and should not be overlooked. Abnormal Development During examination of about eight thousand laboratory reared specimens of 0. moubata, Robinson (1943A) encountered two examples of partial twinning of the posterior area in a third instar nymph and in a fifth instar nymph that subsequently molted to a male and female, each with abnormalities in internal anatomy. These specimens were normally fertile. Another peculiarly humped third instar nymph normally molted to a male that showed suppression of the postanal region. This male failed to copulate although the genital system was well developed and the sperm normal. Robin. son (1944B) also noted many abnormalities of the legs during handling of about ten thousand tampans. Most cases were deficiencies due to partial regeneration of a leg damaged in a previous nymphal instar. Two cases of supernumerary segments of legs were also observed and illustrated. In the same batch a nymph that was much more complete. ly twinned posteriorly than previous examples was observed. This specimen molted to a partially twinned female, mated normally, refused to feed, and deposited a small egg batch (not particularly = 176 = unusual for unfed females). Some of these eggs developed into normal ticks but most were not delivered into the arms of gene's organ and therefore did not hatch. In the same paper, Robinson reviewed reports of partial twinning in other tick species. leg anomalies have been reviewed by Campana (1947). Symbiotes Tissue cells of many normal insects and ticks harbor living microorganisms that for the most part exert no harmful effects on these cells. In fact, some of them may be distinctly benefi- cial to the hosts, carrying out their part of a mutually helpful relationship. / Steinhaus (1947) 7 In some respects, the relationships between arachnids and their symbiotes are very similar to those between insects and theirs. Among noteworthy differences, however, appear to be the absence of mycetomes in ticks, though some mites have these struc— tures. Furthermore, most tick symbiotes occur in the malpighian tubules and in the ovaries instead of in the alimentary tract, though this may not be true for certain of the rickettsiae. The two families of ticks are similar with respect to symbiotes; in both the same organs are associated with microorganisms. They differ, however, in the manner of ovarial infection. [ Steinhaus (1947) 7 Intracellular clusters of large masses of typical rickettsiae were discovered in salivary gland acini of 0. moubata by Hertig and Wolbach (1924). Intracellular symbiotes were not found in larvae (?nymphs) of 0. moubata by Cowdry (1925C,1926A,1927), though they were demonstrated in Argas persicus and in Otobius megnini., Other extensive reviews of symbiotes in ticks are those of Mudrow (1932) and Jaschke (1933). In 0. moubata, unlike ixodid ticks, symbiotes, probably of a bacterial nature, do not occur in the anterior ends of the malpighian tubes but rather in about one-fifth of the length of the tubes just posterior of the anterior ends. In this, 0. moubata differs from A. persicus, in which Jaschke observed intra cellular symbiotes in masses aS large as five microns in diameter and containing as many as forty individual organisms each. ae TET ve In 0. moubata and other argasids, symbiotes may migrate from the malpighian tubes to the ovaries and developing eggs, thus differing from ixodids in which they directly invade the first sex cells (Mudrow 1932). This worker sought the explanation of symbiotic bacteria in the realm of physiologic relations of nw trition. (See three paragraphs below). Argasid symbiotes do not appear to be as pleomorphic as those of ixodids and are usually of the rod or coccus type though they are grouped into apparently gelatinous masses or colonies. Rows or chains of granules or filamentous bundles are not seen in these masses. Tick symbiotes have not been artificially cul. tivated although Steinhaus attempted to do so with those from Argas sicus by utilizing fluids and tissues of the chick embryo. / Steinhaus (1947) 7 With reference to the “bactericidal action" in the guts of insects and A. persicus and 0. moubata (Duncan 1926), the reader is referred to subsequent findings in the following series of papers on work done with A. persicus: Anigstein, Whitney, and Micks (1950A,B), Whitney, Anigstein and Micks (1950), and Micks, Whitney, and Anigstein (1951). The intestinal tract of blood engorged ticks exhibited significantly higher antibacterial titer than those that had not been fed. Study of animal blood itself revealed erythrocytic enzymatic hydrolysates showing marked in vitro antibacterial effect over a relatively wide spectrum of most gram-positive and a few gram-negative organisms. The active principle of the hydrolysate appears to be a peptide amino acid complex, called sanguinin, which, as a powerful enzymatic inhibitor, represses the growth of several organisms including streptococci, both in vitro and in vivo. The role of symbiotes in producing growth-promoting substances in 0. moubata (and in bedbugs) has been studied briefly by De Meillon and Goldberg (1947A,B). Feeding nymphal and adult ticks on thiamin.deficient rats resulted in almost doubling the time necessary for completing the tampan's life cycle, increasing the interval between blood meals and molting, and an additional molt before reaching maturity. Normal growth and reproduction, however, follow feeding on riboflavin-deficient rats (De Meillon, Thorp, and Hardy 1947). The purpose of these experiments, fol- = L/S = lowing work by Brecher and Wigglesworth (1944) on the blood-sucking hemipteron Rnodinus prolixus, was to test the ability of symbiotes in the tick to produce growth-promoting vitamins in the absence of these substances in host blood. Thiamin, it appears, cannot be manufactured by symbiotes under these conditions but riboflavin can be produced in sufficient quantities for normal development and reproduction. Incidentally, it was noticed that the severity of host skin reaction to bites of 0. moubata is greater in animals that are deficient in thiamin than itis in normal rats. In the former, an extensive hemorrhage develops at the site of each bite (De Meillon and Goldberg 1947A,B, De Meillon 1949). Laboratory Rearing Methods This subject has been discussed in more or less detail by all students of the life cycle, mentioned above. Methods for rearing OQ. moubata, care of hosts, caging, precautions, host diet and handling, etc., have been presented by Harvey (1947). Artificial Feeding A capillary tube method for the artificial feeding of 0. moubata and other ticks for studies of disease transmission and physiology has been developed by Chabaud (1950A). Prevention and Control* Prevention Travellers in infested areas should be cautious especially in choosing sleeping and sitting sites. Indigenous habitations whenever possible should be avoided for sleeping, and care should *Although it is not the policy of this study to deal with control and prevention subjects because these are more logically included in a separate report now being prepared, an exception is made in the case of 0. moubata. The control and preventive measures re— quired for this species are unique among African ticks, and its biological and host predilections are different from all others. Moreover, it is possibly the only medically important African tick that has little or no veterinary importance. - 179 - be taken to protect one's self from tampan bites in rest houses, berracks, meeting places, and sometimes in European houses. In Tanganyika, Morstatt (1914) noted, those huts that were exposed to the rain were free of ticks while others in places more pro- tected from the elements harbored tampans. Morstatt suggested camping in grassy spots some distance from huts. Any program of labor introduction from an infested erea should include an initial inspection of newcomers! personal effects, bed— ding rolls, and extra clothes. Strict sanitary measures are of proven success in labor camps. If floor and walls are hard, dry, and free from all cracks and if dust and unnecessary objects that might provide concealment are removed, the tampan's hiding places may be kept to a minimun. Frequent inspection of personal effects, which should be kent in tightly closed boxes or cabinets, or hung away from walls, are of proven success. Persons living in barracks should be warned to report the presence of ticks. Beds must be provided and mos— quito nets may be necessary. In infested buildings, placing of bedlegs in cans of kerosene has been recommended to deter hungry tampans. Special tickproof construction of military huts in heavily infested East African areas has been recommended (Hynd 1945). The base is a six-inch deep bitumen floor (or cheaper hard—beaten tar and earth) with a metal strip inserted at the outer edge nid— way throush its thickness and projecting three inches outwards to prevent ticks from reaching the floor level from outside. A second strip, about one foot above the floor level and extending both inside and outside, helps to confine the searching area for ticks brought in on clothes and gear. Hynd found that the tampan climbs upwards only when it is not able to burrow into the ground. It searches for hiding places in wall cracks or roofing but can not circumvent horizontal metal strips extending outward from walls. Jack (1928,1935,1942) suggested that pigsties be constructed of smooth concrete that is easily cleaned and does not provide a hiding place for tampans. A valuable account of buildins methods to eliminate tampan infestation has recently been published by Annecke and Quin (1952) and Annecke (1952). Reinforced concrete buildinss, which replaced wattle and daub huts at a cost of L 13 per person, are said to have reduced deaths from relapsing fever on a large South African citrus plantation from forty to a single case annually. Addi- tional benefits resulted from employees! social and economic betterment. These buildings were constructed by casting walls in a steel franed mold and raising precast concrete roof sections over them. Floors were made from a vermiculite and cement mixture. Acceptability to indigenous labor, ease of cleaning, and crack proof construction were important considerations. As the economic level of African labor rises, such prevention methods will become normal and expected, but today they are revolutionary. The effect of domestic animals on 0. moubata populations in houses appears to be moot. Under usual conditions it seems that domestic animals allowed to live in human habitations tend to allow an increase of ticks in these buildings. An exception is cited by Walton (1950A), who believes that in areas of high humii dity additional moisture provided by domestic animals in huts is enough to discourage the tamoan. Rooms in which goats are kept in humid Kenya hills are free of ticks (Teesdale 1952). Inasmuch as chickens often are said to be a favorite nymphal host, they probably should be excluded from buildings except pos sibly for periodic forays to feed on ticks near the surface of the ground. Knowles and Terry (1950) reported that chickens in Tanganyika are heavily infested with nymphal O. moubata, but Phipps (1950) found no significant relation between the presence of ticks and fowls in the same area. Chemical Control Gammexane (Hexachlorocyclohexane, BHC or benzene hexachloride) is generally considered to be the most promising chemical for controlling 0. moubata. Ticks coming in contact with 0.5 per cent dust lose co. ordination after five or six hours. During the first day they lose muchadf their body weight by excessive coxal fluid loss, def- ecation of milky fluid, and possibly by increased integumental alicia permeability. Their color darkens; they become more or less im mobile and usually die in a little over a week. Those that live fail to produce viable eggs. Application of 0.5 per cent gamme— xane dust to floors and lower parts of walls sprinkled from per forated cigarette tins at the rate of three or four pounds of dust per hundred square feet is recommended, but frequent checks en be made where reinfestation is liable to occur (Jepson 1947). Application of Jepson's findings on a township scale in Tanganyika was described in detail by Knowles and Terry (1950) using "G dust” (D220 compound one part, and diatomite four parts). These authors found that although total eradication is probably impossible, a townshipwide control program can almost eliminate relapsing fever in a fairly static population and is much cheaper than hospital treatment of the disease. Sampling methods and application methods were also stressed. Factors to be considered in a control program and need for further research were discussed by Phipps (1950), who indicated the necessity of using tested diluents and making accurate sur veys before and after treatment.’ The incidence of ticks and dis. ease should also be checked before large scale programs are instituted for, in Phipps* opinion, not all places in which ticks abound are disease foci. In Annecke and Quin's (1952) extensive chemical control program on a heavily infested South African citrus plantation, it was found that BHC applied as a spray to inner walls of huts in concentrations of 300 mg. gamma isomer (or upwards) per square foot (emulsion of 17% BHC with 4% gamma isomer) gave effective control. With lesser concentrations, tick populations increased rapidly after seven weeks. Surviving ticks from sprayed huts deposited considerably fewer eggs than normal females. Nevertheless, the same authors report, reintroductions were so frequent that the authorities finally resolved to construct tickproof habitations. In spite of comparatively high initial cost in comparison with daub and wattle huts, this was found to be the only realistic approach to reducing deaths and man hours lost to relapsing fever. = Le2y= Subsequently, Annecke (1952) reported that all huts treated with 600 me. gamma isomer per square foot BHC remained free of 0, moubata for at least 27 months. When 300 mg. per square foot ~ were used, huts remained free of ticks for twelve months. The small amounts of BHC used in malaria control programs, 25 to thirty mg. gamma isomer per square foot BHO wettable powder, applied twice three months apart, has little effect on O. moubata, but Annecke (loc. cit.) believes the cumulative effect may reduce or destroy the tick. The above remark immediately suggests the potentiality of tampan resistence to chemicals. So far as known, none has yet been demonstrated. A control program in Fort Jameson township of Northern Rhode sia has recently been reported by Holmes (1953). Heavy spraying with a five percent BHC solution resulted in general diminution of the tick population but did not result in its elimination. It was found that the most economical form for achieving a hundred percent kill was five percent BHC powder mixed with 95% (by weight) sawdust diluent, or with chaff left from pounding corn. To obtain a sackful of this mixture, a container holding 6¢ pounds of forty percent BHO cattle dip wettable powder was mixed with fifty pounds of sawdust. This mixture was then laid as a four inch wide bar rier, thick enough to insure that ticks mist come in contact with it when passing, at the base of all inside walls of the house. The barrier was maintained for three weeks and frequently inspected, especially where it passed door openings, to insure that it was not scattered. This time period should reach all nymphs hatching from eggs laid before the chemical is applied (more or less ex. ceptionally a somewhat longer hatching period may be involved, but for practical purposes this time is probably usually effective - HH). Elimination of tampans was obtained by this method, but re— infestation occurred within a year to a year and a half. Fleas, Congo floor maggots, and bedbugs were also killed. In the fol- lowing year, the number of cases of relapsing fever from treated houses was considerably reduced. The cost of this program was so low that other authorities commenced similar control programs. ~- 163 A control program using 0.5 percent gammexane powder (D.034) was undertaken in houses in humid Kenya hills (Teesdale 1952). Reapplications had to be made every month or two to control nymphs that had hatched from eggs, the latter being resistant to the chemical. Killing effects diminished in treated huts fifty days after application. The low cost of gammexane was said to allow its purchase and use by Africans. Although most recent workers favor BHC dusts over spray so- lutions, Anderson (1947) reported good control in Somaliland coffee houses with a three percent solution of gammexane in diesoline,. “666 spray™ (crude benzene hexachloride, 124 gamma isomer) at a dosage of 1,250 mgm. per square foot of soil was recommended by Hocking (1946), who found DDT at the same dosage less effective. On the basis of experiments in two localities in Kenya, Heisch and Furlong (1954) recommend a spray of gammexane wettable powder P.520 rather than gammexane insect powder for tampan control. Investigators have reported that DDT is of less effective ness than gammexane in controlling tampans. Among these, Jepson (1947) found that five percent DDT dust is slower and less ef- fective than gammexane, although after about three weeks a mor- tality of fifty percent to eighty percent obtained. Holmes (193 ) also indicated that gammexane provides a more complete and rapid kill than DDT. Annecke and Quin (1952) considered that various types of DDT applications lacked sufficient residual effect to be considered useful. The inefficacy of many chemicals for killing 0. moubata and the usefulness of gammexane and of E605F (diethyl-p.nitrophenyl. mono=thio~phosphate) for this purpose were reported by Enigk (1948). Belgian tests with three preparations of the gamma isomer have been reported by Pierquin (1950). Sprays of "Cyclotox™ contain. ing a large proportion of the gamma isomer killed about half the ticks in eight to ten days when applied in Belgian Congo huts. In the laboratory, where the ticks could be kept in closer contact with the chemical, all died after varying lengths of time with different concentrations and preparations (Himpe and Pierquin 1951). The authors conclude that spraying a volatile substance on soil is of less value than mixing it with soil. = Leis The possibility of controlling 0. moubata by feeding hosts on certain chemicals was explored by De Meillon (1946). Fifty mgm. of pure gamma isomer of gammexane were mixed with agar and water and fed to rabbits four or five times. The ticks fed only briefly and showed either incoordination or death afterwards. The domesticated tampan's predilection for human blood obviously limits the application of this interesting approach. 0. moubata is also susceptible to arsenic compounds in the blood of animals. Injections of neoarsphenamine have been used in rabbits for this purpose (De Meillon, Thorp, and Hardy 1948). The failure of 2:3 dimercaptopropanal (British anti-lewisite) to alter the toxicity of neoarsphenamine was described by Thorp, De Meillon, and Hardy (1948). In testing insect and tick mortality when exposed to dry insecticidal film, Busvine and Barnes (1948) found that 0. moubata nymphs are resistant to DDT but susceptible to gammexane pyrethrins, Busvine and Nash (1953) also determined that films of oil solutions are better than dry films for testing insecticides because they give a sharper dose/kill relation. The value of certain derivatives of phenol and naphthalene as ag Pa reL ae in hut floors has been suggested (Robinson 1L944A ). Derris powder failed to affect nymphal 0. moubata in Russian laboratory tests (Mironov, Nabokov, and Kachalova 1040). Pyrethrum sprays and dusts are highly toxic (Robinson 19420 ,D,1943B,1944B)) but field tests have not been undertaken, probably due to high cost of pyrethrum and effectiveness of cheaper BHC. Sulfur dioxide or cyanide fumigation has little effect on QO. moubata, and sprays of kerosene and formalin are not success— ful (Hopkins and Chorley 1940). A spray consisting of 30 cc. turpentine, 50 cc. twenty-five percent alcohol, 5 cc. kerosene and a little white soap was suggested by these authors, though it is expensive and troublesome to prepare. Their best re commendation was a coarse, roughly filtered spray consisting of 15 pounds of paradichlorobenzene in one gallon of kerosene applied under high pressure at the rate of twelve to fourteen gallons per two thousand square feet of surface (also reported by Hargreaves 1936). AEBS The ovicidal value of concentrated vapors of chloroform, sul. furous acid, ammonium sulfide, ethel dixanthogen, BHC, formalin, and phenol has been reported (Enigk and Grittner 1953). A report of the resistance of 0. moubata to various sheep dips (Blacklock 1912) is of little contemporary interest. Disinfection of Personal Effects Blankets, bedding, and clothing may be disinfected by ex posure in a tight container to a temperature of $2°C. for half an hour (Hopkins and Chorley 1940). Burning In a carefully conducted experiment, Garnham (1926) found, contrary to certain textbook statements, that burning of African huts infested with 0. moubata is an effective means of killing these ticks. He suggested pulling down the straw roofing and piling it inside mud hut walls for burning. This may be the only means of control where the indigenous population does not work for pecuniary gain. Otherwise burning is uneconomical un. less the situation is serious. More permanent housing, in which infestation may be controlled or better still prevented, should be substituted whenever possible. Burning infested buildings has long been the indicated control method in many parts of Africa. Flame throwers are sometimes used to good effect where ticks are lodged in shallow cracks in buildings that withstand fire. Jack (1931) obtained control against tampans in nine-inch walls of pigsties by burning brushwood on both sides of them. Burning against only one side and spraying with a ten percent emulsion of paraffin (i.e. kerosene) had failed. Hand picking A reward of sixpence for every twenty ticks collected on a South African farm yielded 73,000 ticks oreyear and over half a million in several years. Laborers placed a thin layer of drift manure along the inside walls of their huts and there collected the ticks as they came to hide. Small holes dug inside and out side the doors and filled with drift manure were also found to be favorite hiding places (Annecke and Quin 1952). = L6G. DISEASE RELATIONS Man (In nature ) O. moubata is the only known tick vector of African tick-borne relapsing fever (Borrelia duttonii) of East, Central, and South Africa. A few cases of this disease have been reported from Equa. toria Province in the Sudan. No evidence supports certain pub. lished maps showing known extensive distribution of tick-borne relapsing fever in many areas of the Sudan. It should be noted that populations of this tick from burrows of wild animals have not been found infested with spirochetes. It is claimed that some specimens naturally infected with rickettsia, Coxiella burnetii, the causative organism of Q fever, have been found in Ruanda-Urundi, and that in Kivu others have been taken infected with an organism referred to as “Bashi virus— rickettsia™. The etiologic agent of food poisoning, Salmonella enteritidis, has been recovered from this tick in Africa. 0. moubata has not actually been found infected in nature with the pathogenic organisms of any other human disease, but experi_ mental data strongly indicate further research in this respect. Tampan bites may cause considerable irritation. Circumstan. tial evidence suggests that persons long victimized by bites of this tick may develop an immunity to them. Fowls QO. moubata is an experimental vector of fowl spirochetosis (Borrelia anserina). It is of negligible importance in transmission of Salmonella bacteria and of Aegyptianella pullorum (a protozoan). The bacterium that causes avian cholera (Pasteurella avicida) sur- vives only a few days in 0. moubata. asi = Experimental Disease Relations Viruses and Rickettsiae The use of 0. moubata and other Ornithodoros species for trans— porting a number of pathogenic organisms for experimental purposes has been suggested. This species is easily infected with Q fever (Coxiella burnetii) and is capable of transmitting the organism by its bite. A diagnos— tic test for Q fever, using 0. moubata for feeding on a suspected host, has been developed. — Rickettsia prowazekii, the causative organism of classical typhus, and R. typhi, that of murine typhus, develop in 0. moubata. The former eet can be transmitted transovarially and latter can be found in eggs and coxal fluid of infected ticks. No multiplication of R. tsutsugamushi, the causative organism of scrub typhus, occurs in 0. moubata; The tick is therefore not a likely vector of this unusually host-specific rickettsia. O. moubata maintains infection with the virus of Russian spring-summer (Far Eastern) encephalitis. It has also been shown that these ticks can be infected with western equine encephalitis virus. The virus causing murine poliomyelitis ete Columbia SK) is destroyed or inactivated in blood ingested by the tampan. Yellow fever is not transmissible by this tick. A Congo rickett— sia of the boutonneuse fever type (boutonneux-pourpre) remains virulent in ticks fed on infected guinea pigs for two months but not for six months. Bacteria Tularemia (Bacterium tularense) is transmitted by the bite of O. moubata and is also transovarially transmitted to progeny. Bacillus anthracis, the causative organism of anthrax, is not transmissible and apparently kills the ticks, although it does remain virulent in the tick's body long after death. The etiologic agent of food poisoning, Salmonella enteritidis, has been recovered from this tick in Africa. Lose Spirochetes of Relapsing Fever O. moubata is capable of harboring and transmitting other Borrelia species besides B. duttonii. This is of considerable interest since in other parts of the world many species of Borrelia are host-specific and tick hosts are spirochete-specific,. Spirochetes Other Than Relapsing Fever Weil's disease (Leptospira icterohaemorrhagiae) survives for about forty days in living 5 moubata, remains virulent in the body long after the tick's death, and can be transmitted when infected ticks bite. Filariae Filariae may develop in the body cavity of 0. moubata but transmission appears to be unlikely. Trypanosomes Virulent trypanosomes may remain in the tick's gut for as long as five years, but transmission is apparently impossible and transovarial infection does not occur. Toxoplasmosis It appears that Toxoplasma gondii cannot be transmitted by 0. moubata although the organism survives in the tick for almost two weeks after artificial inoculation. Tropical Ulcer The suggestion has been made that tick bites, such as those of O. moubata, may be initially responsible for tropical ulcer. IDENTIFICATION O. moubata might be confused only with 0. savignyi in the African fauna. However, since 0, mubata lacks eyes, which in O. savignyi are present as two pairs of small, round, equal sized, es ksss) = shiny black spots in the lateral groove, one pair above coxa I and one pair between coxae III and IV, there should be no real difficulty in distinguishing the species. Tarsal differences mentioned in Nuttall et al (1908) are too variable for applica. tion. Adults average about 8.0 mm. long by 6.0 m. or 7.0 m. wide, although freshly engorged females may reach 11.0 mm, in length. O. savignyi is usually somewhat larger, averaging about 11.0 mn. Tong. the mamnillated integument, conspicuous tarsal dorsal protuberances, and absence of cheeks easily distinguish both 0. moubata and O. savignyi from all other African species. Males are often a little ue than females and their genital aperture is a short, rounded opening as compared with the broad slit of the female. Nymphs have no genital aperture, but in larger instars a small round depression is present in its place. In a study of the sexual differences in this species, Nuttall (in Cunliffe 1921) has shown that the average male is smaller than the average female, though extremes overlap, and that the same is true for the size of the genital aperture of the two sexes. An estimate of the stage of development of immature stages can be made on the basis of differences in mouthparts, legs, and spiracles (Cunliffe 1921). = 190 — ORNITHODOROS (ORNITHODOROS ) SAV IGNYI (Audouin, 1827). (Figures 8 to 12) THE EYED. TAMPAN* DISTRIBUTION IN THE SUDAN Central and northern Provinces (King 1908,1911,1926). Localities reported in literature and from which I have seen specimens are: Darfur: 15 miles northwest of Fasher (SGC). Nyala (horse; SVS). Wo Locality (BMNH). Kordofan: El Obeid (camel yard; SGC, SVS). “Several local. ities™ (King 1911, Balfour 19118). Blue Nile: Wad Medani (camel yard; SGC, HH). Wad Raiya and Kosti (SGC). Kassala: Karora and Goz Rageb (on human; SGC). Khor Mashi hills south of Tokar, Bir Qui Tiri (under tamarisk tree near well; SCC ). Khartoum: Khartoum (cattle quarantine station; HH). Khartoun, Shambat (H. W. Bedford 1939). “Oasis near Khartoum™ (King 1911, Balfour 1911E). Northern: Dongola (D&nitz 1906). Wadi Halfa and Abu Hamed (camel yards; HH). DISTRIBUTION O. savignyi is distributed locally through arid parts of North, East, ABaeamame Gace the Near East, India, and Ceylon. The Near *In South Africa, called "The Sand Tampan” (Theiler 1952A,B). = Ug so or Middle East probably was its original home. Transportation by caravans, lack of field search and of literature reports, and con. fused identification in Africa have combined to provide a still uncertain picture of this tampan's actual distribution within the noted range. During Neumann's time 0. savignyi was frequently confused with 0. moubata and acceptance of many ie records and of some even more recent reports is questionable. These two species never occur in the same ecological niches. They are close neighbors in some areas, as Somaliland, where 0. moubata inhabits huts next to trees under which 0. savignyi hides. Brumpt (1936) summarized the known geographical distribution and medical relations of 0. savignyi. He noted especially that even though this tampan is Pome found along remote camel trails, it is not known from Morocco in spite of considerable search for it there. Brumpt (loc. cit.) also considered it surprising that 0. savignyi has not been carried to Madagascar but that 0. moubata is common in some areas of that island. Once the very different biology of these two species is understood, a reasonable explana. tion for this distribution pattern may be offered. O. moubata is a highly domesticated parasite that inhabits man's dwellings and frequently hides among his personal effects. It was probably transported from Africa to nearby Madagascar among gear in seagoing vessels. This tick has not been able to survive elsewhere outside of tropical and southern Africa, where it is endemic. O. savignyi,on the other hand, appears to have erratically invaded Africa from the East. It prefers more arid outdoor condi. tions than are found in most parts of Madagascar, and part of its spread probably has been by camels, which are not used in Madagascar. In southern Africa, the range of 0. savignyi appears to be related to environment and wild animals rather than to the comparative recent introduction of camels, movements of domestic stock, or treks of hunters (Theiler, unpublished). In African regions where 0. savignyi does occur, populations are often spotty, = ISeee even in areas that appear favorable. This tampan's predilection for resting outdoors in the soil probably more closely confines its apread to overland routes than does 0. moubata's propensity for hiding in goods or personal effects. NORTH AFRICA: EGYPT (Savigny 1826. Audouin 1827. Koch 1875. Brumpt 1908A. Neumann 1911. Yakoub 1945. Halawani 1946. Davis 1947. Taylor and Hurlbut 1953. Hoogstraal 1954A. Davis and Hoogstraal 1954. Theiler and Hoogstraal 1955. Taylor et al 1955. Hurlbut 1956. Taylor, Work, Hurlbut, and Rizk 1956). TIBYA (Franchini 1927,1929A,E,1932B,1933A,B,C ,D ,1934B,1935A,B,C , 1937,1938&. Zavattari 1930,1932,1933,1934. Tonelli-Rondelli 1930B,1932A,B,D,1935. Gaspare 1933,1934. Garibaldi 1935. Theiler and Hoogstraal 1955). TUNISIA, including Ile de Djerba (Weiss 1911B,1912. Galli-Valerio 1911A. Nicolle, Blaizot, and Conseil 1912,1913A,B. Absence in oases; Langeron 1921. Nicolle and Anderson 1927. Colas-Belcour 1928,1929A,C ,1930,1931). ALGERIA (Chalon 1923. Catanei 1929). / Apparently absent in Morocco (Brumpt 1936).7 WEST AFRICA: NIGERIA (Northern Province only: Simpson 1912A. Alcock 1915. Leeson 1953), FRENCH WEST AFRICA (Brumpt 1936. Kone 1949. Rousselot 1951,1953B). Absent in Gold Coast: Selwyn-Clark, Le Fanu, and Ingram (1923) and Corson and Ingram (1923). Absent in Liberia: Bequaert (1930A).7 CENTRAL AFRICA: No substantiated records. Not known from Belgian Congo, but should be searched for there (Bequaert 1930A, 1931). O. savignyi caecus listed by Schwetz (1927) is a synonym of O. moubata. Waeeee (1953) lists 0. savignyi, probably after Nuttall et al (1908) or Howard (1908) but oe records are most likely repetitions of early misidentifications. 7 EAST AFRICA: SUDAN (Not Koch 1875 or Neumann 1896, “Upper Nubia,” cf. britrea. Donitz 1906. Brumpt 1908A. King 1908,1911, 1926, Neumann 1911. Balfour 1911E,1912. Brumpt 1936, Kirk 1938B,1939. H.W. Bedford 1939. Hoogstraal 1954B. Theiler and Hoogstraal 1955). ETHIOPIA (Nuttall et al 1908. Neumann 1911,1922. Wenyon 1926. Bruns 1937 statements questionable). ERITREA (Koch 1875 - 193 - and Neumann 1896 from River Anseba, “Upper Nubia™, Franchini 192D, E. Tonelli-Rondelli 1930A. Niro 1935. Stella 1938A,1939A,1940). FRENCH SOMALILAND (Neumann 1922. Brumpt 1936. Stella 1932A,1939A, 1940). BRITISH SOMALILAND (Pocock 1900. Drake-Brockman 1913A,B, 1914,1915A,B,1920, biology and medical implications in part mixed with O. moubata. Neumann 1922. Stella 1938A,1939A,1940. Ander— son 1947. Heisch 1950A. Falcone 1952, erroneous disease rela tions). ITALIAN SOMALILAND (Brumpt 1908B. Lees 1914. Paoli 1916. Franchini 1925,1927,1920 ,E,1934. Tonelli-Rondelli 1930A, 1931. Mattei 1933. Niro 1935. Massa 1936A,B, medical implica. tions questionable. Moise 1938,1950. Stella 1938A,1939A,1940. Lipparoni 1951,1954. Giordano 1953). KENYA (The 0. savignyi of Karsch 1878 refers to 0. moubata. Dénitz 1906. Neave 1912. Neumann 1912. Anderson 19244,B.. Mackie 1927. Brassey-Edwards 1932. Lewis 1931A,C,1939A. Heisch 1937,1951A. Walton 1951). UGANDA (Bruce et al 1911. Neave 1912. Mettam 1932). Pena ?TANGANYIKA: ?As 0. morbillosus: Gerstacker (1873). Neumann (1901,1907C ,1910B,1911). Moward (1908) ?quoting Neumann. Absent in Bukoba: Morstatt (1914). The presence of this tick in Tanganyika remains questionable. 7 SOUTHERN AFRICA: / ?ANGOLA: Neumann (1896) listed specimens from Landana that Bequaert (1930A) quite logically believes were QO. moubata. Subsequent statements (Santos Dias 19500) are merely @ repetition of Neumann and disease relations are incorrect. “RHODESIA"; Report of Lounsbury (19000), obviously referring to 0. moubata, quoted by other authors. No subsequent reports of 0. “Savien ?7MOZAMBIQUE: Neumann (1896) ?should be 0. moubata. une (19000 ) should be 0. moubata. Theiler (1943B) states 0. savignyi doubtfully recorded but possibly present. Santos Dias eae evidence not presented. Mozambique records are considered incorrect by Theiler and Robinson (1954) aN NYASALAND (Wilson 1950B). BECHUANALAND (Bedford 1926,1927, 1932B,1934. Theiler and Robinson 1954). SOUTHWEST AFRICA (Louns. bury 19006 , possibly referring to 0. moubata. As O. pavimentosus: Neumann 1901,1911. Donitz 1907C,1910B. Howard 1908. Sess: orff 1914. Bedford 1926,1927,1932B,1934. As 0. moubata: Monnig 1949. - 194 - Leeson 1953. Theiler and Robinson 1954. Theiler and Hoogstraal 1955). UNION OF SOUTH AFRICA /“Lounsbury 1899C,1900B,C (confused with O. moubata), 1903B,1904A. Howard 1908. Donitz 1910B. Bed ford 1920,1926,1927,1932B,1934. Alexander 1931. Bedford and Graf 1934,1939. R. du Toit 1942B,C,1947A,B. Theiler and Robin son 1954. Theiler and Hoogstraal 1955.7 NEAR EAST; ADEN (Nuttall et al 1908. Patton and Cragg 1913. Cunliffe 1922. Hoogstraal ms.). YEMEN (Mourit 1953. Hoogstraal, ms.). PALESTINE (Theodor 1932. Smith 1936 quoted by Brumpt 1936. Bodenheimer 1937). IRAQ (Leeson 1953). MIDDLE EAST: INDIA (Christophers 1906. Neumann 1911. Patton and Cragg 1913. Donovan 1913. Fletcher 1916. Cross and Patel 1922. Rao and Ayyar 1931. Sen 1938. Sharif 1938. Kapur 1940. Joshi 1943). CEYLON (Nuttall et al 1908. Brumpt 1936. Crawford 1937. Chow, Thevasagayam, and Tharumarajah 1954). HOSTS Camels are most frequently mentioned as hosts. O. savignyi appears to be present in most areas in which dromedaries are used. Fowls are sometimes attacked and all domestic animals may serve as hosts (Lounsbury 1900B). For instance, this tampan is common in cattle yards at Mawar, India (Joshi 1943) and under trees where mules are tethered in Somaliland (Lipparoni 1951). Human beings are frequently bitten, especially when they sleep in camel yards or sit under trees commonly used by domestic animals for shade. Any laboratory animal may serve as a host. Dogs are satisfactory laboratory hosts (Lounsbury 1904A). Game animals are said to be attacked, but evidence is scant. The rhinoceros, lion, and buffalo may serve as hosts in Kenya according to Walton (1951). Neumann (1912) reported numerous specimens from a Kenya locality where a giraffe had been standing. BIOLOGY Life Cycle Eggs of 0. savignyi are deposited in sandy soil where adults hide. individusl Tenctss, observed by Cunliffe (1922) at 30°C., - 195 ~ laid from 100 to 417 eggs, averaging 219. Other individuals that had fed four times laid five egg batches totalling about 900 eggs over a thirteen month period (Patton and Cragg 1913). The embryonic development, described by Christophers (1906), is essentially like that of O. moubata. Larvae, like those of 0. moubata, are nonmotile and do not feed. Although some undergo ecdysis in the egg, most larvae free themselves from the eggshell before molting to nymphs (Cunliffe 1922). After splitting the eggshell, larvae molt to nymphs in five hours (Davis 1947) to ten hours (Patton and Cragg 1913). Four nymphal instars over a period of about 84 days and seven nymphal feedings were observed by Patton and Cragg (1913). Cun. liffe (1922), on the other hand, noted that males appeared after four to six molts and females usually after the sixth molt. Rea sons for differences in number of instars among argasids remain to be ascertained. The very active nymphs commence feeding two or three days after molting and require fifteen to thirty minutes to reach re— pletion. Adults normally feed for similar periods but the pre. sence of both stages along remote camel trails would indicate that on occasion some tampans may remain longer on the host. Females in Cunliffets (1922) studies lived between 292 and 420 days at 30°. and for an average of 775 days at 22°, Spermatophores superficially similar to those of 0. moubata are utilized by this species (in Egypt), although Christophers (1906) and Nuttall and Merriman (1911) questioned their presence. Other details of the life cycle have been reported by the above mentioned observers and by Rousselot (1953B). However, more extensive and refined studies are still necessary. Ecology 0. ae among the tick species studied by Lees (1947), shows the greatest ability to limit water loss at high temperatures. The critical level for this species is 75%. while for 0. moubata = 196.= it is 63°C. Even xerophilic hyalommas abruptly increase water loss at 45°. (cf. page 154). This factor explains in part how the eyed tampan can exist in deserts where little other life is sustained. Cunliffe's (1922) studies, from which he concluded that the temperature and humidity requirements of both these tampans are much the same, should be repeated with special at— tention to extreme levels. Famous for the viciousness of its attack, this tampan is usually well known wherever it occurs. Natives quickly lead one to infested animal corrals, trees under which man and beasts rest, and well sides where the eyed tampan is superficially burrowed awaiting its prey. In Somaliland, Lipparoni (1951) reported, Oo. ees is common under trees where soldiers tether their mules O. moubata infests huts beside these trees. Although the adventitious presence of 0. savignyi in buildings must be expected, early records from human Hata appear to be based on misidentification. For instance, Drake-Brockman's various reports of 0. savignyi in British Somaliland buildings have been questioned by Taicreon (1947). Anderson found 0. savignyi ex. clusively outdoors in the same area, and 0. moubata, previously thought to be nonexistant in the area, exceedingly common in huts and coffee houses. We have never observed 0. savignyi in sites directly exposed to the sun. Indeed, at the Khartoum quarantine one may see a long, seething line of thousands of hungry tampans helplessly confined to the shade of a row of acacia trees. A few yards away, separated only by the hot, nine o'clock sun, newly arrived cattle tied to a post fence tempt the tampans to cross the glar— ing strip. The next morning, in the coolness of seven o'clock, those tampans under the trees are all blood bloated and resting comfortably in the sand, others are dragging back from their hosts across the now nonexistant barrier, and the legs of the cattle are beaded with yet other podshaped ticks taking their fill of blood in a regular line just above the hoof. Laboratory rats and mice, as noted by Heisch (1950A), assail this tampan. Rats feast on nymphs and adults. Mice commonly assault nymphs, but only particularly bold mice attack adults. These rodents, in turn, facilely escape bloodthirsty but lumber— ing adults, although small, active nymphs more easily attack them. Predators in nature do not appear to have been reported. - 197 - REMAR KS Christophers! (1906) extensive study on morphology and diges— tion of 0. savignyi has been reviewed in the section on 0. moubata, which also contains a number of other data pertaining to bo species. The nymphal instars may be approximately determined accordi to Cunliffe's (1922) data, also presented by Campana-Rouget (1954). The haller's organ of QO. savignyi has been described and il. lustrated (Schulze 1941). Jakob (1924) used this tampan to illustrate certain theories separating the ixodids from the argasids on the basis of differences in external grooves, ridges, and prominences. He did not believe, as a result of these studies, that Argas developed from Ornithodoros, but rather that both genera had a common origin in the Uropodidae, a member of which, Discopoma africana Vitzhum, was illustrated. DISEASE RELATIONS MAN. The bite of 0. savignyi may have severely painful seque— lae but this tampan has never been found infected with pathogenic organisms in nature, and transmission of pathogens has not been demonstrated until recently. Even the earlier assertions that the eyed tampan transmits human relapsing fever (Borrelia spp.) have been cast into considerable doubt by subsequent research. DOMESTIC ANIMALS. Camels and cattle suffer greatly and may even be killed by the volume of blood lost to numbers of eyed tampans in their pens. EXPERIMENTAL. Leishmania donovani, which causes kala azar in human beings, does not develop in 0. savignyi. Trypanosoma cruzi undergoes development in this tick in the laboratory. I. evansi cannot be transmitted from the tick to animals except by inocula. tion of a suspension of infected ticks. Experiments on transmission of heartwater (Rickettsia ruminan. tium) of cattle by means of O. savignyi have been unsuccessful. - 198. West Nile virus remains viable for at least three months in o. SoS » and transmission of the virus to mice by the bite of parenter jminfected ticks has been demonstrated. Similarly, specimens experimentally infected with Sindbis virus transmit the organism when biting. Spirochetosis of chickens (Borrelia anserina) is not trans mitted by this tampan. IDENTIFICATION See remarks under 0. moubata, pages 189 and 190. - 199- FAMILY I XODIDAE INTRODUCTION All tick genera, save those in the family Argasidae and Nuttalliella, in the family Nuttalliellidae*, fall into the family Ixodidae and are referred to as “hard ticks™ or “ixodids™. The use of the term ixodid is not confined to the genus Ixodes. In Sudani Arabic, hard ticks are called "gurad” ( 3/-> All ixodid genera that normally inhabit Africa also occur in the Sudan with the exception of Rhipicentor**, This genus is represented by R. gladiger (Neumann, = bicornis Nuttall and Warburton, ode} in neighboring Belgian Congo (Bequaert 1931) and further south in Africa. R. nuttalli Cooper and Robinson, 1908, occurs on various animals in South Africa and in Southwest Africa. *The family Nuttalliellidae (Schulze 1935) contains only a single, exceedingly rare species, Nuttalliella pomeges Bedford, cain described from Little Namaqualand, Southwes rica. This family appears to be a "missing link". A scutal outline is present but not structurally differentiated from the léathery, papillated inte gument. Though the mouthparts are anterior as in Ixodidae, the palpal segments are movable as in Argasidae, but an inner groove on the second segment is suggestive of the reduction in Ixodidae, in which the terminal segment is essentially merely a small ap. pendage of the penultimate segment. The biology of this strange species is unknown. **Rhipicephalus (Pter — Sune Neumann, 1913, a remarkable aberrant parasite o est Africa, is frequently treated as a monotypic genus and a aettnee as a subgenus of Rhipicephalus. See Neumann (1913) for description of male and Colas-Belcour (1932) for description of female, nymph, larva, hosts, biology, and disease relations. = 200 = In addition to African genera discussed herein, several others have been described from other continents, chiefly by Schulze. Many of these genera, however, meet with little general acceptance among contemporary specialists. A useful summary, including Schulze's and others genera, may be found in Baker and Wharton (1952). In general, however, "textbook genera”, as used by New mann and by Nuttall and Warburton, usually suffice and are of considerable practical and scientific value. They should be little tampered with if at all. Special groups of ticks may be readily categorized at the subgeneric level to provide a useful sounding board for their acceptance by specialists. Ixodid ticks occur throughout the world wherever terrestrial vertebrates are found. They attack most land mammals and some marine forms. Some ixodids parasitize birds and reptiles, some feed on amphibians. Adults of few if any ixodids regard man as a host of predilection but many attack him in the absence of other available hosts. Larvae and nymphs are much more common and se— rious pests of man than adults. Morphologically, these ticks differ from argasids by the presence in all stages of a dorsal scutum. Ixodid mouthparts are always anterior and visible from above; the body is oval. Larvae have six legs; nymphs have eight legs and a female type scutum but lack both porose areas on the basis capituli and the genital aperture of the female. The female scutum covers only the anterior portion of the dorsum, the male scutum extends to the posterior margin of the body. Eyes may be present or ab sent. Nymphs and adults have a spiracular plate situated lateral. ly, posterior of each hind leg. Palpi, clearly segmented and movable in argasids, are limited in ixodids to segments 2 and 3 and to a more or less well developed segment 1; segment 4 is inserted ventrally into a pit of segment 3. Biologically, the majority of ixodids display comparative uniformity within rather narrow limits. Exceptions are notorious but few. The number of species is small, somewhere around five hundred. Ixodids are highly adapted to parasitic life but one is prone to wonder how they have succeeded in the battle for survival. Their aimless wandering habit and dropping from hosts and their indiscriminate oviposition proves fatal to many indi- =) 20 viduals, even though they survive long periods without food. Their extended life cycle subjects them to many adversities depending on season and the peregrinations of their hosts. The requirement of two or more kinds of hosts, often with divergent habits, limits their distribution to certain faunal areas. The comparatively. large size of females makes them subject to injury and tempting food for predatory arthropods and vertebrates. Certain pathogenic organisms, fungi, and hymenopterous parasites kill them. Ticks have little protection against an enemy except their ease of con cealment. They are particularly susceptible to attack during the lethargic premolting and the weak postmolting period. On the other hand, many biological features enable ticks to survive especially well. They lay numerous eggs and withstand a comparatively wide temperature and humidity range with greater ease than many other arthropods. They survive for months or years without food and often gain considerable protection from the con. cealed places in which they feed on the host. They frequently are offered a wide choice of appropriate hosts. Should they annoy the host, the animal is usually powerless to rid itself of the para site. Ticks! slow metabolic rate has certain advantages and the leathery, usually inconspicuous, integument offers some protection from living enemies, water, and chemicals. Parthenogenesis, pos— sibly a common occurrence though not well studied, may aid sure vival. The genus Ixodes, as a biological unit, shows much divergence from the usual patterns of ixodids. The biology of this genus must be considered independently but in relation to other genera. Review summaries frequently leave the student with the impression that the unique biological characters of Ixodes ticks are also characteristic of other ixodid genera. The degree of host specificity in ixodid ticks varies fron genus to genus or within certain subgroups of various genera. Generalizations on this point should be very carefully qualified. Eggs are laid only once; promiscuously; at one time and place; and are always numerous, sometimes numbering over 10,000. Eggs hatch in from two weeks to several months, depending mostly on climatic factors. = On Larvae are active and sometimes are easily visible as hun. dreds or thousands of them rest on grass awaiting a host. Larvae often parasitize small mammals, birds, and reptiles, sometimes in their nests or dens. Only a few kinds choose larger animals for hosts. Thickness of the host skin is possibly an important fac- tor in restricting larvae to smaller animals. Larvae seldom feed within a week after hatching. After some days of feeding, larvae molt to nymphs on the same host (two-host type of life cycle), or drop from the host and some time later molt to nymphs (three-host type of life cycle) on the ground. Exceptions are the single-host boophilid ticks (and a few others) that molt and remain on a single host during their lifetime. Nymphs that have molted on the ground seek a new host, sometimes the same type they fed on as larvae but, more commonly, animals similar to hosts preferred by adults. After several days of feeding, nymphs drop to the ground; be- come quiescent for a time, then molt to adults. There is only a single instar in the nymphal stage (see footnote, page UA) Some species may undergo one or another of these variations in life cycle, depending on climatic, seasonal, or nutritional fac tors, most of which are still poorly evaluated. There is evidence that in certain three-host species (in which larvae characteristically feed on small mammals) larvae that feed on a somewhat larger animal, such as a hare, may re- main on the host, molt to nymphs, and continue feeding. They drop from the larval_-nymphal host only before molting to the adult stage. Nymphs and females become tremendously engorged and new cuticle is developed during the rather slow feeding process in order to accommodate the huge volume of blood ingested. Males become only slightly distended if at all, apparently, as in argasid ticks, by stretching the integument although they may remain in a feeding position for months, even after the slaugh. tered host's skin has been removed. As reservoirs of a great variety of pathogenic organisms, ixodids are pre-eminently important, whether they act as vectors or not. As vectors, they transmit a greater variety of organisms o 20g — than any other arthropod. This variety is said to be greater on continental Africa than anywhere else in the world. Other injury, apparently due to toxins, in the form of tick paralysis may be locally important. Death, lameness, or serious debilitation of the ost by exsanguination or as the result of secondary infec tions at the site of attachment is not uncommon. Economic loss due to numerous punctures of animal hides by the mouthparts of large ticks is frequently reported. = 20/4 AMBLYOMMA INTRODUCTION Of about a hundred Amblyomma species in the world, some twenty occur in Africa and eight in the Sudan. The specific identity of most common African species appears to be settled and only in ex ceptional instances are specimens likely to be confused. One of the chief remaining taxonomic problems among common African amblyom mas is the A. marmoreum group, in which the range of species varia. tion needs to be determined for several somewhat differing forms. Observations from the present study indicate need for further re search on the relationship between A. variegatum and A. pomposun and suggest that the latter may be no more than a subspecies o the former. Recently a few workers have designated certain African populations by subspecific ranks that challenge further investiga. tion. Several West African species are known from so few specimens that their validity is questionable. This genus has been the subject of an extensive review by Robinson (1926) comprising volume four of Nuttall's Monograph on Ticks. The African species have been keyed by Rageau and Ver. vent (1953). The immature stages of most African amblyommas remain to be described with satisfactory criteria for distinguishing them. Economically, two African species have thus far been shown to harbor or transmit human disease organisms. A. hebraeum is considered an important boutonneuse fever ("tick typhus") vector in South Africa and A. variegatun has been found naturally im fected with Q fever in = Equatorial Africa near the Sudan border. Several species are important transmitters of veterinary diseases, cause damage to animal skins, or debilitate animals through the volume of blood withdrawn or by initiating wounds that develop into ugly secondary sores. Biologically, many gaps exist in our knowledge of African Amblyomma distribution, host-preferences, especially of immature stages, and life history. Birds are important immature-stage = 205 — hosts but the full extent of their importance as hosts remains to be studied. Nymphal preferences may differ from those of adults, although in some species this may not be true. Host size appears to be of some importance, for most larvae parasitize small animals; nymphs attack larger animals; and adults feed on the largest available animals, except carnivores. Immature stages, however, more frequently parasitize carnivores. Cattle and, to some extent, other domestic animals are important adult hosts and wild ante. lopes are also frequently parasitized. The African tortoise, rhinoceros, elephant, and buffalo harbor species mostly restricted to themselves. In general, the domestic animal parasitizing spe— cies are well represented in study collections; others are seldom collected. ' Amblyomma ticks are usually three-host parasites and, so far as known in Africa, there is usually only a single generation am nually. KEY TO SUDAN SPECIES OF AMBLYOIIMA MALES 1. Lateral grooves extending anteriorly at least to middle of SCUCUM sre wisie o'cie.eieciolere oieieloiere cieleisie eielercieverereiee Lateral grooves ADSEN Gereleleicisielsicicicieleielole cveleveleleielelelelercielevelelereleteiat/| 2. Eyes small, hemispherical, situated in a welll defined depression. (1.6%, Orbitbedys.cer sisis.cersicwic visielcee> Eyes flat or very slightly convex, not in a GEDr ESS ONicjeceicielsieleloloicicielcieleleeieie eielslcicicie ciele ejele eieieleleleieisielo') 3. Festoons with two colors. (Scutal or- namentation always as illustrated).......esseeseeeA. LEPIDIM Figures 6S and 69 Festoons only dark CONGR ECerecieleleieiololoielele eieiere sicreleieleielelercieteleletere se 2206 he De Te Scutum with numerous coarse punctations, and with a red lateral spot. (Rare in Sudan). ccocescccscccccccccccccccccccccccecccccccehs POMPOSIM Figures Scutum smooth, with few scattered coarse punctations or none; usually without red lateral spot. (Common in Sudan)...........2..eeA. VARIEGATUM Figures Scutum smooth, with only very fine punctations. Eyes slightly convex. (Ghiefisy, Prom bullalo,)s ..ccisse o:0.010.0:<0 006 sieie.60000 ech, COHAERENS Figures Scutum with scattered large punctations. Eyes flat. (Chiefly from cold-blooded MGISUOAGOS ats clesints,g1elvVele aie 6/10 eis:e'e cle caiaiate/a’cveieieiee'e Dew sid cates eeO Smaller than A. marmoreum group, maxi- mam size 5.5 mm. x 4.5 mm. Dark scutal areas more widely separated by pale areas than in A. marmoreum (see Figures )....e.+-eA. NUTTALLI Figures 76 and 77 Size at least 6.0 m. x 5.0 m. Dark scutal areas less widely separated from each other by pale areas than in A. MOULALID occwciciccvcvecescceseissisceicccesecececeschs MARMOREUM Figures Scutum dark, ornamented areas small, punctations only small. (Medium size, drab tick, chiefly from elephants)......eseeeeeeeA. THOLLONI Figures 88 and 89 Scutum extensively ornamented, some large punctations present. (Very large, colorful tick, chiefly from PIT CEFOS|) 6 o/aeieieis foe oi cjainieisee wis via\sieicie)ain,c/e)e csietioy RHUNOCEROLLS Figures ol = 207 . 1. he FEMALES Eyes in a well-defined depression (i.e. orbited), hemispherical or strongly CONVEX alerclole oleleleioieiclelelerelcielereleielelolerelelelelaielevalereleiaeleleisveles Eyes not in a depression, flat or slightly GCONVGXeretcm ieee w cieleietalelciele sieleleieie ele claveleleieloleicicicieteleveieiele e2) Scutal punctations very coarse, uneven, some confluent; surface rugose poste. riorly; ornamentation absent or con. sisting of only a small pale area in posterior field; length often no greater than width. Eyes may be convex but not hemispherical. (Ex tremely rare in Sudan) so ceseccdeceeeas oosv sce ccrccchs OMPOSUM Figures an Scutal punctations not so coarse, sur. face not rugose, pale areas more ex. tensive (unless faded). Eyes hemis. phemicale “(Gommon')\.%%. co occ 6 ce restates ca eee ete ceed Scutum narrowly rounded posteriorly; large punctations chiefly in medio. lateral area, more or less confluent..............A. IEPIDUM Figures 70 and 71 Scutum more widely rounded posteriorly, large punctations generally distributed but nonconfluent.secccccccccccccccvcccccceesesehs VARIEGATUM Figures 9 and 95 Scutum with central and lateral areas largely pale colored; punctations either large or TENG eis syoie a oie a 6 evereve te oreo latalorate oceles 6/3 616 c's ore elelsieversalereierce Scutum with either central or lateral areas largely dark colored; punctations only ESIGN ote yer otekereteve ev eve) ole veie eo eletolelevoreteioie ei 6c vevevsverevelere(orsierelelerevelercten/: S620 = 5. Scutum with some large punctations scattered over entire surface; length usually equalling or greater tlenwil GielteueyOSh tal cts revels clele’ aveieierelelelaleieieielcieiee eicietee sie sisters oO Scutum with few large punctations only on anterior half; wider than long, mostly pale (reddish brown). Eyes slightly convex. (Very large tick, chiefly from rhinoceros).....eeeeeeeeeeA. RHINOCEROTIS Figures Scutum with extensive pale median area that is broadly rounded poste riorly. (Large tick, chiefly from TOT LOUGS)) mealeialcleleleie’s s/s fe ols clsielcivicialeis e'e\c.\eielsic’s see ahem MARMOR HUN Figures 7% and 75 Scutum with less extensive pale median area that narrows to a point poste— riorly. (Medium size tick, chiefly from tortoise and leguan lizards ).....ee.se-eseeeeAs NUTTALLI Figures 7 Scutum dark centrally, small pale spot on posterior border, lateral fields with small pale spots or entirely dark. Eyes flat. Very narrow pale rings on legs. (Chiefly from elephant )..........A. THOLLONI Figures 90 and 91 Scutum pale centrally, lateral fields with only small pale spots. Eyes slightly convex. Broad pale rings on Hess. (Chiefly from buffalo ).c loss. cee ces AS COHAERENS Figures *In the Sudan, exceptional specimens of A. marmoreum may have a scutal width of at least 1.3 greater than length (Fic¢ure 74), but in these, large punctations are scattered over the entire scutal surface. = X08) & Figures 64 and 65, oc, dorsal and ventral views, Sudan specimen Figures 66 and 67, 9, dorsal and ventral views, Uganda specimen AMBLYOMMA COHAERENS Note: The o illustrated is malformed in the following areas: left scutal margin anteriorly and right scutal margin lateral. ly deformed, festoons on right side absent, only three legs on left side. Diagnostic criteria are not affected by these malformations. PLATE XXIV = ALO) os AMBLYOMMA COHAERENS Donitz, 1909. (Figures 64 to 67) THE EAST AFRICAN BUFFALO TICK LynN ORES. EQUATORIA PROVINCE RECORDS 34 Boma Plains Syncerus caffer aequinoctialis Dec #1 Obbo Grammomys macmillani erythro us Mar Birds #1 Obbo Turdus libonyanus centralis Mar *] Torit Francolinus clappertoni gedgii Dec #7 Tkoto Numida meleagris major Dec The Boma Plains specimens constitute the first and only defi nite record of A. cohaerens from the Sudan. If the tentative im. mature stage identifications are correct, we may expect that A. cohaerens occurs in other parts of Equatoria Province, especially where buffalos roam. The East African buffalo tick has not been found in Bahr F1 Ghazal or in Upper Nile Provinces, although buffalos are present in these Provinces and frequently have been examined for ticks by H. H. King, Sudan Veterinary Service per— sonnel, and the writer. DISTRIBUTION A. cohaerens is an Fast and Central African buffalo parasite that reaches the northern limit of its range in Equatoria Prov- ince and extends southwards into Tanganyika. *Tentative identification by Dr. G. Theiler. ee res CENTRAL AFRICA: /?CAMEROONS: Noted by Schulze (1941) with. out data. Recorded on (?imported) cattle at Yaounde by Rousselot (1951) but not repeated by him (1953B). Rageau (1951,1953A,B), apparently quoting Rousselot. 7 FRENCH EQUATORIAL AFRICA (Rousse— lot 1951,1953B). * BELGIAN CONGO and RUANDA-URUNDI (Nuttall and Warburton 1916. Robinson 1926, Schwetz 1927B,C,1932. Bequaert 1930A,B,1931. Rousselot 1951,1953B. As A. cohaerens and also as A. hebraeum: Schoenaers 1951A,B; see REMARKS below. Theiler and Robinson 1954. Van Vaerenbergh 1954). EAST AFRICA: “EAST AFRICA" (Dénitz 1909). SUDAN (Hoogstraal 1954B). FRENCH SOMALILAND (Hoogstraal 1953D). KENYA (Robinson 1926. Lewis 1934. Weber 1948). UGANDA (Neave 1912. Robinson 1926. Tonelli-Rondelli 1930A. Richardson 1930. Mettam 1932. Mettam and Carmichael 1936. Wilson 1948A, 19500. Hoogstraal 1954C). TANGANYIKA (As A. anceps: D8nitz 1909. Hoogstraal 195/C). a {/ NOTE: The ANGOLA record by Sousa Dias (1950) is actually A. astrion, not_A. cohaerens (Theiler, correspondence). See REMARKS below. 7 ~= HOSTS All authors list the African buffalo, Syncerus caffer, as the chief host of A. cohaerens. The East African buffalo tick is frequently reported to attack domestic cattle in areas where buffaloes are common or after large numbers of these animals have been shot out for disease control. Domestic cattle have been listed as hosts by Robinson (1926), Schwetz (1932), Schoenaers (1951B), and Rousselot (1951,1953B). Cattle are frequently parasitized when near game, especially buffalo (Wilson 1948A,1950C). In SAW = certain Uganda areas, A. cohaerens, together with A. variegatun, is the predominant cattle tick on old buffalo Ean See (Richardson 1930). It would be interesting to know how long the buffalo tick remains in an area as a cattle parasite after its chief host has been exterminated. Other animals are rare hosts of adults. Elephants (Robinson 1926). Tortoise, and domestic sheep and goats (Mettam 1932). Eland (lewis 1934, Weber 1948). Warthog (Theiler, unpublished). Black rhinoceros (Hoogstraal 1954C. Also &t removed from a Tanganyika rhinoceros skin by a taxidermist ~ CNHM collections). Hosts of nymphs (tentatively identified as this species) are ground birds and a tree rat (Equatoria Province records above). Dozens of Thammomys tree rats have been examined by me in the Sudan and Kenya without finding other ticks on them. Several nymphs have been taken from a warthog near Lake Edward, Uganda, by Lt. Col. Don Davis, U.S.A. (HH collection). BIOLOGY Unstudied. In Ruanda-Urundi, A. cohaerens occurs as high as 2200 meters altitude (Schoenaers 19515, as Av hebraeum). DISEASE RELATIONS Unstudied. REMARKS Sections of the mouthparts of A. cohaerens have been illus— trated by Schulze (1936A), who also illustrated the form of haller's organ (1941), and (1950A) discussed the dentition of this species. Schulze also (1932c ) utilized this tick to illustrate his concept of the relation of ornamental design to location of muscle attach. ments. A number of other remarks and illustrations concerning the exoskeleton of this species are presented in the same study. ep Ae es The middle festoon of a few males may be largely pale and other festoons may be equally so; such specimens would key to A. hebraeum. There are no excessively pale festooned individuals in our own collection or in that of British Museum (Natural His. tory), but some material in the Onderstepoort collections shows this variation (Theiler, correspondence). Such a specimen, seen in the Rocky Mountain Laboratory collection, was the cause of mis. identification of A. cohaerens as A. hebraeum in Schoenaers? (1951B) list. A. hebraeum, though colorful, never shows as much iridescence as most other ticks of this genus (Theiler, corres. pondence). Should specimens resembling A. cohaerens be found on the west bank in Equatoria Province, they Should be checked against A. splendidum Giebel, 1877, of the Congo and West Africa (cf. Robin. Son 1926, pp. 123.125). A. splendidum males are somewhat larger, have a vermillion-red spot in the center of the scutum, and never have a falciform stripe. Females are indistinguishable from those of A. cohaerens although they are often a little larger. A. cohaerens also closely resembles A. astrion of West Africa. Sousa Dias (1950) confused A. astrion with A. cohaerens. Recent studies by Theiler indicate the distinctness of the two species. Since A. astrion is unlikely to be found in the Sudan tick fauna, it is unnecessary to mention further detail. However, students who may compare our nomenclature with that of Sousa Dias should be aware that this discrepancy exists. IDENT IFICATION A. cohaerens is easily recognized within the known Sudan tick fauna. Closely related species, some of which may occur in the Sudan, are mentioned above. Males fall into a group in which the eye is not in a dew pression, although it may be very slightly convex; festoons are mixed dark and pale, scutal punctations are only fine, scutal ornamentation is as illustrated (Figure 64) but either with or without a falciform stripe; lateral grooves reach nearly to the eyes. This is a medium size tick, from 5.0 m. to 6.0 m. long and from 4.0 mm. to 4.7 mm. wide. - 214- Females have a triangular scutum with only fine punctations and with an extensive pale central area; the lateral scutal areas are dark except for one or two very small light marginal spots. The eyes are not in a depression though they may be very slightly convex. Leg segments are ringed by broad bands. Females are of medium size, approximately 5.0 mm long and 4.0 mm. wide. The scutum is approximately 2.8 mm. long and 2.9 m. wide. = ils) — Figures 68 and 69, o, dorsal and ventral views Figures 70 and 71, 9, dorsal and ventral views AMBLYOMMA LEPIDUM Sudan specimens PLATE XXV = 2167= 11 NP NMHP Ot =) Pa wJ fe Og al TON Wo tS ee ng AMBLYOMMA IEPIDUM Dénitz, 1909. (Figures 69 to 71) THE EAST AFRICAN BONT TICK. EQUATORIA PROVINCE RECORDS Boma Plains Jebel Kathangor Loronyo Taurotragus oryx pattersonianus Torit Hippotragus equinus bakeri Lafon Hippotrazus equinus bakeri Torit Wcdla cies buselaphus roosevelti Terakeka Damaliscus korrigum tian Opari Felis libyca ugandae Kapoeta domestic cattle Kapoeta domestic cattle Kapoeta domestic cattle Llolepori domestic cattle Tliu domestic cattle loronyo domestic cattle Torit domestic cattle Torit domestic cattle Torit domestic cattle Torit domestic cattle Torit domestic cattle Katire domestic cattle Gilo domestic cattle Kapoeta domestic goats Loronyo domestic goats Kapoeta domestic sheep Kapoeta domestic sheep Loronyo domestic dog Torit domestic dog Kapoeta in rest house Bird Torit Neotis cafra denhami SAS Syncerus caffer aequinoctialis Dec Sones pact Gazella granti brig ual Dec Jan Mar Dec DISTRIBUTION IN THE SUDAN A. lepidum occurs in all Provinces except Northern, though it does arrive at the Wadi Halfa quarantine in Northern Province on cattle from the South (King 1926). All available Equatoria Province specimens originate from Eastern District and Torit District with the single exception of a male from a tiang at Terakeka (H. H. King legit), on the west bank of the Nile. The following Sudan locality records are all from cattle and all from the Sudan Government collection unless otherwise noted: Bahr el Ghazal: Lau and Yirol (SVS), Guar, Gogrial Sub. district (giraffe; SVS), Aliab (buffalo; SVS). Akot (dying bull; SVS). Eight miles west of Yirol (head of greater bustard; SVS). I doubt that A. lepidun is widely established, if at all, in Bahr el Ghazal Province. ach collection consists of only a sanete male, except for one male and female from great herds of migrating cattle at Yirol, 22 April 1954 (SVS). r Nile: Pibor Post. Akobo. Maban (domestic cattle and coats WE y. pariak (SVS). Bor (SVS). Melut (mle). Rom (buffalo). Kaka (roan antelope). Er Renk (domestic sheep). Makier (SVS). Malakal (HH). Specimens from Tonga, that were identified as A. lepidum by D&nitz in 1912, were the basis of King's (1911) Sugar? of A. ae ee from the Sudan according to Nuttall's notes for Lot s logbook in British Museum (Natural History). Blue Nile: Hosh. Tibna. Roseires. Wad el Nail. Singa (camel). Wad Medani (domestic cattle, miles, and camel; SCC, HH. One & feeding between toes of man, August 1954; Bisa El Minesi legit). Abu Hashim (camel). Sennar (camel). Lake Ras Amer (camel). Abu Zor. Hassa Heissa (camel; G. B. Thompson, correspondence). Sennar area (cheetah; Robinson 1926). Kosti (Gordon College collection). Kassala: Kassala (SVS). Darfur: Radom (SVS). = 2icie Kordofan: Umm Berembeita (SCC). {Northern and Khartoum: Specimens from cattle at the Wadi Halfa Quarantine station, from Ethiopian cattle at Khartoum, and from "A.0.F." native horses /French West (?Equatorial) Africa 7 (at Khartoum) are also present in Sudan Government collections. 7 See BIOLOGY below for further remarks on distribution in the Sudan, DISTRIBUTION A. lepidum is an East African herbivore parasite and is not known to occur elsewhere. It becomes uncommon in Tanganyika but is more common locally northwards to the semidesert belt of the Sudan. EAST AFRICA: "EAST AFRICA" (Doénitz 1909). SUDAN (As A. hebraeum variegatum: King 1911. King 1926, Robinson 1926. icopstraal TOSUL-TS51B). ETHIOPIA (Stella 1940). ERITREA (Franchini 192M. Tonelli- Rondelli 1932. Niro 1935. Stella 1940). ITALIAN SOMALILAND (Paoli 1916. Tonelli-Rondelli 1926A. Franchini 1926A,1927, 1920 ,E. Niro 1935. Stella 1938A,1940. See also adult host records below). Note: Numerous reports of A. hebraeum from former Italian East African possessions probably refer in part to A. lepidum and in part to A. gemma, KENYA (Robinson 1926. Lewis 1931C,1939A. Dick and Lewis 1947). UGANDA (Mettam 1932. Lewis 1939A. Wilson 1948A,1950, 1953). TANGANYIKA (Evans 1935. Cornell 1936. Lewis 1939A. J. B. Walker; small numbers; unpublished; see HOSTS below). OUTLYING ISLANDS: ZANZIBAR (Donitz 1909. Robinson 1926). IMPORTED SPECIMENS: Cairo, EGYPT (Donitz 1909). A. cele still arrives almost daily at the Cairo slaughterhouse on an cattle but has not established itself in Egypt (Mason 1915, Hoogstraal 1952A). Almost every specimen is a male. Numerous a) oo males are also not infrequently found on traders! camels reaching the environs of Cairo from the Sudan, This species has been taken from imported cattle at East London, South Africa (Robinson 1926), but is definitely not es. tablished in the UNION OF SOUTH AFRICA (Theiler, correspondence). PALESTINE records (Bodenheimer 1937) probably also represent imported specimens. HOSTS A. lepidum is chiefly a cattle parasite with smaller domes— tic animals a scattering of wild herbivores as second choice. Large ground birds and carnivores are rarely attacked. A single adult has been taken feeding on man. Nymphs have been found on antelopes, bustards, and domestic cattle and dogs. The host pre— dilection of immature stages is still poorly known. Domestic animals: All investigators listed above refer to cattle as hosts of this tick. Sudan records are almost the only ones available for other domestic animals. These include camels, horses, mules, goats, sheep, and dogs. Individual camels on oc- casion are heavily infested. Although camels appear to be rather important hosts in central Sudan, local camels in the north are not known to harbor this tick. Evans (1935) listed sheep and dogs as hosts in Tanganyika. Wild animals: Buffalo (Robinson 1926, King 1926, Wilson 19500, various Sudan records above). Rhinoceros (Wilson 1950). Grant's gazelle and hartebeest (Wilson 19500, Sudan records above). Roan antelope eland, giraffe, tiang, wildcat, and preat. er bustard (various Sudan records above). Burchell's zebra and spotted hyena (Theiler, unpublished records), Cheetah (Robinson 1926). Ostrich (1c and lo in Hoogstraal collection, from "west of Afmadu", Somalia, 1952, Col. D. Davis legit). Of 49 Thom son's gazelles examined in Tanganyika, only two yielded eight males and one female A. lepidum; none were found on numerous other game animals eae there (J. B. Walker, unpublished). Man; Blue Nile Province record above. <7 C20) — Nymphal hosts: The Sudan hosts appear to be the only ones recorded for nymphs. These are Grant's gazelle, Roosevelt's hartebeest, rarely domestic cattle and dogs, and greater bustard. Nymphs were identified by Dr. G. Theiler. BIOLOGY A. lepidum is common in many of the semiarid regions of East Africa. it inhabits economically marginal areas and is not known to be a vector of animal or human disease pathogens. Therefore, not even the industrious veterinarians (in Africa they do most of the legwork on ticks for zoologists and for medical research. ers) have investigated this tick's life history. It would, however, be surprising indeed to find that the life cycle were any other than the three-host type. J. B. Walker reports (cor- respondence) that larvae fed on a rabbit did not molt but that a few larvae in her laboratory have engorged on pigeons. In his interesting and important ecological survey of cer- tain tick vectors of East and Central Africa, Wilson (1953) has noted the common occurrence of A. lepidum in the Karamoja area, the driest district of Uganda. “Wilson includes this species in his discussion of the R. pravus — A. gemma association, reviewed herein on page (cf. also A. variegatum, page 681). L These _ sections should be consulted to obtain a better impression of what is now known of the ecology of A. lepidum. Other species associated with it in Karamoja are Re ©. evertsi, H. rufipes, and H. truncatun. For additional background, the following biological informa. tion, as written in the manuscript before Wilson's (1953) paper became available, remains of value. Lewis (1939A) considered A. lepidum to be a desert species but, on the contrary, it appears to prefer more arid savannah country and, to some limited extent, semidesert regions. In Uganda it is most common in dry thorn country (Wilson 195). Although Sudan cattle bound for the Cairo abattoir constantly carry the East African bont tick through the deserts and cule tivated riparian areas of northern Sudan and Egypt, this parasite = ZA = has not established itself in these places. I failed to find it in the French Somaliland semidesert. In the Sudan, A. lepidum is common in the central grasslands and in less rigorous semideserts but quickly disappears with the approach of extreme desert conditions. It also becomes more rare in mixed forest and in forested savannah country. Except for a few specimens mentioned in the paragraph below, fewer than ten specimens have been found in Sudanese areas with over fifty inches annual rainfall (see Bahr el Ghazal records above). The presence of this tick at Katire and Gilo, in the high rainfall area of the Imatong Mountains, is difficult to explain. These specimens were taken from cattle said to have been in the Imatongs for “a long time*, but most cattle, sent as adults, were soon slaughtered as required for lumbermill workers there. Although A. lepidum thrives in moderately low rainfall areas, it should be borne in mind that in the central grasslands, where this species is common, hosts often graze in marshy areas for months during the wet season, and that the dry season grazing area (toich) is also mucky for extended periods. Just where and when these animals acquire their infestations would be of interest. DISEASE RELATIONS In the Sudan, one frequently finds that large, ugly sores have developed on cattle and horses at the site of attachment of A. lepidun. REMARKS Features of haller's organ of A. lepidum were noted by Schulze (1941), who also (1950A) discussed the dentition of this species. IDENTIFICATION Male: The eyes are small, hemispherical, dark, and in a depression (i.e. orbited). Scutal ornamentation is as illustrated BS COP = (Figure 638); note especially the pigmented spot at scutal mid length that is both within and outside of the lateral zroove; also the six or eight partially pigmented festoons. Some large, deep punctations are scattered over the scutun; these are fewer than in A. pomposum but more numerous than in A. variegatun. The lateral groove is long. This is a medium size tick measuring approximately 5.0 mm. long and 4.0 mm. wide and is easily dis tinguished from all other species. Female: Easily identified in typical specimens but its critical characters are more variable than those of the male. Rather few females are extant in collections and when this spe— cies and A. variegatum are collected from the same herd a few questionable specimens may present themselves. Typically, the scutal contour is much more narrowed pos. teriorly than that of A. variegatum, but intermediate individ— uals do exist. Scutal punctations, usually coarse and crowded only in the lateral fields, sometimes spill over into the cen tral field; small punctations scattered over the scutum tend to become rather large in a few individuals. Pigmentation of most specimens consists of a narrowly elongate area in the central field posteriorly and a pair of small spots laterally, but this character is somewhat variable. Eyes, coloration, and relative size are like those of males. — 223) = (fe; ro Figures 72 and 73, ¢, dorsal and ventral views Figures 74 and 75, 9, dorsal and ventral views AMBLYOMMA MARMOnEUM GROUP Sudan (Juba) specimens from tortoise For note on scutal lencth-width ratio of o, see IDENTIFICATION. PLATE “AVI AMBLYOMMA MARMOREUM Koch, 1844 (GROUP x (Figures 72 to 75) THE LARGE REPTILE_AMBLYOMMA,. Le NAS om Sc! EQUATORTA PROVINCE RECORDS 4 al Torit Varanus 2 niloticus** Dec iL al Torit Kinixys . belliana** Jan Oo & 26 Juba Kinixys s belliana** Dec Torit and Juba appear to be the only locality records avail- able for the Sudan. A number of Varanus lizards and tortoises in Torit and Juba Districts and elsewhere in the Sudan have been examined but this tick was found only on the three reptiles listed above. Sudan Government collections also contain specimens fron Atiambo, formerly in this Province but now a part of Usanda. *Sudan specimens referable to A. marmoreum fall into what is be. lieved to be a group of closely related African species in need of careful study before definite names cen be satisfactorily am plied. References here are for any ticks for which the name A. marmoreum has been used. The term "A, marmoreum group", as here used, includes specimens thet can be keyed to this name in the Robinson (1926) key, or that come close to it but do not equal related species such as A, nuttalli. The A. falsomarmoreun of Tonelli-Rondelli (1935) also falls into this froup. Tne disposi- tion of this group dictated by Schulze (1932A) is hopelessly um natural and practically useless. **The distribution of V. niloticus and related species is stated on page 283. K. b. belliana ranres as far west as Cameroons, where the subspecies nocueyi also occurs; the latter extends westward to Sierra Leone veridge, corresnondenre),. 79 - eS} -_- DISTRIBUTION IN THE SUDAN [ Khartoun: Omdurman (King 1926). Khartoum (Nuttall 1914A, Robinson 1920). Specimens from Omiurman and Khartoum in Sudan Government collections are from captive tortoises from unstated localities and probably represent the records or parts of the collections on which King's, Nuttall's, and Robinson's statements were made. No evidence is at nand to show that this tick occurs in nature in Khartoun Province. 7 DISTRIBUTION a marmoreum ranges throughout the Ethiopian Faunal Region, except in -he Arabian extension of this area. It appears to be more common in eastern and in southern Africa than it is in western and central Africa. Z NORTH AFRICA: ALGERIA (As A. Sparsum: Neumann 1899, This specimen was actually collected in the Paris Zoological Garden* (Bequaert, correspondence). The occurrence of A. marmoreun in Algeria and North Africa is questionable. See footnote, page 228). 7 WEST AFRICA: FRENCH WEST AFRICA (Neumann 1911). SIERRA LEONE (Entomolosical Report 1916. Hoogstraal 1954C ). CENTRAL AFRICA: BELGIAN CONGO (Neumann 1911. Nuttall and Warburton 1916. Bequaert 1930A,1931). Note: According to Theiler Pete ce the record for RuandaUrundi in Santos Dias (1954D) is incorrect. EAST AFRICA: SUDAN (Kine 1911,1926. Nuttall 1914A. Robin son 1926. Hoorstraal 1954B). *Hesse (1920) reported a female from the Leipzi¢ Zoological Gar- dens and Hoorstraal (195/40 ) another from the London Zoological Gardens. eZ Oye ETHIOPIA (Neumann 1922. Stella 1938A,1939A,1940. Charters 1946). ERITREA (Tonelli-Rondelli 1930A. Stella 1940). "“SOMALIL LAND, Gueldessa™ (Robinson 1926). BRITISH SOMALILAND (Neumann 1922. Stella 1938A,1939A). ITALIAN SOMALILAND (Paoli 1916. Tonelli-Rondelli 1932E, see also Tonelli-Rondelli 1935 as A. falsomarmoreum sp. nov. Niro 1935. Stella 1940). KENYA (Neumann 1899,1901,1911,1912. Neave 1912. Anderson 1924A,B. Robinson 1926, Bequaert 1930A. Lewis 1931A,C,1932A, 1934,1939h. Loveridge 1936B). UGANDA (Hirst 1909. Robinson 1926. Mettam 1932. Wilson 19500. Binns 1952). TANGANYIKA (Howard 1908. Neumann 1907C,1910B,1911. Morstatt 1913. Love- ridge 19230, as A. marmoreum is actually A. nuttalli according to Bequaert 1930A. “Robinson 1926. Loveridge 1925. J. B. Walker, unpublished; see HOSTS below). SOUTHERN AFRICA: ANGOLA (Specimens in HH collection). MOZAMBIQUE (Howard 1908. Neumann 1911. Robinson 1926. Santos Dias 1947A,1951A,1952 ,1953B. Hoogstraal 195/C). NORTHERN RHODESIA (Neave 1912. Robinson 1926). SOUTHERN RHODESIA (Jack 1921,1928,1937,1942). NYASALAND (Neave 1912. Robinson 1926, Wilson 1950B). BECHUANALAND (Robinson 1926). SOUTHWEST AFRICA (Tromsdorff 1914). UNION OF SOUTH AFRICA (Koch 1844. Neumann 1899,1901, 1911. Lounsbury 1905*, Howard 1908. DUnitz 1910B. Moore 1912. Bedford 1920,1926,1927,1932B. Robinson 1926. Curson 1928, Alexander 1931. Bedford and Graf 1934,1939. Neitz 1948). OUTLYING ISLANDS: ZANZIBAR (Neumann 1899,1901,1911). HOSTS The chief hosts of the components of the "A. marmoreun group” await to be determined. Tortoises and the rhinoceros were men tioned by all early workers, mostly without further data (see REMARKS). *See Robinson (1926). Se Contemporary reports of the rhinoceros, Rhinoceros bicornis subspp. as a host are chiefly those of Robinson (1020) who re— corded half a dozen collections from Kenya, Nyasaland, and Rhode sia. My collection and that of the Museum of Comparative Zo8logy contain specimens from rhinoceros in Kenya. Miss Walker's Tanga. nyika collections (correspondence) contain 47 males and seven females from four rhinoceros hosts. The white, or square—lipped, foes Ceratotheriun s, simyn, is a host in Zululand (Curson 1928). Recent records from tortoises, Testudo spp. or Kinixys belliana, are those of Neumann (1922), Robinson (1926) with numerous collections from throughout the tick's range, and Hoogstraal (1954C) from Sierra Leone. Bedford's (1932B) state ments and Theiler's unpublished records from South Africa indi. cate that tortoises are commonly infested in Transvaal and are the tick's chief host there. Other scattered records for tor— toises are those of the Sudan specimens above, Mettam (1932) for Uganda, Wilson (1950B) for Nyasaland, and Santos Dias! (1953B) summary of Mozambique ticks in which no other hosts are listed for A. marmoreum. A single collection from Tanganyika consists of ‘Six males and nine females (J. B. Walker, unpublished). The warrener or lezguan lizard, Varanus spp., is sometimes attacked, More recent reports are the single collection of Ro. binson (1926) and that from the Sudan listed above, Mettam's (1932) Uganda note, Loveridge's (1936A) Kenya record, a few lots in the Onderstepoort collection (Theiler, unpublished), and a few lots in the HH collection including one from Ancola. Some snakes are hosts, apparently only exceptionally*. Neumann's (1911) A. sparsum (said to be a synonym of A. marmoreum) *The hosts of A. s. sparsum Newnann, 1699 (according to Robinson 1926 a synonym of A. marmoreum) were reported by Neumann to be Spilotes variabilis and Testudo mauritanica from Alseria and Fast — SQ. Yariabilis is a synonym of S. p. pullatus, a large South American tree snake (Loveridge, correspondence). Therefore, since A. marmoreum (= A. sparsum) does not occur in South America, either Neumann's locality record or host record is incorrect. Tne fact that the “Algerian material of A. sparsum was collected ina European zoological garden (cf. page 226) would suggest that both Gata are difficult to assess. It further suggests that the vali- dity of the synonymy of A. sparsum should be reinvestigated. S225.— came from Spilotes p. latus (= variabilis). Puff adders, Bitis spp., have been reported by Hirst (1909), Robinson (1926), Love— ridge (1928), Bequaert (1930A), and Hoogstraal (195/C); Miss Walker's Tanganyika collection (correspondence) contains four males and a female from one puff adder. Python sp. has been listed by Neumann (1911) and Lewis (1934). Mammals, other than the rhinoceros, are also occasional hosts. Theiler's unpublished records include some adults from domestic cattle and sheep. Lewis (1939A) found specimens rarely on buffalo and on domestic cattle. Two buffaloes were listed by Bequaert (1930A,1931) and one by Robinson (1926); my collection contains 37 specimens from a single buffalo in Kenya. Neumann (1911) re- corded a genet, Genetta pardina, Robinson (1926) an eland, and Tonelli-Rondelli (1930K) a bushpig. In my collection are numerous adults from giraffes in Kenya. Alexander (1931) was unable to induce adults to feed on domestic animals. Among birds, the guineafowl h-.s been recorded as a Uganda host by Mettam (1932). Man was reported as an actual host by Charters (1946) in Ethiopia. Nymphs are sometimes taken with adults from tortoises. Dr. Theiler (unpublished records) has numerous nymphs from fowls. Seven nymphs were removed from the African hoopoe, Upupa africana, in Mozambique (Hoogstraal 1954C). Lounsbury (1905*) and Howard (1908) reported that larvae and nymphs feed readily on lizards, cattle, goats, tortoises, and birds. In the Onderstepoort laboratory, larvae and nymphs feed readily on guineapigs but adults do not (Theiler, correspondence). Kenya larvae and nymphs from females from tortoises feed well on the ears of rabbits, and resulting adults on the scrotum of a ram This is a hardy species, and sixteen-month old nymphs feed quite well while adults remain alive for 23 months without feeding (J. B. Walker, correspondence). See REMARKS below. #See Robinson (1926). = 229 2 The sorting out of these host records awaits a thorough bio. logical and taxonomic study on this interesting group of ticks. BIOLOGY (See also REMARKS below) Life Cycle Lounsbury (1905*) stated that adults will not feed on goats or on oxen if they have not first fed from a tortoise in an earlier stage. Adults attached to the host before seeking the other sex, unlike many other species of Amblyomma. Lounsbury also provided observations on time required for each stage and for feeding. These data are not abstracted here for in the light of confused nomenclature it is questionable whether the species with which Lounsbury worked is the same as that in the Sudan. Note that Alexander (1931) was unable to induce South African adults of A. marmoreum to feed on cattle. Ecology In the Ethiopian Faunal Region, this group of ticks occurs in a variety of faunal areas. Where tortoises are common these ticks are often abundant, but this incidence is by no means uni. versal in Africa. On tortoises, these ticks are usually deep in the host's axillae and it is necessary to kill or anesthetize the animal to see or secure all of the specimens, DISEASE RELATIONS Textbook statements that A. marmoreum actually has been in. criminated as a vector of boutonneuse fever of man refer merely to a remark that this was one of several tick species found on patients. *See Robinson (1926). — 2310) = Experimental attempts to transmit heartwater (Rickettsia ruminantium) of cattle through this species have falled. REMARKS A. marmoreum is considered by most workers to parasitize chiefly the rhinoceros and tortoise but Theiler (correspondence) has found so much variation in morphology and host data on spec— imens sent from various parts of Africa that she prefers to refer to all specimens as “A. marmoreum group” until they can receive more intensive study. Most of her specimens come from tortoises, a few from Varanus lizards and cattle. She considers the rhino ceros to be an accidental host, or else the host of a separate, as yet unrecognized, species or subspecies. She has large num bers of nymphs of this group from fowls. Schulze (1932A) realized the complexity of the marmoreum group and proposed new names for specimens from various parts of Africa. Reasons for these differentiations appear quite invalid. A definitive species name for Sudan material awaits assign. ment. The range of variation in even the rather small series of Sudan specimens at hand casts considerable doubt on the validity of all those “related species” that are based on certain aspects of scutal ornamentation or on coxal spur characters. The structure of the larval eye and its sense organs in spec~ imens of the A. marmoreum group has been described and illustrated by Gossel (1935). Nuttall (1914A) reported on a malformed specimen of A. mar- moreum, from the Sudan, and Schulze (1941) noted characteristics of the haller's organ of this species. IDENTIFICATION Males: Large, at least 6.0 m. x 5.0 mm Scutum with reddish-yellow ornamentation that is variable but essentially as illustrated; pale areas in this species are more separated Zoi from each other by dark stripes than they are in A. nuttalli; festoons bicolored; punctations consisting of few, scattered large and numerous small; lateral grooves deep and long; eyes flat. Females: Large, from 7.0 mm to 30.0 m, long and from 6.0 mn. to 20.0 mm. wide, depending on degree of engorgement. Scutum extensively pale ornamented and with central pale area broadly rounded posteriorly; large punctations scattered over surface including from five to twelve on posterior half; eyes flat. Although the scutal length of A. marmoreum group typically equals its width or is slightly greater than the width, the Sudan specimen illustrated (Figure 74) is exceptional in that its width is considerably greater than length. 5 CE Figures 76 and 77, co, dorsal and ventral views Figures 78 and 79, 9, dorsal and ventral views AMBLYOMMA NUTTALLI Sudan specimens PLATE ¥XVII = 2a) S AMBLYOMMA NUTTALLI D'énitz, 1909. (Figures 76 to 79) THE SMALL REPTILE-AMBLYOMMA LyN 2 OC. EQUATORIA PROVINCE RECORDS Reptiles ¢/ Farajok Kinixys b. belliana Mar ty) Torit Xnie b. belliana Aug 3) Meridi, 50 mi. northeast of Kinixys b. belliana Oct (SVS) alt Torit nese: ne ni Loticus Dec 1 Torit Varanus e. exanthematicus anthematicus Jan Mammal a Torit Ourebia ourebi aequatoria Feb Bird a Torit Francolinus clappertoni gedgii Jan These specimens indicate the presence of A. nuttalli on both the east and west banks of the Nile in Equatoria Province. Sudan Government collections contain specimens only from Ossa River (H. H. King legit, 1913), now a part of Uganda. King (1926) mentioned no Sudan Lo cality records for this species. DISTRIBUTION IN THE SUDAN Kordofan: As A. werneri (Schulze 1932A), a single specimen from Talodi. The host 1s most probably the “Cinixys belliana” mentioned by Werner (1924). SS BV DISTRIBUTION A. nuttalli is widely spread throughout the African continent within the Ethiopian Faunal Region. WEST AFRICA: NIGERIA (Simpson 1912A,B. Robinson 1926). GOLD COAST (Robinson 1926. Stewart 1937). FRENCH WEST AFRICA (Villiers 1955). PORTUGESE GUINEA (Tendeiro 1951C ,D ,1952A,C ,D ,1953,1954). CENTRAL AFRICA: CAMEROONS (Donitz 1909. Rageau 1951,1953A,B). scat T CONGO (Schwetz 192%. Bequaert 1931. Theiler and Robinson 1954). EAST AFRICA: SUDAN (King 1926. As A. we werneris: Schulze 1932A. Hoogstraal 1954B). 7 ies ITALIAN SOMALILAND (See REMARKS below). KENYA (Loveridge 1929. Bequaert 19304). UGANDA (King 1926, see DISTRIBUTION IN THE SUDAN above. Robinson 1926. Mettam 1932, Wilson 19500). TANGANYIKA (Donitz 1909. Robinson 1926. Love ee as A. marmoreum is actually A. nuttalli, see Bequaert 1930A). SOUTHERN AFRICA: SOUTHERN RHODESIA (Dénitz 1909. Jack 1942). MOZAMBIQUE (Santos Dias 1949B,1950A,B,1951A,1952D ,1953B,1955A,B). UNION OF SOUTH AFRICA (Curson 1928. Alexander 1929,1931. Bedford 1932. Neitz 1948). HOSTS Adults All authors list land tortoises (Kinixys spp. or Geochelone pardalis) as the chief hosts of adult A, nuttalli. A record of the side-neck turtle, Pelomedusa s. subrufa (= P. galeata), as a host (Santos Dias 1953B) was based on misidentification of Pelusios Ss. sinuatus, "a species of lesser importance as a host™ (Santos Dias 195 5B ) ° 5 C35) So Infrequent hosts of adults are monitor lizards (Varanus spp.) (Robinson 1926, Tendeiro 1951D, and Sudan record above), Agama lizard (Loveridge 1929, Bequaert 1930A), python (Bedford 15725), hedgehog (Robinson 1926), man (Schwetz 1927C), and one specimen from a domestic goat (Theiler, unpublished). Nymphs and Larvae Immature stages infest tortoises and also Varanus lizards, birds, and hares. Guineapigs may be used for laboratory rearing. Owing to the paucity of field records for immature stages it is impossible to determine their host preference in nature. It is unusual to find a tick that normally feeds on warm blooded animals in the immature stages and on cold blooded animals in the adult stages; the reverse is usually true. Yet Theiler (correspondence) has nymphs from South African hares and from a turkey on a farm where the mountain tortoise is also common. Further field study of this matter is indicated but, as A. nuttalli appears to be curiously localized and seldom abundant, the success of such investigation will depend on local factors. Note that in Equa toria Province, single nymphs were found on each of two species of Varanus lizards, on an oribi (antelope), and on a francolin partridge, but none were taken from the many tortoises examined. Various literature records for "iguana" lizard, a nonAfrican reptile, should be ™“leguan™ or monitor lizard (Varanus spp.). Alexander (1931) was unable to induce adults to feed on laboratory animals. See also BIOLOGY below. BIOLOGY Santos Dias (1950A) reared this species using guineapigs and tortoises as hosts. He subsequently reported (1955B) that the life cycle is a three-host type. A maximum of 22,891 eggs from a single female were noted with the claim that this is the great— est number of eggs yet observed in any of the Ixodoidea. The minimum period for completion of the life cycle is estimated at 134 to 151 days, the maximum period 217 to 296 days. This paper Bey PIs oe is illustrated with photographs of both sexes feeding from the interstices of the host's shields. Our Torit adults, however, “were taken in the host's axillae (during a native big-game hunt and stored in a hunter's ear, plugged with mud, for three hours until our lost vials could be recovered). DISEASE RELATIONS Experimental attempts to transmit heartwater (Rickettsia ruminantium) of cattle by this tick species have failed. It is claimed that specimens have been found infected with Q fever (Coxiella burnetii) in Portugese Guinea. As with the Aponomma parasites of lizards and snakes, it is of interest to conjecture that the small reptile-amblyomma may be a vector of the hemogregarines of tortoises. REMARKS Misshapen specimens have been reported (Santos Dias 1949B, 1950A ,1955A). liana (see Werner 1924) from Talodi, Kordofan, Sudan, ee to be a synonym of A. nuttalli. Following Schulze's practice of applying species names to any variant, he distin guished a single specimen as different from A. nuttalli for the following reasons: the dark marking not blackish-brown, but light red brown on a light reddish brown background; darker markings bounded with a coppery color (in A. nuttalli dark yellow brown without copper borders); median stripes more irregular than in A. nuttalli and broadened at the ends; lateral groove sharply defined against the scutum, in A. nuttalli irregular; and ventral median muscleplate smaller. Amblyomma werneri werneri Schulze, 1932(A), described from nixys b. All characters proposed to separate A. werneri from A. nuttalli fall well within the normal range of variation due to age, nutrition, or methods of preservation. In long series of any Amblyomma species, = 23S some specimens vary in roundness, flatness, development of the ventral muscleplate, and sharpness of the lateral groove. Com parison of many specimens of this genus preserved in alcohol with those preserved as dry specimens shows that those preserved in alcohol frequently develop a coppery sheen due to chemical change. Theiler has made similar observations in this respect. The obscw rity of the color pattern and its overlay with a basic color in some specimens in any extensive collection of amblyommas from even a single host is taken for granted by most students. Using the above mentioned criteria, proposed by Schulze, large collec- tions of A. variegatum and A. lepidum from single herds of cattle have been examined. 1t has been found that each collection con tains no less than four "species and up to seven “species”. Comparison of Sudan specimens with others from various parts of Africa and of the type specimens of A. nuttalli in British Museum (Natural Histo a reveals no significant differences to obtain between any of then. It is for these reasons that it has been proposed (Hoogstraal 1954B) to consider A. werneri werneri Schulze, 1932(A), as a syno nym of A. nuttalli Donitz, 1909. It is also of some interest to consider the status of A. werneri poematium Schulze, 1932, described on the basis of two es from a young rhinoceros, at the Amsterdam zoological gardens, from East Africa. This subspecies was distinguished by "a wonder- ful metallic, copper, partly greenish gloss (with) brown elements of the conscutum bordered in copper", in one of the two specimens, but in the other "the structure producing the metallic coloration was in greatest part destroyed, only in a few places did the greenish coppery sheen show up”, The size of these specimens was also larger than that of the subspecies werneri. I have seen a male specimen taken from a Somali tortoise (#17691, Rocky Mountain Laboratory, Hamilton, determined as A. werneri by Dr. E. Stella). This tick answers the description of A. w. poematium but has a somewhat rugose scutum suggestive of injury during molting or during an immature stage. The specimen resembles a teneral individual, i.e. one that has been preserved shortly after molting while still bloated and before the colors are fast. = eoole Santos Dias (1954G) opines that (1) A. poematium is a sepa rate species, (2) A. schlottkei Schulze, 1932, might be a synonym, and (3) A. faiai Santos Dias, 1951, definitely is a synonym. The specificity of A. poematium is hardly convincing on the basis of descriptions and illustrations, though there is a possibility that comparison of specimens may provide yet unmentioned clues to separate this morphologically from A. nuttalli. Breeding experi- ments are also indicated. IDENTIFICATION A. nuttalli is similar to A. marmoreum in characters mentioned under that species, except for the following: Males: Size is smaller, always less than 6.0 m. long. Pale ornamentation of the scutum is somewhat variable, but all specimens are like the one illustrated in that the dark areas are less extensive and more broadly separated from each other by light areas than they are in A. marmoreun. Females: This sex is also smaller than that of A. marmoreum (body approximately 7.0 mm. long, 5.5 mm. wide; scutum about 3.2 mn. long, 3.3 mm. wide); the central pale scutal ornamentation tapers to a narrow point posteriorly and is therefore very dis. tinctive. = 239) S Figures 80 and 81, @, dorsal and ventral views Figures 82 and 83, 9, dorsal and ventral views AMBLYOMMA POMPOSUM co specimen from Belgian Congo Q specimen from the Sudan PLATE XXVIIT = 240)= ANBLYQMA POMPOSUM Donitz, 1909. (Figures 80 to 83) THE HIGHLAND BONT TICK Lil Or S EQUATORIA PROVINCE RECORD Al Yei domestic cow Jan This specimen, identified by Dr. G. Theiler, is the only one of this species from the Sudan. It represents an apparently rare intrusion into the western half of Equatoria Province from the Belgian Congo. Ecologically, the Yei area, except for a few hill masses, does not seem suitable for the survival of this mountain. inhabiting species though other localities in the eastern high lands of Equatoria Province might well meet its requirements. See A. superbum, REMARKS below. DISTRIBUTION A. pomposum is a highland tick of eastern Central Africa, adjacent parts of East Africa, and northern parts of southern Africa. See also REMARKS and IDEVTIFICATION below. CENTRAL AFRICA: BELGIAN CONGO and RUANDALURUNDI (Nuttall and Warburton 1916. Seydel 1925. Robinson 1926. Schwetz 19274. Bequaert 1931. Weitz 1947. Schoenaers 1951A. Theiler and Ro binson 1954. Note: Santos Dias 1953E refers most of these Congo reports to his A. superbum sp. nov. However, correspondence with curators reveals that-he had not examined the specimens on which these references were based). EAST AFRICA: SUDAN (Hoogstraal 1954B). KENYA (See Note ugpder IDENTIFICATION below). UGANDA (Hoogstraal 1954C). TANGANYIKA (Donitz 1909). SOUTHERN AFRICA: ANGOLA (Leitao 1942. Robinson 1926. Santos Dias 1950. SBacelar 1950. Sousa Dias 1950, this report referred age ee to A. superbum sp. nov. by Santos Dias 1953E; see IDENTIFICATION below. WEEE and Robinson 1954. Rousselot 1953B). MOZAMBIQUE [ Robinson 1912%, ofa Santos a 19474 ,1953B* ,1954A,C#. Ac cording to Theiler (correspondence), A. variegatum govurensis Santos Dias (1950B), from the extreme north of Sul = Save Prov- ince, is synonymous with A. pomposum. Recently, Santos Dias (1953E) has agreed with this view; see IDENTIFICATION below 7. NORTHERN RHODESIA (Robinson 1926. Matthysse 1954. Theiler and Robinson 1954). SOUTHERN RHODESIA (See REMARKS below). HOSTS Domestic cattle are referred to as hosts of A. pomposum by most authors, but Matthysse (1954) found it only on ao hosts, and then rarely, in Northern Rhodesia. Mules (Nuttall and War- burton 1916. Robinson 1926. Sousa Dias 1950). Horses (Theiler, areas records). Sheep, goats, dogs, donkeys (Sousa Dias 1950). Man (As synonymous A. variegatum nocens: Robinson 1912* and subsequently frequently quoted without additional observations). “Striped antelope” (D8nitz 1909). Sable antelope, roan ante— lope, and eland (Robinson 1926, Schwetz 1927A; Congo specimens in Onderstepoort collection). Hartebeest, kudu (Robinson 1926). Zebra (Schwetz 1927A). Buffalo (Jack 1942*). Warthog (Schoenaers 1951A). Ankole topi, Damaliscus korrigum ugandae (Hoogstraal 1954C). The nymphal specimen from a monkey, mentioned by Santos Dias (1954C), should be checked against A. variegatum. “Wild hosts only in Northern Rhodesia” (Matthysse TOELT. BIOLOGY See REMARKS and IDENTIFICATION below. All authors who refer to collecting localities for A. pomposum stress the fact that it is a highland species. ¥This record should be read in conjunction with statements in REMARKS and in IDENTIFICATION below. ee oe DISEASE RELATIONS MAN*; A. osum is said to attack African children's heads and causes ei of the skin. This has not been substantiated. CATTLE: Inflammation and sloughing of mammae. Heartwater (Rickettsia ruminantium). HORSES: Pyolymphangitis. REMAR KS According to Robinson (1912*), A. pomposum (= A. variegatum nocens) occurs in the Rhodesias and Mozambique chiefly in bushveld from 2000 to 3000 feet elevation and seldom above 4000 feet. He further stated that this tick is notorious for the damage it does to stock in the Rhodesias, where it is known as the “Pyaemia tick”. However, Morris (1933,1935,1936) attributed “tick.pyaemia™ in Northern Rhodesia to A. variegatum. In Southern Rhodesia, Jack (1918) referred “ixodic lymphangitis™ to A. variegatum, Sinclair (1916) associated skin diseases of cattle with A. variegatum, and Jack (1928,1937,1942) also mentioned only A. variegatum with reference to abscesses and sloughing of the hosts* skin. In his first two papers Jack did not differentiate between A. variegatum and A. sum, but in his 1942 report he stated that the Local highlands where A. sum would be expected to occur, are free of amblyommas but that some male specimens of A. variegatum from the lowlands may show a tendency to resemble A. pomposum. See also IDENTIFICATION below. Theiler (correspondence) calls attention to the following facts that may modify many of the above reports concerning A. pomposum. Robinson's (1912,1926) remarks concerning A. pomposum in Mozambique an& Southern Rhodesia are based on statements of Mr. E. M. Jarvis. Jack's records for Southern Rhodesia apparent. ly are quoted from the same source, for no further data are pre. sented. The extensive Onderstepoort collection has no speci. mens from Southern Rhodesia. Theiler's correspondence with Dr. Lawrence, Assistant Director (Research) of the Southern Rhode- sia Veterinary Department, indicates that he is not aware that A. *This record should be read in conjunction with statements in REMARKS and in IDENTIFICATION below. yeh ta sum occurs in this territory and that he considers the Jarvis statements as "sheer nonsense™, It appears, therefore, that earlier literature records for A. pomposum in Mozambique and Southern Rhode— sia are open to question. Wherever A. pomposum occurs it seems to be present in good numbers. Its distribution, so far as Theiler has determined (cor_ respondence), is mainly in the Rhodesia Highland Savannah type of vegetation, certainly not in the moist vegetation of the Umtali- Melsetter district and adjoining Manicaland. More extensive search for and study of this species is required. Statements regarding damage to cattle and to children by this tick in Southern Rhodesia appear to be questionable. Sousa Dias (1950) writes concerning A. pomposum, which is com mon in the Angolan highlands, as follows: wrt 1s considered by breeders to be one of the most harmful ectoparasites of stock for it causes wounds that are most difficult to heal. It is probable that (this tick) is one of the factors that favors the dispersal of bovine dermatoses so common in Angola™, He surmises, that A. sum is a heartwater vector in Angola inasmuch as it occurs in heartwater areas in the absence of other recognized vectors. [ Neitz (1947) showed that A. pomposum is a vector of heartwater_7. A. pomposum is close morphologically to A. lepidum and to A. variegatum. The latter, biologically, is a most versatile tick except that it shuns desert and rainforest areas. A. lepidun is a semiarid country and savannah species. A. Iposum appears to be chiefly a highland species. See also remarks on A. superbum in section below. A gynandromorph of this species has been described by Santos Dias (1954). Schulze (1932C) discussed certain features of the ornamentation of A. pomposum in relation to other species in this genus. Sy ih IDENT IFICATION Both sexes have hemispherical eyes situated in a depression, and are distinct from A. variegatum and A. lepidum in possessing very coarse scutal punctations. Male scutal ornamentation, inside the lateral groove, is like that of A.lepidum except that a small red spot may be present laterally in A. pomposum, but no red color is found on A. lepidun. The female scutum may be variable in length-width ratio, that of some specimens being only as long as wide, of others longer than wide; its rugosity is very distinct; it may be unornamented but usually has a small white or pale spot in the posterior field. £ Robinson (1926) stated that QQ are unornamented. Nevertheless a number of those in the Nuttall collection, which comprised his chief reference material, have ornamented spots on the scutun. These, as well as others similar to them have been observed in different collections. 7 Specimens from the Sudan referable to these characteristics should be checked against authoritatively identified specimens from the known range of A. pomposum before this name is applied. Char acters provided here are generalized; an exhaustive survey of the subject is precluded by our uncertainty over variability between this species and A. variegatum, as indicated below. Note In some large collections of A. variegatum, a few robust male specimens, or, more rarely, a few pygmy male specimens, may be sug- gestive of A. pomposum owing to unusually heavy scutal punctations. Associated females are also more heavily punctate and may have a wider scutum than normal. Jack (1942) referred to similar males from Rhodesia (see REMARKS above). I have collected a few lots of such specimens in the Sudan and in the mountains of Yemen (Arabia). The most distinctive collection in this category is one recently presented to me by Dr. C. B. Philip, who collected it from a herd of cattle near Kabete in the mountains of central Kenya. The single female has a wide scutum and is heavily punc- tate but not rugose and the punctations are not confluent. Of = 245 = the males, four are slightly more punctate than is usual for A. variegatum, one is slightly more punctate than the first four, and the last two are so heavily punctate that, alone, there would be little question of their identity as A. pomposum. Such spec. imens, in addition to various queries oy mentioned, suggest the possibility that A. pomposum is a heavily punctate, mountain or heavy forest subspecies of A. variegatum and that intergradation does occur. It appears that A. variegatum govurensis of Santos Dias (1950B,1954H) is an intermediate form between the almost nonpunctate A. variegatum and the heavily punctate A. pomposum. Santos Dias! description adds weight to the concept that A. pomposum is actually no more than a variant form or subspecies of A. variegatum. Rearing of progeny from isolated females in lowlands and in highlands and transporting some of their progeny to different altitudinal levels for development under different ecological conditions may solve this question. Since the above was written, Santos Dias! (1953E) paper des cribing A. superbum sp. nov. has appeared. In it, A. variegatum ovurensis is placed in synonymy under A. pomposum. A. superbum Te considered to differ from both A. variegatum and A. pomposum chiefly on the basis of size, depth, and distribution of puncta. tions. Even more recently, the same author (1954H) has reaffirmed the validity of his variety of A. variegatum, with no indication of what he proposes to do about A. superbum. On ecological grounds, A. superbum (or A. variegatum govurensis) might be a useful niche in which to drop the Sudan specimen and certain other Central African lowland specimens. Variable and confusing series of specimens still confront us. Unfortunately, however, A. superbum does not answer the problems this material poses. No recourse offers itself but to maintain the present systematic status of A. variegatum and A. pomposum, undertake biological studies suggested in the paragraph above, and only then judge the presently considered questionable validity of A. superbum as a real species and the range of variation in A. vari ariegatum and A. pomposum. = 2464 Santos Dias (1953E) further refers the Belgian Congo records of A. pomposum by Nuttall and Warburton (1916), Schwetz (1927A), pesTagee CISsT) and Schoenaers (19514) to A. superbum. Since no adequate descriptions for differentiating Congo specimens were provided by these authors, the validity of this proposed synonymy is highly questionable. Belgian Congo specimens that have been seen in British Museum (Natural History) collections, in Museum of Comparative Zoology collections, and in the HH collection are typically A. pomposum by comparison with specimens from everywhere within the range of this species. In conclusion, one may only belabor the point: the status of heavily punctate specimens morphologically intermediate between A. variegatum and A. pomposum remains to be ascertained by bio- topcase not by museum-type studies. 5 LY = Figures 84 and 85, do, dorsal and ventral views Figures 86 and 87, 9, dorsal and ventral views AMBLYOMMA RHINOCEROTIS an specimens PLATE XxIX oS Ais = AMBLYOMMA RHINOCEROTIS (de Geer, 1778) (= A. PETERSI Karsch, 1878) (Figures 84 to 87) THE RHINOCEROS AMBLYOMMA L NO oveo. EQUATORIA PROVINCE RECORDS ds Torit on grass Dec (SGC) 22 Kajo Kaji on grass - (BMNH) The Sudan Government collection specimens were collected by H. H. King. British Museum (Natural History) specimens were taken by Captain C. M. Stigand. DISTRIBUTION IN THE SUDAN Upper Nile: Bor (King 1926). DISTRIBUTION A. rhinocerotis occurs in central, eastern, and southeastern Africa apparently wherever the mitingeenos is found. WEST AFRICA: LIBERIA: Bequaert (1930A) states that Neumann's (1901,;1911) Liberian records of this species, repeated by Bedford and Hewitt (1925) and by Bedford (1932B) are in error. FRENCH EQUATORIAL AFRICA: Neumann (1899) listed A. aureum (a synonym of A. rhinocero- tis) from "Ngourou Plains, Zanzibar™, It is probable that this local- ity is actually N'Gourou, Ubangi-Shari, French Equatorial Africa./ CENTRAL AFRICA: BELGIAN CONGO (Schwetz 1927A. River Misisi, Schwetz 192%, p. 92, is in Uganda. Tonelli-Rondelli 1930A. Bequaert 1931). NOTE: According to Theiler (correspondence), the record for Ruanda-Urundi by Santos Dias (1954D) is in error. = QE) = EAST AFRICA: SUDAN (King 1926. Hoogstraal 1954B,C). BRITISH SOMALILAND (Neumann 1922. Stella 1938A,1939A,1940). TTALIAN SOMALILAND (Tonelli-Rondelli 1930A. Stella 1940). KENYA (Neave 1912. Neumann 1913,1922. Anderson 1924A,B. Robinson 1926. Bedford 1932B. Lewis 1931C,1934. Weber 1948). UGANDA (Neave 1912. Neumann 1922. Robinson 1926. Schwetz 1927, p. 92, as Belgian Congo. Bequaert 1930A, p. 803. Mettam 1932, 1933. Wilson 19500). TANGANYIKA (Neumann 1901,1907C ,1910B,1911. Neave 1912. Morstatt 1913. Robinson 1926. J. B. Walker ; un published) . SOUTHERN AFRICA: NORTHERN RHODESIA (Neave 1912. Robinson 1926). SOUTHERN RHODESIA (Jack 1942). NYASALAND (Neave 1912. Robinson 1926. Wilson 1950B). MOZAMBIQUE (Karsch 1878. Neumann 1911. Santos Dias 1947A,1953B). UNION OF SOUTH AFRICA (Breijer ee) Bedford and Hewitt 1925. Curson 1928. Bedford 1932B, 1936). OUTLYING ISLANDS: MADAGASCAR: Neumann (1901,1911). Poisson (1927). Locality record probably erroneous, cf. Hoogstraal (1953E). ZANZIBAR: Neumann (1899) probably in error, see WEST AFRICA above./ Note: Neumann (1899) listed JAVA for the synonymous A. aureum but subsequently (1908) he stated that the specimen on which this record was based was actually A. testudinarium. 7 HOSTS All workers list as hosts either the black rhinoceros, Diceros bicornis, or the white or square-lipped rhinoceros, Ceratotherium Simm > the latter in both the northern and eonubema areas of 1ts Ses. Other animals that uncommonly serve as hosts are: eland (Neumann 1907C ,1910B,1911), tortoise (Bedford 1936), and python (Mettam 1932). Domestic cattle: numerous adults, in a single lot; Uganda Veterinary Service collections. BIOLOGY Unstudied. = 250) 6 DISEASE RELATIONS Unstudied. REMARKS The frequent records of specimens taken from grass are due to the large size and conspicuousness of the rhinoceros amblyomma,. This species often has been referred to as A. petersi (Karsch, 1878), but according to Schulze (1932A), this name is synonymous with A. rhinocerotis (de Geer, 1778). This decision is acceptable for the present, but it must be noted that Theiler (correspondence) is far from certain of its validity. Obviously needed is a careful study of the original material and literature by a competent contem porary student with full access to pertinent specimens. The specific name rhinocerotis (de Geer, 1778) frequently has been applied to Dermacentor rhinocerinus (Denny, 1843). However, as Bequaert (1930B) pointed out, Donitz (1910B) has long ago indit cated that de Geer's specimens belonged in the genus Amblyomma be- cause of their longer palpi. The remarkable parallel or convergent evolution of rhinoceros. infesting Amblyomma and Dermacentor ticks, and the relationship of Cosmiomma amense (Denny, 1043), a hyalommalike beast, is worthy of ee study. The capsule of larval haller's organ in A. rhinocerotis has been noted by Schulze (1941), who also (1950A) discussed the adult dentition of this species. IDENTIFICATION Males, at least 8.0 mm. long and 7.0 mm. wide, are as large as any other African amblyomma. “The scutum lacks lateral grooves, has bicolored festoons, small and flat eyes, extensive pale (yellowish) ornamentation on a dark (reddish-brown) background, and a few large scattered punctations. The leg segments have nar row pale distal rings. = 25S Females are also very large, approximately 9.0 mm. long and 8.0 mm. wide. The scutum is largely pale (reddish or golden) with lat— eral margins and small internal areas dark reddish-brown; it bears few coarse punctations on the anterior half but numerous fine punc— tations; eyes are flat or very slightly convex. The leg segments exhibit narrow, pale distal rings. = 252 = Figures 88 and 89, co, dorsal and ventral views Figures 90 and 91, 9, dorsal and ventral views AMBLYOMMA THOLLONI Sudan Specimens PLATE XXX = 259) S AMBLYOMMA THOLLONI Neumann, 1899. (Figures 88 to 91) THE ELEPHANT AMBLYOMMA Lb NN © Fo EQUATORIA PROVINCE RECORDS 19 178 Lotti Forest Loxodonta africana oxyotis Apr SS Tereteina Loxodonta africana OxyotLs Feb a fk Torit Loxodonta africana oxyotis Dec [he X8s Torit Loxodonta africana oxyotis Dec 1* Lokila Chameleo g. gracilis Oct (SVS) King (1926) listed Equatoria Province without localities and his specimen vials include no further data. The nymph from a chameleon was identified by Dr. G. Theiler. DISTRIBUTION IN THE SUDAN Although it may have been reasonable to expect that A. tholloni occurs on elephants in Bahr El Ghazal and Upper Nile Province and on the west bank of Equatoria Province, no specimens have been col. lected to indicate its presence in these places. Ticks from several elephants shot near Yirol and Kenisa in Bahr El Ghazal and Upper Nile Provinces in 1911, 1953, and 1954, have all been R. simus simus, R. simus senegalensis, or intergrades of these two subspecies. DISTRIBUTION A. tholloni occurs through much of tropical Africa, wherever the African elephant, Loxodonta africana subspp., is found, except possibly along the northern and southern margins of the host's range. 5 Gy WEST AFRICA: LIBERIA (Bequaert 1930A). SIERRA LEONE (Simpson oe Robinson 1926). IVORY COAST (Rousselot 1951,1953B. Villiers 1955 )- CENTRAL AFRICA: CAMEROONS (Neumann 1901,1911. Ziemann 1905, 1912A. Rageau 1951,1953A,B). RIO MUNI (Robinson 1926). FRENCH EQUATORIAL AFRICA (Neumann 1899. Tonelli-Rondelli 1930A. Fiasson 1943B. Rousselot 1951,1953B. Rageau 1953B). BELGIAN CONGO (Neumann 1899,1911. Nuttall and Warburton 1916. Roubaud and Van Saceghem 1916. Robinson 1926. Schwetz 1927A,B,C, 1932. Schouteden 1929. Tonelli-Rondelli 1930A. Bequaert 1930A,B, 1931. Rodhain 1936. Fain 1949. Theiler and Robinson 1954. Van Vaerenbergh 1954). NOTE: According to Theiler (correspondence), the record for RuandaUrundi by Santos Dias (1954D) is in error. EAST AFRICA: SUDAN (King 1911,1926. Robinson 1926. Hoogstraal 1954B). KENYA (Neumann 1922. Lewis 1931C,1932. Mulligan 1938). UGANDA (Neave 1912. Robinson 1926. Tonelli-Rondelli 1930A. Mettam 1932. Wilson 1950C. See HOSTS below). TANGANYIKA (Neumann 1899,1907% , 1910B,1911. Morstatt 1913. Robinson 1926. Hoogstraal 1954C. J. B. Walker, unpublished; see HOSTS below). SOUTHERN AFRICA: ANGOLA (Gamble 1914. Robinson 1926. Santos Dias 1950). MOZAMBIQUE (Santos Dias 1947A,B,194& ,194% ,D ,1950B, 1953B,1955A. Bacelar 1950. Tendeiro 1952). NYASALAND (Neumann 1899*, Neave 1912. Robinson 1926. Wilson 1950B). A. tholloni has not yet been recorded from the UNION OF SOUTH AFRICA, but Theiler (correspondence) believes that this is probably because it has not been looked for and that it possibly occurs on Kruger Park elephants and on remote herds in Southwest Africa. The possibility that this tick is incapable of following its normal *The reference to “region du Nyassa" by Neumann (1899), for spec imens collected by Ed. Foa, may refer to Niassa Province, Mozambique. = 255) host into the southern periphery of the host's range should be considered in view of its apparent total absence on elephants in Bahr El Ghazal Province of the Sudan./ HOSTS All authors list the African elephant, Loxodonta africana subspp., as chief host. Uncommon hosts that have been reported for a few specimens are: gazelle (Neumann 1901,1907% ,1910B), antelope and domestic horse (Neumann 1911), rhinoceros (Neumann 1922), leopard (Robinson 1926), bushpig and large lizard (Schwetz 19278), domestic dog (Santos Dias 1953B) and buffalo (Hoogstraal 19540). Dr. Theiler has larvae, nymphs, and a male specimen from a bird, Pitta reichenowi, another indication of ground birds as hosts of immature Amblyomma species. In our collection are several nymphs and a male with massive legs taken from a hippopo. tamus in Kazinga Channel of Lake Edward by Lt. Colonel Don Davis, U.S. Army. Miss J. B. Walker has a collection consisting of seven males and four females from a Tanganyika black rhinoceros; also others from an elephant there. The small nymph collected at Lokila, Equatoria, from a chameleon by E. T. M. Reid is an unusual record. Santos Dias (1948) states that larvae and nymphs are rarely found on ele— phants. The only other records for nymphal hosts in nature are those of the bird and hippopotamus listed above, and one nymph in the Onderstepoort collection (Theiler, unpublished) with adults, from an elephant at Toro, Uganda. BIOLOGY Adult specimens of the elephant amblyomma may be found on any part of the host's body, Immature stages previously have been reported only by Santos Dias (1940), who states that larvae and nymphs are rarely taken on elephants. Santos Dias experiment— ally reared larvae and nymphs on guinea pigs and reported that six months were required to complete the three-host life cycle. He observed a chalcid wasp parasite, Hunterellus hookeri, infesting nymphs. Fiasson (1943B) reported 3000 egrs from an engorged fe— male. = -250.— Mr. J. Owen, who furnished the 197 specimens from a single elephant in a plains herd passing through Lotti Forest, reported that his "boys" could have collected at least twice as many from this elephant if they had had more containers for them. No spec- imens other than the few listed could be found on the three other Equatoria Province elephants noted above. Numerous other newly killed elephants in this Province have been examined with. out finding ticks of any sort. DISEASE RELATIONS A. tholloni is possibly a vector of Nuttallia loxodontis of elephants. REMARKS The stage to stage growth of A. tholloni has been charted by Campana-Rouget (1954). Misshapen specimens have been described and illustrated by Santos Dias (1947B,194% ,1955A). Larval and nymphal stages of A. tholloni were described and illustrated by Santos Dias (1949). Variations in male scutal patterns from Mozambique were il. lustrated by Santos Dias (194.78). Within the geographical range of the elephant amblyomma there are two scutal color patterns, one drab and lightly pigmented with small areas of color, the other brightly marked, usually with more extensive pigmented areas. The bright form is particularly common among numerous specimens seen from West Africa and rare among those from East Africa; the drab form is common in East African specimen and rare in West African material. After having examined all of the numerous A. tholloni specimens in British Museum (Natural History) collec tions, which represent almost all areas of the geographic range of this species, one may only conclude that these two color patterns do not appear to be genetic variants and are probably not associated with temperature or rainfall factors or with methods of preservation. They may possibly derive from nutri- tional factors. A biological study of living specimens is the 5 AN) & only means of determining the reason for these two color patterns. Since the above was written, Rageau (1953B) has reported that ameroons specimens and others that he has seen from French Equatorial Africa all show the reduction in ornamentation that has appeared to me to be more common in East than in West African specimens./ Robinson (1926) stated that a large pale spot in each lateral field of the scutum and a stout spur on coxa IV are female diag nostic characters. I have examined Nuttall's lot 3381 in British Museum (Natural History), on which Robinson's species definition and illustrations were based, and find it to be the most heavily and liberally ornamented material, along with a few others fron Sierra Leone, of any representatives of this species in the cole lection. Actually, lateral field pale spots are absent in most of these specimens. The stoutness of the spur on coxa IV is also a variable character and the specimen selected by Robinson is an extreme example. In most specimens, this spur is merely a small pointed projection from or near the posterior coxal margin. Ali, Sudan specimens at hand are drably colored and their pig- mented areas are no more extensive than those illustrated (Figure 90). On some, ornamentation is almost obsolete. IDENTIFICATION Males. No other African amblyomma can be confused with this species because of the smallness of its pigmented areas, flat eyes, and absence of both lateral grooves and of large scutal punctations. Males measure about 5.0 mm. long and 4.0 mm. wide. Females are equally easily distinguished from all others in Africa by the absence of large scutal punctations, triangular scutal shape with narrow posterior margin, and distribution of color pattern that is usually only an irregular spot in the pos terior point but sometimes also has small lateral spots. The cer- vical grooves are short and eyes are large, flat, and pale. Fe. males, unengorged, are usually about 6.0 mn. long and 4.7 mm. wide. = 250 = Figures 92 and 93, o, dorsal and ventral views Figures 94 and 95, 9, dorsal and ventral views AMBLYOMMA VARIFGATUM Sudan Specimens PLATE XXXTI WD F293) FW pr WN Oo 15 OO mi oo BHR FOOrW)D AMBLYOMMA VARIEGATUM (Fabricius, 1794). 7 o HW (Figures 92 to 95) THE TROPICAL BONT TICK EQUATORIA PROVINCE RECORDS Tkoto MAN (feeding on) Feb Torit Atelerix pruneri oweni Nov Torit Euxerus erythropus leucoumbrinus Dec Kapoeta Lepus capensis subsp. Apr Jebel Kathangor Crocuta crocuta subsp. Dec Torit Genetta tigrina ae uatorialis Dec Torit Canis aureus soudanicus Dec Torit Canis aureus soudanicus Apr Kapoeta Herpestes sanguineus sanguineus Apr Boma Plains Holo Syncerus caffer aequinoctialis Dec Syncerus Caffer aequinoctialis Mar Koss Valley Syncerus caffer aeauinoctialis Jan Laboni Syncerus caffer aequinocti is Feb Torit Syncerus caffer aequinoctialis Jan Kapoeta Hippotragus equinus bakeri Dec Lafon Hippotragus equinus bakeri Dec Torit Hippotragus equinus bakeri Apr Torit Hippotragus equinus bakeri Jan Jebel Kathangor Gazella granti brighti Dec Torit auth cerpe grimma roosevelti Feb Tora hync tragus cuenther i smitnhii Dec Torit Rhynchotr guentheri smithii Apr Lokila Ourebia ourebi aequatoria Feb Torit AlceLaphus buseLaphus roosevelti Apr Loronyo Alcelaphus buselaphus roosevelti Jan Loronyo Taurotragus oryx pattersonianus Jan = 200) = (2) NIN Now 26 nN joa) OOF OO OND 22 ake. 3 Ovi (en 1 10 Ad 30 Lip 9B 13 3S 209.6 34 78 ERE l- Gage 3 a 1 3 ee) Sat ya byel9 Panee 56 152 8 10 15d 103 2 4 98 148 Iyt 3 4 8 Dee, <7 af Kapoeta Nagichot Nagichot Torit Torit Torit Torit Torit Goniryo Katire Gilo Juba Juba Rejaf Gondokoro Tombe Muni Terakeka Yei Yei Yei Yei Lorella (Yei River) Kajo Kaji Kajo Kaji Meridi Meridi Amadi Yambio Li Rangu Kapoeta Juba Kaguada Katire Kajo Kaji Kajo Kaji Katire Katire Juba Katire Torit domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic on crass Seasity cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle cattle sheep sheep sheep goats goats dogs dogs dogs horses pig BIRDS 1 Torit Tchagra senegala erlangeri Jan 2 Torit Sphenorhynchus abdimi1 Jan 5 Re) RSE Re BSc 3 ee Kapoeta Lissotis m. melanogaster Dec (2) iW Torit domestic chicken Jan 7 Katire domestic turkey Jan DISTRIBUTION IN THE SUDAN A. variegatum occurs in Equatoria, Bahr El Ghazal, Kordofan (= Nuba Mountains) and Upper Nile Province, and has been found on Upper Nile cattle at the Wadi Halfa quarantine, but is not es- tablished in Northern Province according to King (1926). The following are Sudan localities from which specimens, all from cattle unless otherwise noted, have been studied. si Bahr El Ghazal: Meshra el Req (SGC). Lau, Akot, Yirol, Tali Post, Kuru, Lake Nyubor, Peth, Raga, Khor Shamman, and Boro (SVS). Wau (domestic horses, sheep, donkeys, pigs, cattle; SVS, HH). Busseri (domestic horse; SVS). Njambo (buffalo; SGC). Raga (domestic goat, cattle; SGC). Aweng (domestic dog, cattle; SVS). Fanjak (SVS, HH). Guar and nearby areas in Galual_Nyang Forest (common on many individuals of tiang, buffalo, giraffe and roan antelope; also on warthog, hartebeest, and domestic dog; nymphs from spurfowl, Francolinus clappertoni; SVS, HH). 15 miles north of Tonj (tiang; SVS). Atet (Ttiang; SVS). Malek (nymph on Franco- linus sp.; svs). Lau River, 37 miles west of Yirol (nymph on cane rat; SVS). Nymphs are very numerous on tiang in the dry season and were also taken on buffalo and engorging on man. In the wet season, adults replace nymphs on wn animals. Upper Nile: Khor Atar and Taufikia (SGC). Duk Fadiat (cattle, HH; eae pig; SVS). Malakal (SVS, HH). Ler (SVS). Bor (SVS, HH). Darfur: ‘Idd el Ghanam (SGC). Radom (SVS). Blue Nile: Roseires (SGC). Pel oe Kordofan: Talodi (SVS). Cattle from Kordofan at the Wadi Halfa Quarantine (SCC). Khartoum: Abyssinian cattle at Khartoum (probably at the Quarantine Station) (SGC). Balfour (1906) reported specimens on trypanosome-infected cattle arriving from the south./ It will be noted that 12°N. is about the northern limit of this species. Rare isolated populations may exist slightly further north in the Sudan. DISTRIBUTION A. variegatum is distributed generally throughout the Ethiopian Faunal Region except in northern Sudan, most of Southwest Africa, much of Mozambique, and the entire Union of South Africa. The range includes mountainous Southwest Arabia and the tropical bont tick has become established in the Madagascan archipelago. It has also established itself from imported specimens in the West Indies and in the Cape Verde Islands. A record from Guatemala (Neumann 1899) has not been confirmed in later literature. “AFRICA" (Fabricius 1794) NORTH ATLANTIC OCEAN: CAPE VERDE ISLANDS (Tendeiro 1954X). WEST AFRICA: NIGERIA (Neumann 1899. Ziemann 1905. Simpson 1912A,B. Jonnston 1916. Connal and Coghill 1917. Robinson 1926. Pearse 1929. Beaton 1939. Mettam 1947,1948,1951. Unsworth 1949, 1952. Gambles 1951). FRENCH WEST AFRICA (As A. venustum: Koch 1844. As Ixodes elegans: Guerin.Meneville 1643. Neumann 1899, 1911. Joyeux 1915. Robinson 1926. Andre and Lamy 1931. Brumpt 1934. Lloverol, Philippe, and Adjovi 1942. Girard and Rousselot 1945. Rousselot 1951,1953B. Villiers 1955). TOGO (Ziemann 1905). SIERRA LEONE (Neumann 1899. Simpson 1913. Yorke and Blacklock 1915. Entomological Report 1916. Rou binson 1926). GAMBIA (Simpson 1911. Robinson 1926). GOLD COAST (Simpson 1914. Macfie 1915. Beal 1920. Robinson 1926. Stewart ee Og 1933). PORTUGESE GUINEA (Howard 1908. Tendeiro 1947,1948,1949B, 1951A,C ,D,F ,1952A,C ,D,E,1953,1954. Bacelar 1950). BIJAGOS ISLANDS (Tendeiro 1953x). CENTRAL AFRICA: CAMEROONS (Ziemann 1912A. Warburton 1927. Joncheres 1934. Bardez 1934. Rageau 1951,1953A,B. Rousselot 1951,1953B. Unsworth 1952. Dezest 1953). RIO MUNI ("North Central Rio Muni": Hoogstraal collection). FRENCH EQUATORIAL AFRICA (Fiasson 1943B. Blanc, Brunneau, and Chabaud 1950A. Giroud 1951. Rousselot 1951,1953A,B). BELGIAN CONGO and RUANDA-URUNDI (Newstead, Dutton, and Todd 1907. Massey 1908. Roubaud and Van Saceghem 1916. Nuttall and Warburton 1916. Van Saceghem 1918. Seydel 1925. Robinson 1926. Schwetz 1927A,B,C ,1932,1933B,1934. Schouteden 1929. Bequaert 1930A,B,1931. Tonelli-Rondelli 1930A. Bouvier 1945. Giroud and Jadin 1950. Giroud 1951. Jadin and Giroud 1951. Schoenaers 1951A4,B. Rousselot 1951,1953B. Reference to Belgian Congo by Berge and Lennette 1953 should be French Equatorial Africa. ert and Robinson 1954. Santos Dias 1954D. Van Vaerenbergh 1954). EAST AFRICA: SUDAN (Balfour 1904,1906. King 1908,1911,1926. Hoogstraal 1952A,1954B). ETHIOPIA (Pavesi 1884A. Neumann 1899,1902B,1911,1922. Robin. son 1926. Stella 1935A,1939A,B,1940. Roetti 1939. Charters 1946. D'Ienazio and Mira 1949. Hoogstraal 1954C). ERITREA (Franchini 1929D,E. Tonelli-Rondelli 1930A. Niro 1935. Stella 1938A,1939A, 1940. Ferro-Luzzi 1948). FRENCH SOMALILAND (Robinson 1926. Stel la 1940. Hoogstraal 1953D). ITALIAN SOMALILAND (Franchini 19264, 1927,1929C. Niro 1935. Stella 1938A,1939A,1940). KENYA (Neave 1912. Neumann 1922. Anderson 1924A,B. Robin. son 1926. Daubney 1927,1930A,B,1933,1934,1936B. Walker 1927, 1929. Tonelli-Rondelli 1930A. Lewis 1931A,B,C ,1932A,B,1934, 1939A,B. Daubney and Hudson 1931A,B,1934. Roberts 1935. Love— ridge 1936A. Fotheringham and Lewis 1937. Mulligan 1938. Dick and Lewis 1947. Weber 1948. White 1949. Binns 1951. van Someren 1951. Worsley 1952. Wilson 1953. Wiley 1953. Hammond 1954. See also IDENTIFICATION under A. pomposum, p. 245). a COs UGANDA (A. Theiler 1910A. Bruce et al 1911. Neave 1912. Neumann 1922. Robinson 1926. Richardson 1930. Mettam 1932,1933. Carmichael 1934. Mettam and Carmichael 1936. Wilson 1948A,B,C, 19500 ,1953. Clifford 1954. Hoogstraal 1954C. Taylor 1954). TANGANYIKA (Gerst&cker 1873. Neumann 1911. Neave 1912. Morstatt 1913. Jarvis 1918. Robinson 1926. Moreau 1933. Cornell 1936. ery ray 1941B. Beakbane and Wilde 1949. Wilson 1953. Smith ESD ic SOUTHERN AFRICA: ANGOLA (Neumann 1899,1911. Santos Dias 1950B. Sousa Dias 1950). MOZAMBIQUE (Karsch 1878. Howard 1908. Neumann 1911. Robinson 1926. Theiler 1943B. Santos Dias 1947A,19492, 1950B,1954H,1955A. Bacelar 1950. Wilson 1953). NORTHERN RHODESIA (Neave 1912. Robinson 1926. Morris 1933, 1935 ,1937,1938,1939,1940. LeRoux 1934,1937,1947. Matthysse 1954. Theiler and Robinson 1954). SOUTHERN RHODESIA (Koch 1903. Sin clair 1916. Jarvis 1918. Jack 1921,1928,1937,1942). NYASALAND (Old 1909. Neave 1912. De Meza 1918A,B. Robinson 1926. Wilson 1943 ,1946,1950B). UNION OF SOUTH AFRICA: Absent (Alexander 1931). “Rarely present" (Theiler 1943B). Dr. Theiler (1950 correspondence) states that A. variegatum is actually absent from the Union and from SOUTIL WEST AFRICA. See REMARKS below. Early literature records for this species in the Union of South Africa are: Howard 1908, Galli- Valerio 1909, Moore 1912, Bedford 1920, Curson 1928, Cooley 1934, Bedford and Graf 1939.7 OUTLYING ISLANDS: ZANZIBAR (Neumann 1899,1911. Neave 1912. Aders 1917). MAURITIUS (Neumann 1899,1911. De Charmoy 1914,1915. Robinson 1926. Moutia and Mamet 194.7). MADAGASCAR (Neumann 1899, 1911. Joyeux 1915. Robinson 1926. PBiick 1935,1948A,C,1949. Buck and Metzcer 1949. Millot 1948. Zumpt 1950B. Courdurier, Blick and Quesnel 1952. Hoogstraal 1953E). REUNION (Neumann 1899. Millot 1948. Gillard 1949). COMORES GROUP (Millot 1948). ARABIA: YEMEN (Franchini 1930. Girolemi 1952. Mount 1953. Sanborn and Hoogstraal 1953. Hoogstraal ms.). "SOUTHERN ARABIA" (Hoogstraal 195/C). = 265 = IMPORTED SPECIMENS: Records in the literature for EGYPT (Guerin-Meneville 10201843), one of the type localities of synon. ymous Ixodes elegans) should be discounted. Although A. variega- tum frequently arrives at the Cairo abattoir on cattle from the Sudan and from other areas of East Africa, the species has never become established here (Hoogstraal 1952A). Extremely few fe males are found by the time cattle reach Cairo. A. variegatum has become established and is a serious problem in the (St. Kitts, Guadeloupe, Antigua). As early as 1895, Barber wrote an account, both pleasant and critical, of the ravages of "the gold tick”, A. variegatum (= Hyalomma venustum) in Antigua. See also: Neumann ; ) Ticks in the West Indies (1914), Ford (1919), Saunders (1914A,B,1915,1919), Senevet (1938), and Mauze and Montigny (1954). It appears that A. varie. atum has been found in GUATEMALA (Neumann 1899,1911), but There [e been no subsequent reports of its presence there. There has been some question about West Indies records among American work. ers who have not visited these islands. Numerous specimens from West Indies may be seen in British Museum (Natural History) col. lections. Three males are stated to have been found on a dog in south. western FRANCE (Lamontellerie 1954). MISCELLANEOUS: The distributional map of Tendeiro (1947) which includes Egypt and Sinai and omits the Sudan and parts of Ethiopia, Eritrea, and the Somalilands, should be modified. TUNIS has been listed as a collecting locality based on spec imens labelled from a hedgehog on Djerba Island collected by A. Weiss (Galli-Valerio 1911A). Colas-Belcour and Rageau (1951), with ample reason, consider this record doubtful. It may be based on misidentification of an immature Hyalomma sp. HOSTS Where it occurs, A. variegatum is often the most common tick on cattle. Its incidence on Stier domestic animals varies locally but is usually less than on cattle. Among wild animals, the buf- falo and numerous kinds of antelopes are important hosts. Other = 200'= wild animals are either rather seldom infested by adults or those that more frequently harbor them, such as the rhinoceros, general. ly are not mumerous in nature. Carnivores are only exceptionally attacked. Man is rarely utilized as a host by adults, though nymphs attach more frequently and larvae are sometimes serious pests. Our knowledge of the host preference of immature stages is fragmentary. Nymphs feed on moderate size to large animals including all domestic animals and larvae attack mostly birds and small mammals from the size of hares to goats. Adult Hosts Domestic animals: Cattle (Practically every reference in the DISTRIBUTION section above pertains to parasitism of cattle by A. variegatum and these need not be repeated here. Selected refer— ences to parasitism of other domestic animals are presented below inasmich as many phases of these relationships are much less obs. cure than those with cattle). Camels (Robinson 1926%. Hoogstraal, ms.). Sheep (Robinson 1926, Schwetz 1927, Daubney 1930A, Daubney and Hudson 1931A,B,1934, Lewis 19310 ,1932B,1934, Tendeiro 1948, Rousselot 1951, Sudan records above). Goats (Robinson 1926, Schwetz 192%, Lewis 1934, Beaton 1939**, Tendeiro 1948, Sudan records Bboye). Horses (Simpson 1911, Robinson 1926, Schwetz 1927B, *Hosts listed by Robinson (1926) are based chiefly on the extensive data in the Nuttall collection now in British Museum (Natural History), where it is available to those who would make a further study of host—- relationships. **Though pinpoint blemishes in the tanned skins of goats are attri- buted by the (Nigerian) trader to the bites of ticks, this animal has been found to remain uncommonly free of ticks, particularly in the dry season. In the rains, when all domestic animals become grossly infested if not hand dressed, the goat is usually only parasitized by ....... A. variegatum and then to any extent only in the hollow of the Peedi e€ clefts of the hoof, and in the perineum. These exceedingly tenacious parasites set up a local inflammation with pus formation due to infection of the wound by organisms of necrosis, particularly in the feet. Severe lameness may be caused, and virus diseases, e.g., heartwater may be trans. mitted. (Beaton 1939). = 267 =< Blick 1935,1948A,C, Gillard 1949, Rageau 1951, Rousselot 1951, Sudan records above). Donkeys (Robinson 1926, Tendeiro 1948, Rousselot 1951, Sudan records a ve). Dogs (Simpson 1912B, Robinson 1926, Tendeiro 1948, Sudan records ACHE BMNH collections contain four adults from a dog from Senegal). Cat (Robinson 1926). Pigs (Schwetz 1927A, Lloverol, Philippe and Adjovi 1942, Rousselot 1951, Sudan records above). Man; In Madagascar and French West Africa (Joyeux 1915). Antelopes: Reedbuck (Robinson 1926*, Weber 1948, Wilson 1950B, Santos Dias 1953B). Bushbuck (Robinson 1926). South African bush. buck (Santos Dias 1953B). Steinbuck (Lewis 1932A). Waterbucks, various (Robinson 1926, Tendeiro 1952C, Santos Dias 1953B). Roan antelope, various (King 1926, Lewis 1934, Sudan records above). Sable antelope (Robinson 1926, Wilson 1950B, Santos Dias 1953B). Hartebeest, various (Simpson 1914, Robinson 1926, Lewis 1934, Sudan records above). Kongoni (Robinson 1926). Tiang (Sudan records above). Eland, various (Robinson 1926, Bequaert 1930B,1931, Lewis 1934, Weber 1948, Wilson 1950C, Sudan records above). Nyala (Santos Dias 1953B). Duiker (Loveridge 1936A, Wilson 1950B, Sudan records above). Grant's gazelle (Lewis 1934). Bright's gazelle, Smith's long-snouted dikdik, and Roosevelt's duikerbok (Sudan records above). Oribi, various (Wilson 1950B, Santos Dias 1953B, Matthysse 1954, Sudan records above). Other mammals: Giraffe (common on many Bahr El Ghazal gi. raffes examined, records above). Zebra (Neumann 1911, Robinson 1926*, Lewis 1932A,1934, Weber 1948, Matthysse 1954). Black, or narrow~lipped rhinoceros (Karsch 1878, Neumann 1911, Robinson 1926, Lewis 1932A, Wilson 1951C). White, or square-lipped rhino_ ceros (Tonelli-Rondelli 1930A). Elephant (Robinson 1926). Buf. falo (King 1926, Robinson 1926, Richardson 1930, Bequaert 1930B, 1931, Fiasson 1943B, Wilson 1950C, Rageau 1951, Santos Dias 1953B, Sudan records Eber). Warthog (Massey 1908, Rabinson 1926, Santos Dias 1953B, Sudan records above). Bushpig (Tonelli-Rondelli 1930A, Matthysse 1954, Sudan records above). Antbear (Lewis 1932A). Leopard (captive) and hares (Tendeiro 1947,1952C). Cheetah (Lewis 1934). Lion (Theiler, unpublished). are a YES COE Pe Po I WB ok 2 RR oa Sa NE eh a *Hosts listed by Robinson (1926) are based chiefly on the extensive data in the Nuttall collection now in British Museum (Natural History), where it is available to those who would make a further study of host-relationships. = 20ot= Birds: Spurwing geese (tick identification questionable: Bedford 1932B). Gray hornbill (Lophoceros n. nasutus) and ground hornbill (Bucorvus abyssinicus), domestic and wild chickens, spur- fowl, guineafowl (Tendeiro 7,1948,19520). Snake: Bitis arietans (Neumann 1911). Snail: A curious case, said to be parasitism by A. variegatum on a snail, Limicolaria adansoni Pfr., in Senegal, has been reported by Neumann (1911) and by Andre and Lamy (1931). This record bears further investigation. Nymphal Hosts Nymphs feed on a great variety of mammals of medium and large size, including occasionally man. Birds are frequently parasitized but reptiles are rarely attacked. Man: In French West Africa, nymphs do not attack man so frequently as do larvae (Joyeux 1915). In the Sudan a single nymph was taken feeding on man in Equatoria and several in Bahr El Ghazal Province. Domestic animals: Nymphs are common on domestic stock in Kenya (Lewis 1934). Survey of our Equatoria and Bahr El Ghazal Province records above shows a small number of nymphs on cattle and often many on goats, especially during the dry season. Few were found on other domestic animals, pigs, horses, dogs, and sheep. Sudan Government collections contain nymphs from cattle, sheep, goats, dogs, pigs, donkeys, and horses. Tendeiro (1948) listed cattle and goats as nymphal hosts. Among a collection from Madagascar, Zumpt (1950B) found nymphs from cattle and dogs. Theiler (correspondence) has specimens from elsewhere in Africa from the same animals as well as from camels and domestic cats. Poultry has been listed as a nymphal host by Wilson (1950B), Tendeiro (1948), Hoogstraal (1953E), and we found this stage on chickens and turkeys in the Sudan (records above). Daubney and Hudson (19314,B,1934) referred to the comparative rarity of im mature stages on sheep in Kenya. Fiasson (1943B) noted immature specimens on sheep at Libreville. = 209%— Wild mammals: Insectivores: Hedgehog (Wilson 1950B, Sudan record above). Madagascar tenrec (Hoogstraal 1953E). Carnivores: Jackal (Neumann 1902B, Lewis 1934, Matthysse 1954, Sudan records above). Cheetah (Wilson 1950B). Long-eared fox, Otocyon megalotis (Lewis 1934). Spotted hyena and mongoose (Sudan recente above). Lagomorphs: Hares (Wilson 1950B, Matthysse 1954, Sudan record =o . Rodents: Cane rat (Wilson 1950B, Bahr El Ghazal record above). Ground squirrel (Sudan record above). Antelopes: Harte. beest, Thomson's gazelle, klipspringer (Lewis 1934). Pec tese defassa waterbuck (Tendeiro 1947). Waterbuck in Uganda and “forest antelope from Rio Muni (HH collection). Oribi (Hoogstraal 1954C, Sudan record above). Sudan records are from Bright's gazelle, Roosevelt's duikerbok, Smith's long-snouted dikdik, oribi, common eland, Roosevelt's hartebeest, tiang, and buffalo. Nymphs are especially numerous on tiang in Bahr El Ghazal Province during the dry season. In Northern Rhodesia, Matthysse (1954) found all stages on zebras. Wild birds; Coucals, various (Theiler, unpublished. Hoogstraal 1953E). Ground hornbill, Bucorvus abyssinicus (Warburton 1927). Raven (Hoogstraal 1954C). Spurfowl (Pternistis sp. or Francolinus sp.) (Lewis 1934, various Sudan records above). Long-legged bus— tard and hooded vulture (Theiler, unpublished). Guinea fowl (Lewis 1934 and Sudan records above). Abdim's stork, lesser bus. tards, tchagra shrike (Sudan records above). An undetermined passerine bird (arveola) (Tendeiro 1952C). For domestic birds, see Domestic animals above. Reptiles: Chameleon in Madagascar (Hoogstraal 1953E). Larval Hosts Man has been listed as a larval host by Wilson (1950B). Larvae commonly attack man at the beginning of the dry season in Upper Guinea, French West Africa (Joyeux 1915). In Cameroons they are serious pests of man (Rageau 1953B) and attach on the legs and about the belt. Ziemann's (1912B) mention of being badly bitten by tick larvae in the Cameroons may refer to this species. Ac. cording to De Meza (1918A), in Nyasaland larvae are serious pests of people working about cattle. Larvae just visible to the eye burrow under the skin of human legs and cause severe irritation that may be associated with rash and pus if the ticks are numerous. ee Oe Birds: Spurfowl (Pternistis sp. or Francolinus sp.) (Lewis 1932B). Helmeted guinea fowl (Lewis 1934). "Rural chickens" (galinha do mato) (Tendeiro 1952C). Lesser bustard (Sudan record ISR E 7 Mammals: Hare (Lewis 1934). Reedbuck (Lewis 1931C). Zebra and jackal (Matthysse 1954). Genet, jackal, and mongoose (Sudan records above). Domestic goat (Lewis 1934, Sudan record above) and cattle (Lewis 1931C, Sudan record above). Domestic dog in Madagascar (Hoogstraal 1953E). Stage not stated but probably immature Hedgehog (Atelerix spiculus) and gerbil (Taterillus gracilis angelus) (Pearse 1929). Hedgehog (Atelerix albiventris Wagner) - adansoni Roch.) and white-tailed mongoose (Ichneumia albi- auda) (Rousselot 1951). BIOLOGY Life Cycle This three-host tick is frequently the most common cattle parasite within its range. Hosts of each stage are listed above. Most workers have experienced difficulty in rearing A. variegatum but J. B. Walker states (correspondence that she finds this species quite easy to rear. With nonfeeding ticks maintained at from 25°C. to 27°C., the minimum periods for the life cycle are as follows: eee PERIOD DAYS (Minimum) Walker Lewis Lewis 1932A 19398 Preoviposition 12 TS Clee) Oviposition to hatching 53 86 (19.26% ) Larval prefeeding period ia tl Larva feeds 5 7 EES Premolting period 14 22 (25.27 ) Nymphal prefeeding period U 7 Nymph feeds 5 7 oul) Premolting period 19 2b (2276) Adult prefeeding period 7 ; Adult (Female) feeds 12 10 a= 22 Total 141 195 = The prefeeding periods in the above table are arbitrary and in nature may be shorter than the fisures indicate. Walker fed larvae and nymphs on a rabbit and adults on aram. Lewis (1932A) used hares, chickens, and sheep as hosts. Nuttall (1915) recorded attachment periods twice as long as Lewis! and stated that males may remain on the host for fron four to eight months, and may even die there. oft 30°C., larvae energe from egss after sixteen to 31 days; at 15 C. larvae fail to hatch. On the other hand, Mettam (1933) reported that in his Usanda laboratory "times occupied during feeding, moltinc, etc. are much shorter than the one obtained by ....... Lewis”, but these observations apply only so far as the nymphal stace. Larvae feeding on man drop off the host after twelve hours (Joyeux 1915). Under laboratory conditions, the longevity of A. variegatwa (2? unfed adults — HH) is 732 days (Lewis 19393). In those parts of its range with but one rainy season annually, the tropical bont tick has only a single generation a year, as reported for Nyasaland by Wilson (19508) and for Vorthern Rhodesia by Matthysse (1954). In Kenya and Uganda, with two rainy periods each year, multiplication is faster and two or three generations may breed during a twelve months' period (Wilson 1953). Females engorge and oviposit during the wet months, larvae engorge early in the dry season, and nymphs live through the dry season. The periods of preoviposition and embryonic development in nature should be more carefully investigated. A delay or dia pause phenomenon of some three months or more for these combined periods in the rainy season appears likely. Ecology The typical seasonal cycle, as explained for Northern Rhodesia by Matthysse (1954), applies to the Sudan and other single rainy season areas of Africa. Details for other areas with two rainy seasons are not certain, and there appears to be more overalppins of different stages in such situations. Adults appear towards the end of the dry season, first males and then females. Populations increase in nunbers and remain high through the rainy season and decrease rapidly in the dry season, although a few specimens may be found even then. Larvae and nymphs gradually become more numerous in the dry season, and while some nymphs are found during the rains they are scarce. Adults and nymphs are most common on the udders, scrotun, flanks, dewlap, and brisket; larvae feed on the ears and head of the host (Wilson 19483,1949). Beakbane and Wilde (1949) also noted adults on the perineum and indicated means of control with respect to the feeding sites of ticks infesting cattle. In Cameroons, larvae of this tick have been observed in im. mense numbers on tall herbare along paths, waiting for a suitable host to pass (Razeau 1953B). Similar, vivid remarks by Ziemann (19128) for Cameroons suggest that his observations may also have referred to A. variegatun, "The fully fed female of A. variezatun works her way into the soil to lay her esrs, and unfed adults are freouently seen, waiting for a passing host on the foliase of bushes three or four feet high” (Lewis 1932). Eis. = A. variegatum has been found from sea level to 8500 feet elevation. in the Yemen (Arabia), this species is common on cattle in well vegetated valleys and hillsides between 2500 and 5000 feet elevation, but absent in deserts at lower elevations and rare in more barren higher elevations (Hoogstraal, ms.). Franchini's (1930) record of A. variegatum from Hodeida, on Yemen's coastal plain, is due either to erroneous locality labels or represents specimens from highland cattle arriving for slaugh- ter. This species is common at Asmara, Eritrea, 7500 feet eleva. tion (HH collecting), but Schoenaers (1951B) states that it does not occur over 2000 meters (6500 feet) elevation in Ruanda_Urundi. In Fast and Central Africa, Wilson (1953) has nicely defined the presence of two very distinct ecological relationships between ticks and cattle. One of these, the R. appendiculatus_A. variecatum association, occurs in areas with cnet well above 25 inches per annun (and is of considerable importance in relation to East Coast fever and heartwater transmitted to cattle by the respective ticks). The second, the R. pravus (= R. neavi)-A. gemma association, occurs where rainfall very seldom exceeds twenty or 25 inches per annum (and is of negligible veterinary importance). For a summary of Wilson's second association, see R. pravus (page 681). The distribution of the R. appendiculatus-A. a asso— ciation corresponds to what veterinarians previously referred to as "dirty areas" (i.e. East Coast fever endemic areas). As stated above, rainfall here is at least 25 inches annually, usually well above this figure, and falls below this level only once in every twenty or 25 years. This association includes the highlands of Kenya and Tanganyika, a 25 to thirty mile belt bordering Lake Victoria in Kenya, Ucanda, and Tanganyika, and continues down the Rift Valley in the country adjacent to Lake Tanganyika and Lake Nyasa, and (a short distance) into Mozambique. It also includes the humid seacoast plains, which are only a few miles wide in Kenya but much wider in Tanganyika. Within drier areas (i.e. those of the first association) are isolated islands in the “rain shadow" of hills and mountains where higher precipitation results in more dense vegetation than that of the surrounding plains. In these islands, the PR. appendiculatus-A. variesatun association persists. The soil and vegetation on which this association oc- curs vary tremendously with slope of terrain, altitude, underlying rock formation, and temperature. The single common factor in their Sli ecology is relatively high rainfall. Within their areas of dis. tribution, these ticks vary in relative prevalence and density, due in part to approaching thresholds or extremes within the range of tolerance. A very definite line of demarcation, un. effected by the movement of cattle along trade routes, exists between these two zones. Huts of pastoral peoples in which cattle and other domestic animals frequently rest harbor the tropical bont tick. Smith (1955) noted its presence in human habitations and a similar situation may have accounted for Robinson's (1912) remarks con. cerning parasitism on children by the so called variety nocens. The red-billed oxpecker or tickbird, Buphagus e. erythro- rhynchus (Stanley), which attends all the larger herbivores except the elephant and the hippopotamus, has been shown by Moreau (1933) to be a predator of some importance on A. variega. tum and other economically important ticks. Of the 58 ticktbinis ‘examined in Tanganyika, 186 specimens of A. variegatum were found in the stomach contents of sixteen; the number of ticks per stom. ach ranged from one to 109. In Kenya, van Someren (1951) took specimens of the tropical bont tick from stomachs of a few of the same birds that he ex. amined. He also found unidentified ticks in Tanganyika birds, B. a. africanus. van Someren's interesting biological study of The red-billed oxpecker includes observations on the birds! habit of irritating sores on domestic animals. Buphagus erythrorhynchus and B. africanus subspp. range through African savannahs requently are seen clinging to the flanks or legs of domestic and wild animals. Lang (1924) noted the tick bird acting as a sentinel for elands. Loveridge (1928) reported specifically unidentified ticks in stomachs of Tanganyika tick birds and Dr. J. P. Chapin found ticks in their stomachs in the Congo and in Kenya (Bequaert 1930B). I am told that the best account of tick-eating habits of these birds is quoted in Banner- man's Birds of Tropical West Africa (Volume 6, page 105). Assertions that the white heron, or cattle egret, Bubulcus spp., is a tickivore are not supported by evidence. Dr. Chapin = 275 = found no ticks in cattle egret stomachs and their actions do not indicate that they commonly feed on these parasites (Bequaert 19302, Plowes 1950). In Southern Rhodesia, "Egret" (1938) reported, these birds may pick grasshoppers off grazing cattle put do not search for ticks on them. Colleasues and I have found no ticks in stomachs of many cattle egrets examined in Africa and elsewhere and Kirkpatrick (1925) found none in stomachs of many Egyptian birds of this type. REMARKS Special attention is called to the taxonomic and biological status of heavily punctate specimens of A. variegatum, discussed under A. pomposun (page 245). Remarks by Theiler (1951 correspondence) are of value in understanding distributional factors of this species and in il. lustrating the care that mist be taken in evaluating older lit. erature. She writes: "..... statements made (Theiler 1943B), to the effect that this species is found in the Union of South Africa, were based on records in the literature, i.e. before we hed studied the South African tick survey material ..... Our abundant material did not produce one specimen from anywhere in the Union or in SouthWest Africa. Bedford's statements are based on incorrectly identified nymphal material. Robinson's record in the Monograph, I take to be a record of an introduc— tion into the country, which certainly has not been able to maintain itself. Nor does A. variezatum seem to be as widely spread in Southern "hodesia as one gatnered from some of the earlier workers. On the contrary, A. hebraeum seems to be more prevalent than was first thought. When reading reports of ear— lier workers (18961906), one must bear in mind that they are still reporting on a period during which cattle had been, or were being, introduced from East Africa and from Madagascar, bringing A. variegatum and Boophilus fallax with them. Possi- bly some of these ticks come into areas in which they could maintain themselves for a year or two, or possibly even longer." Brumpt (1922,1934) reported on gynandromorphic specimens of A. variegatum. Malformed specimens have been noted by Santos Dias TiS eSED, and Tendeiro (1951F). Certain aspects of the chitin = 210 = ization of the exoskeleton and gut and of the musculature of this species have been discussed by Ruser (1933). A. variegatun has been used to illustrate the double oblique-striation sf tick musculature by Kruger (1935). Variations in scutal ornanentation among specimens from a restricted area were illustrated by Tem deiro (1949B,1951F). The eye structure and related sense organs were described and illustrated by Gossel (1935). DISEASE RELATIONS MAN and ANIMALS: Q fever (Coxiella burnetii). Larvae and nymphs commonly attack man under Local conditions and may cause severe irritation. CATTLE: Heartwater (Rickettsia ruminantiun). Bovine lymphar citis, large septic sores, and severe inflammation of mammae. Possibly transmits a fungus, Cryptococcus (= Actynomyces) farci_ minosus. A. variezatum has been mentioned in connection with bovine rickettsiosis (Rickettsia bovis), but the tick's role does not appear to have been defined. This tick does not transmit East Coast fever (Theileria parva). SHEEP: Nairobi sheep disease (virus). Heartwater (R. runinantiun). GOATS; Heartwater (R. ruminantiun). Severe secondary in. fection and lameness. PIGS: Possibly a vector of porcine piroplasmosis (Babesia trautmanni). WILD ANIMAL INJURY: In the Sudan I removed a live nymph from a thish abscess that was almost completely overgrown by the skin of the jackal host. = ZI IDENTIFICATION Males: There is no question of identity of this important species in the Sudan tick fauna or throughout most of its range. The combination of characters including hemispherical, orbited eyes, long lateral grooves, entirely black festoons, and paucity of large scutal punctations easily distinguishes A. variezatun. The scutal ornamentation is as illustrated (Figure 92) except that in about five percent of specimens an additional coppery spot may be found just inside (not outside) of the lateral grooves at the level of the scutal midlength.. Specimens with these spots are readily distinguished froin A. lepidum and A. pomposum by their dark festoons and lack of large scutal puncta- tions. Note: According to Jack (1942) males from eastern parts of Southern Rhodesia approach A. pomposun in that they have coarse scutal punctation. No females were available for comparison. A few confusing specimens such as these from the Sudan, East Afri- ca, and Yemen (Arabia) have been observed (See IDENTIFICATION of A. pomposum, page 215). Females: This sex is sometimes more difficult to identify with certainty than the male. Hemispherical, orbited eyes are also found in A. lepidum and A. pomposun. In the latter, the very rugose, broad, short scutum easily separates it. The pos— terior margin of A. variegatum is comparatively sore broadly rounded than that of A. lepidum. One may have considerable dif- ficulty in deciding whether a female scutum is widely or narrow— ly pointed posteriorly. There appears to be some variation in this character, but a thorough study has been impossible be- cause, in all available collections, no more than eighty female specimens of A. lepidun have been seen. No large series from any single area has been represented and until such time as more material comes to hand it is preferable to hold the study of this feature in abeyance. SZ ioe APONOMMA INTRODUCTION African Aponomma ticks, small, eyeless parasites of large snakes and of monitor lizards (Varanus spp.), are seldom rep- resented in collections. They are markedly host-specific and rarely feed on animals other than their normal hosts. Originally considered as Amblyomma, species of the genus Aponomma are now treated as a separate generic offshoot from or omma. In this genus the eyeless condition is considered by some to have resulted from disuse since certain members feed under the host's scales. Parallel instances also occur among those Amblyomma ticks that have indistinct or vestigial eyes and that parasitize scaly hosts. This interpretation might hold for those Aponomma species that attach below snakes? scales but may hardly account for the eyeless condition of others attacking only scaleless lizards. Possibly Aponomma ticks became adapted to lizards as a concomitant to the c= of vision. The biology of aponommas is poorly known. Adults and nymphs are frequently found on the same host. The nomenclature of the few African species has been confused until the recent works of Theiler (1945A,B). Most African spec- imens are now easily identified to species. More recently, Santos Dias (1955C) has redescribed the type material of A. ochraceum Neumann, 1901, from Tanganyika and Zanzibar, and of A. fraudigerum Schulze, 1935, from a host, Varanus griseus, presumed to have come from North Africa.* WV. fae ranges over the entire northern Sahara region, reach. ing the Mediterranean only in southern Tunisia, Libya, and Egypt, and extends to the Central Provinces of India (Loveridge, corres pondenge). The source of the type material of A. fraudigerun, Fuhlsbuttelterrariun, probably refers to the grounds of an animal dealer at Fuhlsbuttel, a suburb of Hamburg, and the collector, Karl Peter, is probably not the famous African explorer but rather the animal dealer (Theiler, correspondence). - 279 - Aponomma species are more numerous in the Oriental Region than in Africa but their taxonomic status is uncertain. For Indochina, Toumanoff (1944) lists three species, Anastos (1950) records four from Indonesia, and Roberts (1953) studied nine from Australia. None occur in Europe and Neumann's (1899) lo cality label for A. laeve from Patagonia was possibly incorrect. In the Americas, four species have been described by Schulze (1932A,1936E,1941X) but specimens available in collections throughout the world are few indeed. Specific concepts in this genus are much in need of careful study. KEY TO SUDAN SPECIES OF APONOMMA MALES AND FEMALES Shiny dark brown with green or copper markings on scutum. (Usually on Var anus 1igards ec Without caudal appendage. Hypostome dentition 4/4 (see Note eae Palpal basal segment without ventral bristle— bearing protuberance on inner margin. Adanal shields without posterior spur..............B. ANNULATUS Figures 1 2. Hypostome dentition 3/3 (rarely 3.5/ 3.5). Palpal basal segment with ventral, inner bristle-bearing pro tuberance. Adanal shields with long, narrow internal spur usually extending beyOnde bod ya mar etnies iclslee ere cleicieleleleloleleicicleteclele sielerere Dom DC OLOALTUS Figures Hypostome 4/4. Palpal basal segment without ventral, inner bristle—bearing protuberance. Adanal shields with inner margin hardly extended if at all and not reaching body margin.......esscccecseeesebe. MICROPLUS** Figures 112 and 113 *Preserved specimens must be entirely free of surface sheen from liquid preservatives before identification can be attempted. The caudal appendage of c'B. decoloratus is very variable in size and the palpal basal bristle-bearing protuberance of either sex can be seen only when the mouthparts are absolutely clean; the bristles are frequently broken off. Turning the specimen at an angle to the light may be necessary to see this character as well as the groove of coxa I. *%B, microplus does not occur in the Sudan but there is some likeli-_ hood that 1t may reach the Sudan within a few years. The inclusion of B. microplus in this key makes it serviceable for all known species, according to contemporary concepts of the genus. = 295s FEMALES 1. Coxa I with shallow, rounded emar- gination separating internal and external spur. Palpal basal segment and hypostome as| in male... 2 ..ccccccvsiscccccccvocesbs ANNULAIUS Figures 106, 107, and 117 Coxa I with deep elongate “inverted VE shape cleht dividimo mS OUr Siete crise c16/e/e oles e.aleleise'sisiele'elelelsvelseleiela = 2. Hypostome 3/3 (rarely 3.5/3.5). Palpal basal segment with ventral, inner bristle-bearing protuberance.........seeeeeBe DECOLORATUS Figures 110, 111, and 116 Hypostome 4/4. Palpal basal segment ventrally with inner margin concave and lacking bristle—bearing protw DETEMIC Ciayatetera/stsleisie)6 © cis'viel else eleisie sie cles elvis eeivisicieienioes De PLO RObiUs Figures 114, 115, Note The number of longitudinal files of denticles on each side of the hypostome is expressed by a formula such as 4/4 or 3/3. The number of these files is constant in most species of ticks, but the number of denticles in each file may be more variable. The formula 3.5/3.5 indicates that there are three complete and one shorter file of few denticles anteriorly on each side of the hypostome. Following Minning, the length/width ratio of the female scutum has usually been considered to be an important diagnostic character. Examination of large numbers of field-collected specimens from Afri- ca and elsewhere reveals wide variation in this feature within each species and no valid data for diagnostic purposes. =, 296T= Figures 104 and 105, 3, dorsal and ventral views Figures 106 and 107, 9, dorsal and ventral views BOOPHILUS ANNULATUS an specimens PLATE XXXIV - 297 - BOOPHILUS ANNULATUS (Say, 1821) (= B. CONGOLENSIS Minning, 1934). L N ¢@ @ 3 i: al 1 99°39 TO Ome 10 1 11 (Figures 104 to 107 and 117) THE TEXAS FEVER TICK EQUATORIA PROVINCE RECORDS Meridi Taurotr derbianus gigas Feb (SVS) Nagichot domestic cattle Dec Katire domestic cattle Jan Torit domestic cattle Nov Tombe domestic cattle Mar ee Muni domestic cattle Mar (SVS) Terakeka § domestic cattle Mar (SVS) Yambio domestic cattle Jan Meridi domestic cattle Feb (SVS) were collected at Wau from donkeys: 2 abl 4 ab 6 11 6 2 7 29 18 2 34 19 3 DISTRIBUTION IN THE SUDAN In addition to the above Equatoria Province specimens, the following others have been seen, all from cattle except those which Locality Province Date Malakal Upper Nile Jan 1952 Ler Upper Nile Jan 1952 Bor Upper Nile Jan 1952 Akobo Upper Nile Mar 1914 Lau Bahr El Ghazal Jan 1914 Wau Bahr El Ghazal Sep 1950 Wau Bahr El Ghazal Feb 1953 Wau Bahr El Ghazal Feb 1953 Wau Bahr El Ghazal Jul 1953 Fanjak Bahr El Ghazal Jan 1953 Fanjak Bahr El Ghazal Mar 1953 Yirol Bahr El Ghazal Apr 1954 Talodi Kordof an Dec 1926 Gollector A. A. P. Cc. Je E. He Webb (SGC) T. M. Reid Hoogstraal M. Dahab (SVS) W. Polden (SVS) Durran (SVS) J. Soutar (SGC) (svs) Gordon College Collection E. T. M. Reid H. Hoogstraal E. T. M. Reid E. T. M. Reid J. W. Gowland Wadi Halfa Quarantine Station from Kordofan cattle = 2967— (s7s) The Sudan distribution picture is a curious one. Although we have a fair number of collections from cattle in that half of Equatoria Province that is east of the Nile, none contain spec. imens of B. annulatus except two from high altitudes (lo from Katire, 3500 feet elevation, lo from Nagichot, 6500 feet eleva. tion) and one from Torit, at the general two thousand foot level of the plains in this area. Yet collections from the southern part of Upper Nile Province, which is just north of the east bank of Equatoria Province, include a good number of specimens. Cli- matic, faunal, or floral differences can hardly account for the rarity of B. annulatus in Torit and Juba districts. The most easterly record, Akobo, Upper Nile Province, is on the Ethiopian frontier (P47IN., 33°OL'E.). On the west bank of the Nile, this species is here recorded from several localities to as far north as Talodi, Kordofan Province (10°37!N.). Some specimens reach the Halfa Quarantine station in Northern Province but Boophilus ticks have never been collected from indige- nous cattle in Northern Province. DISTRIBUTION Thus far we know B. annulatus only from West Africa, Central Africa, and certain parts of southern Sudan near the periphery of East African biotic Provinces. As early as 1905, Donitz recognized that this tick inhabited only "tropical Africa™ and was absent in eastern and southern Africa. He tentatively applied the name B. australis Fuller to it, although he stated clearly that he could not difterentiate African material from descriptions of American B. annulatus. Unfortunately, he had no American specimens for comparison for it appears that this perspicacious student might otherwise have saved future generations much misunderstanding. Instead, this species subsequently has been either completely overlooked or subjected to numerous ambiguous remarks and names. Many African specimens undoubtedly have been identified as B. decoloratus, and earlier workers who have recognized specimens as different from B. decoloratus have referred to them by various names. The actual species with which various investigators were dealing cannot be determined without seeing their specimens. For instance, Nuttall's lots identified as B. australis, which have Sao te been examined in British Museum (Natural History), contain both B. decoloratus and B. annulatus. B. annulatus is a North American cattle tick that one may assume originally parasitized deer and buffalo. It also has been introduced into the Mediterranean basin. In North Africa and the Near East, it frequently is referred to as B. calcaratus (Birula, 1894), to which Minning appended several subspecies. These all appear to be the same as American and African populations of B. annulatus. The name also has been used by students of the Oriental fauna but examination of pertinent specimen material is indicated to establish the validity of these identifications. Records presented below are the only ones from Africa that are known to pertain to B. annulatus, with a few additional, an. notated references that might be pertinent. Quite possibly other isolated populations are maintaining themselves outside of the presently recognized range of the Texas fever tick in the Ethiopian Faunal Region, having been introduced on cattle from West or North Africa, the Near East, or North America. WEST AFRICA: NIGERIA (Hoogstraal 19540). LIBERIA (Specimens from cattle at Harbel, Firestone Rubber Company Plantation, H. A. Beatty legit; MCZ collections). SIERRA LEONE (Hoogstraal 1954C). PORT GUINEA: B. (= Margaropus) annulatus listed by Monteiro a Costa (1926) and Santtanna Barreto (1929); quoted by Tendeiro (19514), but not subsequently repeated in faunal lists by this author although it would not be surprising to find this tick here./ NOTE: Records below are for "B. congolensis”. CENTRAL AFRICA: FRENCH EQUATORIAL AFRICA (Minning 1934. Rousselot 19558. Theiler and Robinson 1954). CAMEROONS (Rageau 1953A,B). BELGIAN CONGO (Theiler and Robinson 1954. Minning 1934 stated, apparently as a guess, that the B. annulatus calcaratus specimens of Newstead, Dutton, and Todd 1907, from Coquilhatville, are "B. congolensis"). EAST AFRICA: SUDAN (Hoogstraal 1954B,C. Balfour 1911F referred to "B. australis™ in the Sudan, but since he also stated that B. decoloratus 1S absent there his remarks are difficult to interpret). = 10,0 HOSTS Cattle (All references). Rarely giant eland and domestic donkey (Sudan records above). In America, other domestic animels, ere and buffalo have been reported as infrequent hosts (Cooley 1946 e BIOLOGY Life Cycle Like other boophilids, the Texas fever tick is a single host parasite. Its life cycle has not been studied in Africa, where it is not known to occur under cold conditions. After dropping from the host, the female commences oviposition in about three or four days but after twenty to forty days in winter (southern United States). The oviposition period ranges from eight or nine days in summer to 42 days in winter. The number of eggs average 1911, with a maximum of 3806. With abundant mois. ture, eggs may hatch in as little as from 17 to 21 days, but up to 44 days is more common. Winter incubation may require between five and six months. A few hours after hatching, larvae collect in masses at the tip of grass, and may remain alive from 49 to 159 days awaiting a host. Once an animal is found larvae prefer- entially attach to the legs, belly, or dewlap, but if numerous they are found everywhere on the body. The larval-nymphal molt occurs seven to twelve days later and nymphs molt to adults five to ten days afterwards. Females feed for from four to fourteen days, during which time they mate, and then drop to the ground, oviposit, and die. These data are extracted from the very com. plete work of Hunter and Hooker (1907) in the United States. Ecology Most African specimens are from collections containing more numerous B. decoloratus, with the exception that during the rainy season the numbers of B. annulatus have in some instances exceeded those of B. decoloratus. In Cameroons (Rageau 1953B), this tick is less common than B. decoloratus but both species are found to- gether. Our first suspicion, that this might be a species ac. = 201 = climated to humid West Africa, is negated by finding it in Sudan localities with long, severely hot, dry seasons. The Texas fever tick occurs in the Sudan in areas with from 800 mm. to 1500 m. annual rainfall. The ecology of B. annulatus is unstudied in Africa and the results of research on this subject are awaited with considerable interest. In North and Central America, B. annulatus is entirely a tropical and subtropical tick that dies out when introduced into the northern states. DISEASE RELATIONS Unstudied in Africa. B. annulatus in America is the famed vector of Texas fever of cattle (Babesia bigemina). REMARKS Minning (1934) considered West African populations to differ from New World B. annulatus and called them B. congolensis. Nw merous B. annulatus specimens from the United States and Gentral America have been examined in collections of British Museum (Natural History), Rocky Mountain Laboratory, Museum of Comparative Zo8logy, as well as Louisiana material kindly presented by Dr. F. C. Bishopp from United States Department of Agriculture collections. None of these can be distinguished from African ™"B. congolensis". The chief characters presented for differentiating males are the pointed outer spur of coxa I of “B. congolensis” and the blunt outer spur of coxa I of B. annulatus. In a few African specimens this spur is blunt, in many American specimens pointed. The American specimens in the Nuttall collections at British Museum (Natural History) are mostly blunt-spurred with few pointed-spur specimens, but those at the Rocky Mountain Laboratory in Montana are almost entirely pointed-spurred. Specimens from Louisiana in the present collection have pointed spurs. It is evident, therefore, that this character is a variable one with no diagnostic significance as to species. The shape of the eyes of African and American specimens is similar. Examination of other so called dif- ferentiating characters has also failed to reveal differences. a Oe ae Similarly, no points of differentiation may be detected be— tween female specimens from Africa and America. The chief diag- nostic characters proposed for these, arching of the eyes and shape of the distal margin of the third palpal segment, appear similar, as are all other morphological features of specimens from both continents. Minning stated that the scutum of Amer- ican B. annulatus bears hairs and the scutum of ™B. congolensis™ does not. Several African specimens with scutal hairs =a some American B. annulatus without scutal hairs are available, though they may have been rubbed off the latter. Theiler (1943B) has already observed that the relation of the position of the eye to scutal margin in Boophilus ticks is subject to variation according to degree of engorgement. As already stated (page 296) no differences between length/width ratio of the female scutum can be determined in field-collected material of each species. As stated above, it appears that there are no constant differences between American and African populations of B. annw latus. Likewise, it is impossible to differentiate between Amer— ican and African populations of B. annulatus, and specimens col. lected in North Africa, Southern Europe, and the Near East, which Minning (loc. cit.) referred to as B. schulzei and as B. calcara. tus subspp. = ¥ In their classic work on this parasite in America, Hunter and Hooker (1907) open by observing “It is safe to state that no more important problem than the eradication of ...... (B. annulatus) ...... confronts the farmers of any country. Not only the cattle raising industry but the whole economic condi- tion of a large section of country is affected™. The Texas fever tick is now completely eradicated from the United States except for periodic introductions from Mexico, where it still exists. At the turn of the century it ranged through Mexico and the sixteen southern states of the United States from the Atlantic to the Pacific. The history-making discovery by Smith and Kilbourne (1893) that Babesia a is the cause of Texas fever of cattle was followed by their finding that this tick is the vector. The life history and morphology of the tick have been reported handsomely by Curtice = 3037— (1892), Salmon and Stiles oe Hunter and Hooker (1907), and Hooker, Bishopp, and Wood (1912). The latter report contains a useful summary of the antecedent literature. IDENTIFICATION Males are easily identified in the African fauna. They have no caudal appendage on the posterior body margin and the inner margin of the adanal shields does not project posteriorly as a spine. Palpal segment I is without a ventral bristle—bearing protuberance. The hypostome formula is 4/4 and the denticles are noticeably finer than those of B. decoloratus. In size, males are about the same as those of B. decoloratus. Females are usually readily separated from B. decoloratus by absence of a deep gap between the inner and outer spurs of coxa I. A shallow concave emargination replaces this gap. There is no bristle-bearing protuberance on the internal ventral margin of palpal segment I; this margin is fairly elongate and mildly con. cave. The scutal margins anterior of the eyes are usually straight and parallel. The scutum is definitely longer than wide. The hypostome formula is as for the male. The color of this spe cies is usually paler than that of B. decoloratus. - 304 - Figures 108 and 109, &, dorsal and ventral views Figures 110 and 111, 9, dorsal and ventral views BOOPHILUS DECOLORATUS udan specimens PLATE XXXV = 3 05i— BOOPHILUS DECOLORATUS (Koch, 1844). (Figures 108 to 111, and 116) THE BLUE TICK L WN Q of EQUATORIA PROVINCE RECORDS iS) al Tereteina Alcelaphus buselaphus roosevelti Feb ap Gal Torit Aicelapnus buseLaphus roosevelti Dec (2) 10 3 Meridi Taurotragus derbianus gigas Feb (SVS) U2 Nagichot domestic cattle Jul 9 1 Nagichot domestic cattle Dec (2) aL Tleu domestic cattle Oct Ae 49) Katire domestic cattle Jan g Katire domestic cattle Oct 30 Gilo domestic cattle Dec LT 4 FLOTAt domestic cattle Jan 5B 4 Torit domestic cattle Feb 4 10 Torit domestic cattle Nov i 9p i Torit domestic cattle Dee (2) 6 Juba domestic cattle Dec ie Tombe domestic cattle Mar (SVS) 13 a Muni domestic cattle Mar (SVS) 11 Terakeka domestic cattle Mar (SVS) Pe Kajo Kaji domestic cattle Dec 10 Yei domestic cattle Mar (Sscc) 2 Sea Meridi domestic cattle May (SVS) 4 Yambio domestic cattle Jan 2 Li Rangu domestic cattle Apr ale Kapoeta domestic sheep Dec 3} 2 Juba domestic horse Jan 3002 More (Yei River) domestic donkey Jan (SGC) Al Gilo domestic dog Dec DISTRIBUTION IN THE SUDAN Equatoria, Bahr el Ghazal, Upper Nile, Kordofan and Blue Nile Province; specimens also taken from cattle arriving at the Halfa =e 300 Quarantine in Northern Province, but it is by no means certain that the species is established there (King 1926). Specimens have been seen from the following localities, except Upper Nile: Fangak and Ler (SVS). Maban (domestic pigs; SVS). Bor, Pariak, Malakal (SVS, HH). Akobo and Taufikia (SCC). Daga Kigille Road (Tora hartebeest; SGC). pune Nile: Fung District (Singa headquarters) (= Funo) (King 1926). Bahr El Ghazal: Wau (domestic donkeys, horses and cattle; SVS, HH). “Busseri (domestic horse; SVS). Fanjak (domestic dogs and cattle; SVS, HH). Galual.Nyang Forest (hartebeest and sick domestic donkeys; SVS; tiang; HH). Sixty miles north of Aweil (domestic goat; SVS). Khor Shammam, Boro, near Raga, Lau, Karu, Akot, Yirol, and Peth (SVS). Kordofan: Talodi (SVS). El Obeid (SVS and Gordon College collection). Khartoum: Khartoum (horse; Gordon College collection). Khartoum Quarantine Station (on Darfur cattle; HH). Darfur: Radom, Nyala, Sibdo, and Zalingei (domestic horses and cattle; SVS). Near Safaha (domestic sheep; SVS). Kassala: Kassala and Port Sudan (SVS). Only four female specimens are known from this Province. { Northern: Quarantine Station specimens from southern cattle; Scc./ Since it appears that B. decoloratus is not established in Northern Province and is very rare 1n Kassala Province, Sudan evidence corroborates earlier expressed beliefs that 15°N. is about the northern limit of this species. The most northern Sudan locality in which B. decoloratus commonly occurs is El Obeid, at 13°11'N. In Kassala Province, the blue tick is rare and is found, in association with either local humid coastal - 307 - situations or with cattle movements, as far north as Port Sudan, which is at almost 20°N. latitude. Small populations also exist in more humid riverside garden areas in and around Khartoum. DISTRIBUTION The blue tick is distributed throughout most of the Ethiopian Faunal Region, barring the northern and southern periphery of this region on continental Africa. Within its range, B. decoloratus occurs widely everywhere except in more open, dry areas, and in tropical forests. It is usually common where a fair degree of shade and humidity is present. Outlying specimens have been re ported from other continents but whether these are from imported hosts or from established populations is not clear. NORTH ATLANTIC OCEAN: CAPE VERDE ISLANDS (Howard 1908. Bacelar 1950). WEST AFRICA: NIGERIA (Simpson 1912B. Unsworth 1949,1952. Mettam 1951. Gambles 1951). GOLD COAST (Simpson 1914. Beal 1920. Stewart 1933,1934. Minning 1934). FRENCH WEST AFRICA (ee 1934. Rousselot 1951,1953B. Villiers 1955). LIBERIA (Minning 1934). PORTUGESE GUINEA (Monteiro da Costa 1926. Tendeiro 1946A, B,1948,1951C ,1952A,C ,D,E,1953,1954. Bacelar 1950). SIERRA LEONE (Hoogstraal 1954C). CENTRAL AFRICA: CAMEROONS (Ziemann 1912A. Minning 1934. Rageau 1951,1955A,B. Rousselot 1951,1953B. Unsworth 1952. Dezest 1953). FRENCH EQUATORIAL AFRICA (Minning 1934. Fiasson 1943B. Rousselot 1951,1953A,B). BELGIAN CONGO and RUANDA-URUNDI (Newstead, Dutton and Todd 1907. As B. capensis: Massey 1908. Roubaud and Van Saceghem 1916. Nuttall and Warburton 1916. Seydel 1925. Schwetz 1927A,B,C,1932, 1933B,1934. Schouteden 1929. Bequaert 1930A,B,1931. Minning 1934. Van Slype and Bouvier 1936. Bouvier 1945. Fain 1949. Schoenaers 1951A,B. Rousselot 1951,1953B. Theiler and Robinson 1954. Santos Dias 1954D. Van Vaerenbergh 1954). EAST AFRICA: SUDAN (Balfour 1911F incorrectly stated that B. decoloratus does not occur in the Sudan. King 1908,1911,1926. Hoogstraal 1954B,C). = 50's) ETHIOPIA (Minning 1934. Stella 1938A,1939A,B,1940. Roetti 1939). ERITREA (Franchini 1929D. Niro 1935. Stella 1938A,1939A, B,1940. Ferro-Luzzi 1948). FRENCH SOMALILAND (Hoogstraal 1953D). BRITISH SOMALILAND (Stella 1940). ITALIAN SOMALILAND (Niro 1935. Stella 1938A,1939A,B,1940). KENYA (Neave 1912. Montgomery 1919. Anderson 1924A,B. Lewis 1931A,B,C ,1932B,1934,1939A,B,1943. Fotheringham and Lewis 1937. Mulligan 1938. "Kenya Vet. Serv.” 1939A,1949,1952. Piercy 1948. Weber 1948. Binns 1951,1952. van Someren 1951. Wiley 1953. Hammond 1955). UGANDA (A. Theiler 1910A. Bruce et al 1911. Neave 1912. Richardson 1930. Mettam 1932,1933. "Minning 1934. Mettam and Carmichael 1936. Wilson 1948B,C 19500). TANGANYIKA (Donitz 1905. Neave 1912. Morstatt 1913. Knuth and du Toit 1921. Moreau 1933. Minning 1934. Cornell 1936. Lewis 1939A. Reichenow 1941B. J. B. Walker, unpublished; see HOSTS below). SOUTHERN AFRICA: ANGOLA (Howard 1908. Manetti 1920. Santos Dias 19500. Sousa Dias 1950. Bacelar 1950. Theiler and Robinson 1954). MOZAMBIQUE (Howard 1908,1911. Nuttall 1911B. De Oliveira 1915. Present in Tete District only: Theiler 1943B,1949A. Santos Dias 1947B,1950B,1952D,1953A,B,1954H. Bacelar 1950). NORTHERN RHODESIA (Neave 1912. Chambers and Smith 1914. Le Roux 1937, Matthysse 1954. Theiler and Robinson 1954). SOUTHERN RHODESIA (Robertson 1902,1904A,B. Koch 1903. Donitz 1905. Bevan 1912,1915,1919. Edmonds and Bevan 1914. Jack 1921,1928,1937,1938, 1942. Lawrence 1935,1942). NYASALAND (Old 1909. Neave 1912. De Meza 1918A,B. Wilson 1943,1946,1950B). SOUTHWEST AFRICA (Howard 1909. Tromsdorff 1913,1914. Minning 1934. Schulz 1939. Theiler 1949A). SWAZILAND (""Agricola™ 1946. Barnard 1949). UNION OF SOUTH AFRICA (Koch 1844. Dixon and Spruell 1898. Lounsbury 189% ,1900A,C ,1904A,1905B. Neumann 1901,1911. A. Theiler and Stockman 1904. A. Theiler 1905A,C,1906,1909B,€ ,1910B,C ,D,E, 1911B,1912A,1921. Laveran and Vallee 1905. Warburton and Nuttall 1909. A. Theiler and Christy 1910. Donitz 1910B. Moore 1912. Van Saceghem 1914. Bedford 1920,1924A,1926,1927,1929B,1931B, 1932B,1934. Cowdry 1925€,1926A,1927. Curson 1928, P. J. du Toit = 509° and Viljoen 1929. P. J. du Toit 1931. Alexander 1931. Bedford and Graf 1934,1935,1939. Pijper and Dau 1934. Cooley 1934. Minning 1934. M.D. 1936. Pijper and Crocker 1938. Neitz and du Toit 1938. J. H. S. Gear 1938. McIntyre 1939. R. du Toit 1942B ,€ ,1947A,B,1948. Cluver 1944. Theiler 1949A: important biological survey. Meeser 1952. J. Gear 1954). On arsenic resistance and BHO control and resistance in South Africa: du Toit, Graf, and Bekker 19Z1. Bekker 1942,19L4, 1975 ,1927,1953. “Anxious™ 1943. Bekker and Koch 1943. Dell 1943. Mullins 1944. Bagshawe-Smith 1944. Omer-—Cooper and Whitnall 1945. Whitnall and Fenwick 1945A,B. Whitnall and Brad ford 1945,1947A,B,1949. Bekker Graf 1946. “Agricola™ 1946. Thorburn 1947,1952. Whitnall 1947. Cooper 1947,1953. Thorburn 1947. Whitnall, Bradford, McHardy, Whitehead, and Meerholz 1948, 1949A,B. Whitnall, Thorburn, Whitehead, McHardy, and Meerholz 1949. Bekker, Graf, Malan and Van der Merwe 1949. Daly 1950. Whitnall, McHardy, Whitehead, and Meerholz 1951. Kruger 1951. Fiedler 1952. Whitnall, Thorburn, McHardy, Whitehead, and Meerholz 1952. Hitchcock 1953. Blomefield 1954. Busvine 1955. Metcalf 1955. OUTLYING ISLANDS: ZANZIBAR (Neave 1912. Aders 1917). MALAGASY GROUP: From present evidence, B. decoloratus is not established in the Madagascan archipelago (Hoogstraal 19555). Neumann (1901) listed Madagascar and Mauritius. Buck (1935) listed the former island, De Charmoy (1914,1915) and Moutia and Mamet (1947) the latter, and Gillard (1949) mentioned Reunion. Minning (1934) referred Neumann's material to B. microplus (= B. fallax). Buck (1948A,C) reports that P. decoloratus has never been found on Madagascar or on other islands in this group. NOTE: Specimens of B. decoloratus from North Syria, West Kurdestan, and Buenos Aires (South America) have been noted (Minning 1934), but without mention of whether populations are maintaining themselves in any of these localities. The blue tick is sometimes found on cattle from the Sudan and East Africa at the Cairo abattoir. This species has never been found on native cattle in Egypt. - 310 = HOSTS Cattle are the chief host of the blue tick and most papers listed in DISTRIBUTION above refer mainly to this animal and ef- fects of this tick's feeding upon it. Domestic horses and, less frequently, sheep and goats are attacked. Among wild animals, antelopes are important hosts but few others are infested. Wild carnivores are almost never parasitized by this tick. This is a single host tick, therefore hosts of the immature stages are the same as those of adults. Domestic animals: Cattle (Most papers listed in DISTRIBUTION above). Horses (Howard 1908, A. Theiler 1911, Lewis 1931C, Mettam 1932, Minning 1934). Mules (Howard 1908). Donkeys (Howard 1908, Minning 1934, Sousa Dias 1950, Sudan records above). Sheep (Howard 1908, Schwetz 192%, Mettam 1932, Lewis 1934, Minning 1934, Wilson 1950B, Sousa Dias 1950, Sudan records above). Goats (Howard 1908, Lewis 19310, Bedford 1932B, Mettam 1932, Cooley 1934, Tendeiro 1948, Wilson 1950B, Sousa Dias 1950, Sudan records above). Pigs (Knuth and du Toit 1921. Sudan records above). Dogs (Lounsbury 1904A reared the blue tick on dogs, but only a few specimens com pleted their life cycle on these animals. Howard 1908, Bedford 1932B, Mettam 1932, Sousa Dias 1950. Bahr el Ghazal specimens in Sudan records above were collected from dogs belonging to Dinka herdsmen; these animals sleep in cattle huts). Camels (King 1926). Water buffalo (Theiler, unpublished). Man; (Nuttall 1911B, Bedford 1920). Antelopes: Grant's gazelle (Weber 1948). Sable antelope (Bedford IOS, Cooley 1934, Jack 1942. Roan antelope (Simpson 1914, Jack 1942). Blue wildebeest (Bedford 1932B). Gnu (Santos Dias 1950B). Hartebeest (Lewis 1934, Santos Dias 1950B, Sudan rec ords above). Nyala (Santos Dias 1950B,1952D). Tiang (Sudan records above). Impala (Bedford 1932B, Cooley 1934, Jack 1942, Meeser 1952, Santos Dias 1952). Reedbuck (Santos Dias 1950B). Bushbuck (Mettam 1932, Lewis 1943, Santos Dias 1950B). Waterbuck (Bedford 1932B). Eland (Lewis 1943. Sudan records above). Sitatunga (Bequaert 1931). Oryx (Minning 1934). Duikers (Mettam 1932, Cooley 1934). Topi (In Miss J. B. Walker's collections from nu Merous game animals in Tanganyika, the blue tick is represented only by several females and a male from two of these hosts). - 3ll- Other wild animals: Hares (Tromsdorff 1914, Bedford 1932B, Cooley 1934). Zebra (Bedford 1932B). Bushpig (Santos Dias 1950B). Buffalo (Old 1909, Schwetz 1927B,C, Mettam 1932, Jack 1942). BIOLOGY Life Cycle B. decoloratus is a one-host tick. Females lay their eggs on the ground. When larvae find a suitable host they remain on it, either on the dewlap and neck or on the ears, at the tip or along the upper edge of the pinna. On the ears, larvae are often associated with immature stages of R. appendiculatus. At this locus larvae molt to nymphs and nymphs ort to adults. Adults usually move to the belly and flanks of the host, and mate on the animal. The life cycle is completed entirely on the host except for oviposition, in from three weeks (A. Theiler 1911B) to a month (Lewis 1939B). Lounsbury (1905B) stated that females leave the host 23 days after having attached as larvae, but males may remain on the host for another month. He listed the pre— oviposition period as six to nine days. Eggs hatch after five weeks and larvae molt one week after having completed feeding. Wintertime egglaying and hatching is much slower. Unfed larvae may survive for over eight months (Theiler 1949A). Ecology A. Theiler (1911) reported such a heavy infestation of B. decoloratus on a horse that it died of acute anaemia. Half of the ticks were collected; they weighed fourteen pounds. More recent literature on arsenic-resistance in South African blue ticks also frequently refers to and illustrates markedly heavy infestations of host animals. Because of their unique life cycle, boophilids are readily controlled by dipping cattle, and there are numerous reports of the eradication of B. decoloratus from large areas. New control problems are posed, however, by the development of resistant strains. = S12 ne From Theiler's (1949A) survey of conditions under which the blue tick exists in South Africa she concludes that the most im portant factor in limiting its spread is increasing aridity. In most parts of South Africa, the critical level is represented by an annual rainfall below fifteen inches. B. decoloratus is absent in deserts except for introductions that do not become established. It is present at all altitudes from sea level to high mountains, and can withstand both frost and high temperature. Similar though less detailed observations were reported by A. Theiler (1921). In tropical African forest, the blue tick occurs probably only in open, cleared areas (Theiler and Robinson 1954). The data confirm Lewis! (1939) findings for Kenya, where B. decoloratus inhabits moist regions, highlands up to over 8000 feet altitude, and forests and glades but seldom open, dry, scrub areas. In Kenya, it is also resistant to heat and cold provided moisture is available. In RuandaUrundi the altitudinal range does not exceed 6500 feet (Schoenaers 1951B). In the Belgian Congo, Bequaert (1931) found the blue tick to be so common that it was unnecessary to list all the localities from which specimens were secured. Yet Van Vaerenbergh (1954) reports this species as generally distributed in the Congo and Ruanda-Urundi but represented by a small number of specimens or absent from many lots in his collections. The discrepancies in the inferences of these two authors undoubtedly result from dif. ferences in collection areas and methods, interests of collectors, number of ticks taken and hosts examined, and other factors. Sim ilar situations have provided widely differing reports for the in cidence of other species of ticks elsewhere, as for instance R. S. simus on cattle in Nyasaland and in South Africa and R. s. Sanguineus on dogs in eastern and southern Africa. It is obvious. Ty ete to generalize on population abundance in large areas and difficult to evaluate generalizations not supported by data. After having seen Dr. Bequaert's extensive collections, it should be confirmed that the blue tick is certainly numerous in many Congo areas and that it is uncommon or absent at high elevations and in dense forest areas. In Nigeria this is said to be the second most common tick from domestic animals and represented 28% of the specimens collected from domestic animals in all Provinces (Mettam 1951). Only A. variegatum at 45%, is more common in these collections. = - 313 - Wilson (1953) has stressed that the factors governing the dis. tribution of this tick in East and Central Africa require further study. In untreated Northern Rhodesian cattle, ™blue ticks in all stages were present ..... throughout the full year's observation. These ticks quickly became very abundant, the cattle being gross. ly infested through May, June and July. There was no evidence of decrease in infestation throughout the dry season apart from a slight decrease in late July and August. The blue tick infesta. tion did not build up during the wet season". / Matthysse (1954)7 The redubilled oxpecker or tickbird, Buph e. Se chus (Stanley), which attends all of the larger ee excep the elephant and hippopotamus, is a predator of some importance on B. decoloratus and on other economically important ticks. Of 58 tick-birds examined in Tanganyika, 51 blue ticks were found in the stomach contents of thirteen; the number of ticks per stomach ranged from one to ten (Moreau 1933). In Kenya, van Someren (1951) found 38 blue ticks in stomach contents of four out of twelve of these birds that he examined. See p. 275 for a discussion of this sub. ject. REMARKS Schulze (1936A) remarked that the spurlike prolongation of the male adanal shield is sometimes separated from the base in a posi- tion similar to that of the subanal shields of Hyalomma species. Two hundred and fifty males have been examined from various parts of Africa without seeing a similar condition. Schulze also (1932C) compared the adanal shields of B. decoloratus with morphological peculiarities of fossil Bopueyane: The triple capsule of the haller's organ in B. decoloratus has been described and illustrated by Schulze (1941). Double diagonal striations of the nerve fibers have been noted and illustrated by Kruger (1935). The blue tick has been employed by Gossel (1935) to delineate features of the eyes and their related cells in ticks. Abnormal specimens have been noted (Warburton and Nuttall 1909, and Bedford 1924A). - 3uU- DISEASE RELATIONS MAN: Evidence that this tick transmits boutonneuse fever (Rickettsia conorii) appears to be entirely presumptive. The bite may in itself result in severe inflammation, but man is probably seldom attacked by this tick. Q fever (Coxiella burnetii) is claimed to have been found in specimens from Portugese Guinea. CATTLE: Redwater or Texas fever (Babesia bigimina). Spiro— chetosis (Borrelia theileri). Gallsickness (Anaplasma marginale). Not a vector or heartwater (Rickettsia ruminantium). The virus of “a specific transmissable petechial fever of cattle” survives in this tick. HORSES, SHEEP, and GOATS: Spirochetosis e. theileri). Not a vector of equine piropLasmosis (Babesia equi). PIGS: Possibly a vector of porcine babesiosis (Babesia traut— manni), Domestic animals may be so severely irritated that their health is impaired and deaths from the bloodsucking of huge numbers of B. decoloratus have been recorded. A toxin from the egg of B. decoloratus has been described. IDENTIFICATION Males are easily determined by the presence of a small tail. like caudal appendage, which varies considerably in size, on the posterior body margin and by a narrow spurlike elongation of the internal margin of the adanal shield, which reaches to or more commonly extends beyond the posterior body margin. The unique, small bristle-bearing protuberance on the internal ventral sur. face of the basal palpal segment is most important; this feature may be difficult to discern in small specimens and the bristles are often broken, though the knobs may usually be seen. Hypo- stome dentition is 3/3, rarely 3.5/3.5. Size is very small, about 2.7 mm. long and 1.5 mm. wide. =, 315 Female characters may be difficult to determine satisfactorily in greatly engorged, preserved specimens. Coxa I has two well. defined spurs discernably separated by a narrow but deep, inverted V-shaped cleft. Palpal segment one ventrally has an internal knob bearing one or two bristles; these bristles may be broken or obs— cured by crowding in preserved, engorged material. The scutum rarely may be approximately as broad as long, but more commonly is slightly longer than broad; the scutal margin, anterior of the eye, curves very slightly outwards. The hypostome formula is as in the male. Engorged individuals may reach 12.0 m. or more in length and about 8.0 mm. in width. The normal slatish color of the engorged female and the blue of the nymph gives this tick its common name. Most specimens in my collection contrast with the usually paler females of the other two boophilids. 3 ERG Figures 112 and 113, @, dorsal and ventral views Figures 114 and 115, 9, dorsal and ventral views BOOPHILUS MICROPLUS Northern Rhodesia Specimens Presented by Dr. G. Theiler PLATE XXXVI - 317 - 3/3 dentition bristle Dearing protuberance dentition inner margin elongate, mildly concave dentition inner margin short, deeply concave 18 cleft Figure 116, B. decoloratus, ventral view (Sudan). Figure 117, B. annulatus, ventral view (Sudan). Figure 118, B. microplus, ventral view (N. Rhodesia). Figure 119, dorsal = ew of same species. DIFFERENTIATION OF BOOPHILUS FEMALES PLATE XXXVII - 318 . NOTES ON BOOPHILUS MICROPLUS (Canestrini, 1888) (= B. FALLAX Minning, 1934). (Figures 112 to 115 and 118) THE PANTROPICAL CATTLE TICK According to current concepts, the three Boophilus species of the Ethiopian and Malagasy Faunal Regions are the OnLy ones known to comprise this genus throughout the world and subspecific desig- nations are of questionable accuracy. The identification of these species has been much confused in literature. The single species not yet known from the Sudan, B. microplus (= B. fallax), threatens to encroach upon these borders, therefore succinct notes on this latter tick are interpolated in spite of the general practice of excluding non-Sudanese species from this report. As elucidated below, it is impossible to differentiate so called diagnostic differences between specimens from African popw lations of B.“fallax™ and specimens of B. microplus from popula tions in the Near and Middle East, various parts of Asia, and South America. Lengthy study of long series of material from these areas causes me to agree with Anastos (1950) that all these populations appear to represent a single species. On the other hand, Theiler reports (correspondence) that not only can she distinguish between B. fallax and B. microplus, but students with only basic knowledge of tick identification easily separate both species when presented as unknowns to be keyed! This matter is one of the very few items concerning which agree. ment has not been reached with Theiler, after an extended exchange of notes and ideas. Theiler states (correspondence) that in her experience in the more southerly parts of Africa, B. “fallax” is usually local. ly restricted, but where it does occur it is frequently numerous. — Sy There is a generalized, vague impression among students of the African fauna that B. fallax is an endemic Madagascan tick that has been introduced into Africa. However, the predilection of the Boophilus ticks for ruminants, absent in the endemic Madagascan fauna, as well as the apparently slow evolution of boophilids would appear to negate any possibility that the species may have evolved there within the few centuries that cattle have been introduced to these islands. It-seems more likely that cattle from southern Asia brought their characteristic ticks with them to Madagascar. For over half a century there have been periods of heavy importation of Madagascan cattle into South and East Africa, and these probably introduced this tick into Africa. Export of cattle from infested African areas to Madagascar may have returned some ticks to the areas of their immigrant ancestors. DISTRIBUTION “B, microplus is found in Central America, South America, Australia, the Oriental Region, in the southern part of Florida, and in parts of Africa ...... After examining specimens of B. fallax Minning from South Africa ....... I am convinced that this species is a synonym of microplus” (Anastos 1950). The following are selected records from the Ethiopian and Malagasy Faunal Regions for "B. fallax". EAST AFRICA: UGANDA (Wilson 1948A,1950C). TANGANYIKA (Minning 1934. Theiler and Robinson 1954). SOUTHERN AFRICA: MOZAMBIQUE (Theiler 1943B. Santos Dias 1953B, 1954H,1955A). NORTHERN RHODESIA (Matthysse 1954. Theiler and Robinson 1954). SOUTHERN RHODESIA (Theiler and Robinson 1954). NYASALAND (Wilson 1946,1950B. Theiler and Robinson 1954). UNION OF SOUTH AFRICA (As B. australis: Fuller 1899 and Lounsbury 1905. As B. annulatus: Dénitz 19108. As Margaropus annulatus australis and as B. microplus: Howard 1908. Minning 1932. Theiler LOJ5B. Theiler and Robinson 1954). MALAGASY REGION: MADAGASCAR (Minning 1934. Buck 1935,1948A,C. th 1s Buck and Metzger 1940. Buck and Ramambazafy 1950. Zumpt 1950B. = 320% Cordurier, Buck, and Quesnel 1952. Hoosstraal 1953E. The B. ‘“caudatus" of Colas-Belcour and Millot 1948 may refer to this spe- cies. ifinning 1934 refers all reports of B. decoloratus fron Madagascar and other islands in this group to B. microplus (= B. fallax). It appears that B. decoloratus has not sctabtished ite self in this archipelago). WAURTTIUS (De Charmoy 1915. -‘toutia and Mamet 1947. Millot 1948. Hoorstraal 1953E). REUNION (Gillard 1949. Hoogstraal 1953E. The "B. caudatus reported by Neumann 1897 may refer to this species). COMORES GROUP (jfinning 1934. Millot 1948. Hoogstraal 1953E). SEYCHELLES (Desai 1952). HOSTS All authors report domestic cattle as the chief host. “finning (1934) also noted specimens from a domestic horse and Theiler (1943B) from domestic sheep and poats. Buck (1935,1949A,C) found material on domestic sheep and Buck and Ramambazafy (1950) on domestic horses. The only wild animal known to have served as a host in Africa is a lion (Theiler 1943B). Anastos (1950) reports chiefly domestic cattle but also a variety of other domestic animals as hosts of B. microplus in Indonesia. He noted that records from wild mammals and birds and from domestic chickens are extant. BIOLOGY IN ETHIOPIAN AND MALAGASY FAUNAL REGIONS Life Cycle This is a single host tick. Engorged females leave the host from 35 to 149 days after having attached as larvae, and there may be from two to three generations a year in South Africa (Lounsbury 1905). Wilson (1946) observed no seasonal periodicity of adults in Nyasaland. He found larvae with nymphs and adults on cattle only once. Nymphs and adults were usually found together. Nymphs and adults are almost constantly restricted to the udders, flanks, and belly; larvae to the inner side of the ears of the host. Elsewhere, a number of biological studies on this species have been reported. Among these are Sapre (1940) for high alti- tudes in India, Tate (1941) for Puerto Rico, Legg (1930) for Australia. Beotouy As already stated, B. microplus occurs only locally in Africa but where present it may be very common. In Northern Province of Nyasaland it is more numerous on cattle than B. decoloratus (WilL son 1946). In the Malagasy Region, B. microplus is Largely a low. land species with scattered foci around urban highland localities (Buck 1948A,C). In Africa it survives best in natural forest conditions (Theiler 1943B). Barly in the century, the range of B. microplus in eastern and southern Africa was wider than it now is, probably because of more frequent importation of infested cattle from Madagascar at that time. Climatic conditions have reduced these populations to their present more localized foci (Theiler 194.38) but exten sions of these infestations should be anticipated. This tick was introduced into southern Africa after the 1896 rinderpest out— break (Theiler and Robinson 1954). REMARKS A misshapen specimen has been described and illustrated by Santos Dias (1955A). My collection contains two gynandromorphs. The synonymous B. australis Fuller, 1899, was described in South African literature. Fuller also gave a differential diag_ nosis for the three forms presently recognized as comprising this genus. DISEASE RELATIONS Unstudied in Africa. The following references are for the Americas, Asia, and Australia. SPR Cattle: Redwater or Texas fever (Babesia bigemina). Babesiosis (Babesia berbera). Anaplasmosis or gallsickness (Anaplasma marginale). ees Sheep: Babesiosis (Babesia ovis). Horses: Biliary fever (Nuttalia equi). Wherever and under whichever name it occurs, this tick appears to be of considerable veterinary importance. IDENTIFICATION Male; This small, yellowish to reddish brown tick varies from about 1.o mm. to 2.5 mm. in overall length, and from 1.0 m. to 1.4 mm. in width. The presence of a short, tapering caudal append age is noteworthy (absent in B. annulatus, present in B. decoloratus, but in B. microplus the internoposterior juncture of the adan shields does = extend beyond the posterior body margin). The hypostome dentition is 4/4 (typically 3/3 in B. decoloratus); and the inner margin of the basal palpal segment ventrally 1S concave (bearing a bristle-bearing protuberance in B. decoloratus). The scutum is quite hirsute. a Minning (1934) indicated, as his primary critical difference between B. fallax and B. microplus, the bluntness or acuteness of the inner spur of coxa 1. ‘this character is variable among spec- imens from Africa and other continents; figure 112 illustrates a specimen from Northern Rhodesia in which this spur is blunt on one side and acute on the other! Female: After some experience females should be easily recog- nized in African collections, provided one carefully and methodical. ly observes clean specimens from which the wet surface film of preservatives has been removed. Like that of B. annulatus, this sex has 4/4 hypostome denti- tion. B. microplus can be distinguished because the spurs of coxa I are Aed by a deep, inverted "V" shape cleft, while those of B. annulatus are separated merely by a shallow, concave emargination (this character should be ascertained by placing the specimen obliquely against the light). No bristle-bearing pro- - 323 - tuberance is found on the inner margin ventrally of the basal pal. pal segment. The inner margin of the basal sesment is short and deeply concave by comparison with that of B. annulatus. The palpi usually appear more compact and less acutely ridged. Minnine'ts (1934) character for separating EB. microplus and B. fallax on the basis of the anterior curvature of palpal secment 3 applies only to his illustrations, not to specimens. The scutun, normally slightly longer than wide, may be widened by engorgement (from 0.40 mm. to 0.58 mm. long and from 0.34 m. to 0.50 m. wide). The eyes are generally oval and raised above the scutal surface but this character may be difficult to discern. Size varies from about 2.0 m. to 12.5 mm. long, and from 1.0 mm. to 7.9 mm. wide, depending on degree of engorgement. Ai SYS DERMACEN TOR INTRODUCTION Dermacentor, a medically important genus in many parts of the world, is represented by only three species in tropical Africa. Small populations of two of these, D. c. circ ttatus and D. rhinocerinus, occur in Equatoria and Bahr Ghazal Provinces of the Sudan. In Africa they are always rather rare and largely con fined to the rhinoceros and elephant. As such, they are of little more than academic interest. The identity of these species may be determined easily from the following keys and illustrations. The third African species, D. hi tamensis (Denny, 1843) (= Ixodes bimaculatum Denny, 1843, and yomma hi tami Koch, 1842), was originally described from Hi tamus amphibius of South Africa. Schulze (1919) erected the genus Cosmiomma for this species on the basis of its Hyalomma-like characters although it lacks accessory shields and subanal shields. Zumpt (1951) sank Cosmiomma under Dermacentor but Theiler states (correspondence) that Schulze's definition justifies its retention as a genus. Other authors have placed it in Hyalomma. Still another study by a qualified student on the Aer material appears necessary be— fore an acceptable systematic niche can be found for D. (C.) hippo potamensis. D. (C.) hi tamensis has been reported from South Africa and between “Zaetbe “Gi.e. East Reviee) vane the Great Lakes (Tan. ganyika). For over a century, the only positively known specimens have been the types described by Denny and by Koch, which have been seen again by later students. In the collections of the East Afri- can Veterinary Organization there is a single male taken from vege— tation at Manyani, Teita District, Kenya, 5 November 1951, D. L. W. Sheldrick legit. This specimen, according to J. B. Walker (cor- respondence), is almost exactly like the type material illustrated by Dénitz (1910B). One or two specimens are in the collection of the Veterinary Department at Kabete, Kenya. D. (C.) hi tamensis is a large, brightly colored Amblyomma like tick. The male scutum is described as pale straw-yellow wit symmetrical black markings and a few small punctations; ventrally 5 BOS the legs are a deep liver-red. The female scutum is lightly punc- tate, largely pale in color with two submedian longitudinal black stripes and a pair of vertical black stripes near the posterior margin. The dorsal surface of the female posterior of the scutum is dull crimson with two large nearly circular, slightly elevated orange spots near the lateral margin. The apex and lateral mar. gin of the palpi are dull crimson. The last tarsal segment and ventral surfaces of the legs are colored as in the male. This all but extinct beast should not be difficult to recognize. Another species, D. niveus Neumann, 1897, parasitizes wild boars in Tunisia, Algeria, and Spanish Morocco (Senevet, Colas— Belcour, and Gil Collado 1933), and various other animals in Ew rope and Asia. It is difficult to determine what "D. reticulatus Neumann," listed by Stella (1938A,1939A,1940) from Ethiopia and by Niro (1935) from Somaliland, actually is. D. reticulatus, which does not appear to be a synonym of D. marginatus Sulz., 1776, as stated by Schulze (1933C), inhabits Europe and Asia. If its range does extend into the Ethiopian mountains, it would represent a unique ixodid distributional pattern, but conceivable on the basis of geographic distribution of other invertebrates. Although it has not been our policy to discuss non-Sudanese species, the above remarks and a few additional taxonomic notes are inserted inasmich as no review of the genus Dermacentor in Africa is available. Extra-Sudan species have not been demons trated to be of medical importance and will not be included in subsequent volumes of this undertaking. In various papers on Dermacentor, Schulze has divided the genus into several genera that show interrelationships within a closely circumscribed group and can, by contemporary concepts, be considered at most only as subgenera. These are of only slight interest in Africa, especially as the mot subject of subgenera will be further revised in future studies. D. circ ttattis was placed in the subgenus Puncticentor, which was subsequently syno- nymized (Zumpt 1951) under the subgenus Amblyocentor, in which D. rhinocerinus had been placed. The usefulness of the latter cate. gory 1s questionable in the absence of study of the entire genus. These same remarks are possibly pertinent to the subgenus Cosmiomma, originally proposed as a full genus embracing only D. hippopotamensis. = 3262 KEY TO SUDAN SPECIES OF DERMACENTOR MATES Ornamentation consisting of eight small, pale spots near scutal periphery; festoons and central scutal area darks ceccccscccccccesels Cc. CIRCUMGUTTATUS Figures 120 and 121 Ornamentation consisting of seven large pale spots, which cover most of scutun, and of smaller peripheral and festoon SOiv Gis leleleleie clelelsleicisicioicleicleleiclelclelelelelaielelelelelelelelercloio'e cle elefel)c RHINOCERINUS Figures 124 and 125 FEMATES Scutum largely dark, ornamented with three pale SPOLScerccscccvcccsccscccccccsscscenle C. CIRCUMGUTTATUS Figures 122 an Scutum largely pale, a few small dark Spots present or abSenteccccccsssccsccccccccsecceeeds RHINOCERINUS Figures 126 and 127 es Se Figures 120 and 121, o&, dorsal and ventral views Figures 122 and 123, 9, dorsal and ventral views DERMACENTOR C. CIRCUMGUTTATUS Uganda Specimens Loaned by British Museum (Natural History) PLATE XXXVIIT Selon DERMACENTOR CIRCUMGUTTATUS CIRCUMGUTTATUS Neumann, 1897. (Figures 120 to 123) THE ELEPHANT DERMACENTOR i Wi ps0 5c; EQUATORIA PROVINCE RECORD Lea Kajo Kaji on grass Dec (BMNH) British Museum (Natural History) specimens, collected by Captain C. H. Stigand in the early 1900's, are the only ones extant from the Sudan (Hoogstraal 1954B,C). This is the mst northern and eastern record for the occurrence of the rare West African elephant dermacentor. It is unlikely to be found else— where in the Sudan other than on the west bank of Equatoria Province and possibly in Bahr El Ghazal Province. DISTRIBUTION D. c. circumguttatus, a Central and West African elephant parasite, extends into East Africa only so far as the western parts of Uganda and the Sudan. In Mozambique, a separate sub. species is tentatively recognized on the basis of somewhat larger size and more irregular spotting. WEST AFRICA: LIBERIA (Bequaert 1930A). SIERRA IEONE (Simp son 1913). IVORY COAST (Rousselot 1951,1953B). CENTRAL AFRICA: CAMEROONS (Neumann 1901. Ziemann 1905. Rageau 1951,1955A,B). FRENCH EQUATORIAL AFRICA (Neumann 1897. Tonelli-Rondelli 1930A. Rousselot 1951,1953B. Rageau 1953B). BELGIAN CONGO (Neumann 1897. Nuttall and Warburton 1916. Schwetz 1927A,B,C. Schouteden 1929. Bequaert 1930A,B,1931. Tonelli. Rondelli 1930A. Rodhain 1936. Fain 1949, Schoenaers 1951A. Theiler and Robinson 1954). . NOTE: According to Theiler (correspondence), the record for Ruanda_Urundi by Santos Dias (1954D) is incorrect. = B28) & EAST AFRICA: SUDAN (Hoogstraal 1954B). UGANDA (Neave 1912. Mettam 1932. Wilson 1948A,1950). HOSTS All authors report this species from elephants, Loxodonta africana subsp. Mettam (1932) also listed the “common duiker™. BIOLOGY Unstudied. In Banningville territory of Belgian Congo "all sick elephants carry this tick and Amblyomma tholloni, sometimes in great numbers” (Fain 1949). DISEASE RELATIONS It has been suggested that either this species or Amblyomma tholloni may transmit piroplasmosis (Nuttallia loxodontis) of elephants in the Congo. REMARKS Schulze (1941) noted features of the tarsus and haller's organ of this tick. Santos Dias (1952F) has separated Mozambique populations into another subspecies, D. circ ttatus cunha-silvai. These specimens are larger than those ar more northerly parts of Africa and exhibit more irregular pale scutal coloration in several smaller, more divided spots rather than in three large spots as in females of the typical subspecies and eight large regular spots as in males of the typical subspecies. These dif- ferences, howeyer, appear to be similar to individual variants of D. rhinocerinus, reported by Bequaert (1930B), and larger sub. sequent collections may indicate the necessity of dropping this subspecies. = BBO) — IDENTIFICATION Characters in the generic key readily identify this tick and separate it from the only other species in the Sudan. - 331 - ic5 127 Figures 124 and 125, o, dorsal and ventral views Figures 126 and 127, 9, dorsal and ventral views DERMACENTOR RHINOCERINUS Sudan Specimens PLATE XXXIX - 332 - DERMACENTOR RHINOCERINUS (Denny, 1843) (= D. RHINOCEROTIS of authors) (Figures 124 to 127) THE RHINOCEROS DERMACENTOR ae NG Or C, EQUATORIA PROVINCE RECORDS lige Magwe on grass Feb (SVS) ee Nimule on grass Jan (just west of) Localities uncertain 4. 2. Acholi (Lado) May (SCC) S412 Atiambo (Lado) Jun (SGC) 8 10 Azzar (Mongalla) May (SGC) Syeartg) How River (Lado) Mar (SGC) The material from uncertain localities near the Sudan_Uganda border, which was the basis of King's (1911,1926) reports of the rhinoceros dermacentor from the Sudan, was collected by him in 1909 and 1911. The recent specimens from Magwe and from the game reserve just west of the Nile near Nimle establish this species on both the west and east banks of Equatoria Province. DISTRIBUTION IN THE SUDAN Bahr El Ghazal: All specimens from grass, near Yirol to 25 miles west of Yirol; 23 November 1954, E. T. M. Reid lerit (7%, 509). 9 June 1954, P. Blasdale legit (19). Material In MH col lection. DISTRIBUTION D. rhinocerinus parasitizes the black and the white rhinoceros in eastern and southern Africa and in the Belgian Congo. - 333 - CENTRAL AFRICA: BELGIAN CONGO (Schwetz 19270. Bequaert 1930A, B,1931. Tonelli-Rondelli 1930A). NOTE: According to Theiler (correspondence), the record for Ruande-Urundi by Santos Dias (1954D) is in error. EAST AFRICA: SUDAN (King 1911,1926. Hoogstraal 1954B). ETHIOPIA (Warburton 1910. Neumann 1922. Stella 1940). ERITREA (Stella 1940). ITALIAN SOMALILAND (Pavesi 1895. Paoli ea Tonelli-Rondelli 1930A. Niro 1935. Stella 1938A,1939A, 1940). KENYA (Neumann 1912,1913,1922. Neave 1912. Anderson 1924A, B. Brassey-Edwards 1932. Lewis 1932A,1934). UGANDA (Neave 1912. Mettam 1932,1933. Wilson 19500). TANGANYIKA (Gerstacker 1873. Neumann 1907C ,1910B,1913,1922. Morstatt 1913. Bequaert 1930A. eer Schulze 19444. J. B. Walker, unpublished, see HOSTS below). SOUTHERN AFRICA: ANGOLA (Karsch 1878. Howard 1908. Bacelar eee MOZAMBIQUE (Howard 1908. Bedford 1932B. Santos Dias 1953B). NORTHERN RHODESIA (Hoogstraal 1954C). SOUTHERN RHODESIA (Jack 1942. Specimens from Sebungwes Theiler, correspondence). NYASALAND (Old 1909. Neave 1912. De Meza 1918A). SOUTHWEST AFRICA (Thejler, unpublished). UNION OF SOUTH AFRICA (Denny 1843. Gerstacker 1873. Howard 1908. Donitz 1910B. Breijer 1915. Curson 1928. Bedford 1932B. The localities in the first four references cannot be accepted without question; only Zululand records are sure (Theiler, correspondence). Specimens of "subspecies permaculatus" (see REMARKS below) from "Zambeze" were reported by Neumann (19070 ,1910B). NOTE: Records from ZANZIBAR (Howard 1908, quoted by later authors) probably result from the use of this name for East Africa by early writers. - 334 - HOSTS The black rhinoceros, Diceros bicornis subspp., and the white, or square-lipped, rhinoceros, Ceratotherium simum subspp., are listed as hosts by all authors. The hosts of our Equatoria Prov- ince specimens listed above, if they were rhinoceros, represent both genera inasmich as only the black rhinoceros occurs near Magwe east of the Nile and only the white rhinoceros occurs across from Nimle west of the Nile. In Tanganyika each of several rhinoceros hosts of various ticks yielded small numbers of this species (J. B. Walker, unpublished). King (1926) suspected, from the places in which he found spec. imens of this tick, that it may also attack the hippopotamus. Evans (1935) recorded domestic cattle and sheep as hosts in Tanga. nyika. Lewis (1934) mentioned a domestic donkey in Kenya and De Meza (1918A) took specimens from domestic cattle in Nyasaland and from elephants in Tanganyika. Neumann (1907 ,1910B8) listed an eland as host, and Mettam (1932) noted the jackal from Uganda. BIOLOGY Unstudied. Specimens are commonly reported from the hosts? genitalia. DISEASE RELATIONS Unstudied. REMARKS That the specific name rhinocerotis de Geer (1778), used by many authors for this species, actually applies to a distinct species in the genus Amblyomma was indicated by Donitz (1910B). Schulze (1932A) suggested that D. rhinocerinus be placed in the genus Amblyocentor on the basis of minor morphological pecu. liarities. There is, however, little utility in fragmenting tick - 335 - genera on the basis of insignificant characters. Amblyocentor is therefore considered as a subgenus of Dermacentor. A few male specimens have the anterior spots of the scutum partially or completely fused, thus resembling the female scutum. Neumann (190% ,1910B) described the subspecies permaculatus on the basis of these differences. Subsequent investigators with the exception of Tonelli-Rondelli (19304), have disregarded this name and considered these characters to be no more than individual variation (Bequaert 1930B). A "provisional name, D. rhinocerotis arangis, was applied by Lewis (1934, p. 39) to specimens of variable ise but after com parison with other specimens this name was withdrawn (footnote of same page). Females have two large patches of reddish brown hairs and scattered lighter hairs near the posterior margin of the body dor sally. These, and the cuticle of this species, have been studied by Schulze (1944A) and Jakob (1924). Schulze (1941) noted features of the tarsus and haller's organ, and (1950A) of the dentition of this tick. IDENT IFICATION Key characters readily separate and identify the two Dermacen tor species discussed in this report. = 336 = HAEMAPHYSAL1S INTRODUCTION Haemaphysalids are so small and inconspicuous, except when the females become greatly engorged, that they are seldom ade— quately represented in collections. Collectors frequently over— look them when larger and more colorful ticks are present. Many species show a marked predilection for seldom examined hosts such as hyraxes, birds, and hedgehogs. Some haemaphysalids appear to be actually quite rare in nature. In tropical and southern Africa, the genus Haemaphysalis is represented by the ubiquitous H. leachii subspp., She a car. nivore parasite, and by approximately fifteen less common species. In the nearby Madagascan archipelago, among whose ten known endem. ic tick species are nine haemaphysalids, most are distinctly re lated to the Oriental fauna. Asia has some fifty or more haemaphy-— salid species, which, in proportion to the total tick fauna, are to that continent what rhipicephalids are to Africa. A dozen forms are listed in the Russian fauna (Pomerantzev 1950). Of a total of eighteen ixodid species in the Philippines (Kohls 1950), not including the cosmopolitan kennel tick, one third are haema— physalids. The Americas and Europe claim only about five species each. Since Nuttall and Warburtonts (1915) revision of this genus, the African haemaphysalid fauna has received but little attention from biologists, systematists, or collectors. Many records in. cluded here represent considerable extensions of known range. Obviously, some few African species remain to be discovered and described. Differentiation of most African haemaphysalids is relatively easy, either by certain combinations of characters or by unique characters for individual species. Morphological char_ acters and facies of most species are comparatively quite constant. An important exception is H. leachii subspp., among the African forms of which there is very Considerable variation. Haemaphysalids are usually three-host parasites, although exceptions do occur. The life cycles of H. 1. leachii and of H. aciculifer have been fairly well studied in the Laboratory, but - 337 - few reliable biological data are available for African species. Factors governing morphological and biological variations of H. leachii subspp. pose an especially intriguing problem. Medically, the only African haemaphysalid of known importance is H. 1. leachii, a vector of boutonneuse fever (tick typhus) of man and of malignant jaundice of dogs. The same species may also be a reservoir of Q fever. The high potential of H. bequaerti as a medically important species has been recently suggested an others probably eventually will be incriminated in disease trans— mission of academic or practical interest. - 338 - KEY TO SUDAN SPECIES OF HAEMAPHYSALIS MALES Coxa IV with needlelike spur that is longer than coxal width. (On SOULHELNG TUM NENTS) tise e cee ieee eeeess cee eececce nelle BCICULINER Figures an Coxa IV with spur much shorter than Coxala wid beOrkwi CHOW SPUl ais csisiclelee «e c.c's 310 «1c s oles oc 'eie cicle'eic'g co Palpal segment 2 with strong dorsal retrograde spur. Lateral grooves extending anteriorly only to scutal midlength. (Equatoria mountains, ATOM) cvcaiavsiaisiolslere aicjslae cio saCeiteteeicinaete Meee ase 8 ee estie PACA Figures 154 and 155 Palpal segment 2 with no dorsal retrograde Spulyemlerteters cl eetoteiciae areleleletoeie cisicicicicicvclcleislerelelete eielclcleleler> Palpal dorsal basal margin forming a straight line; basal salience at right angles to long axis of palpi. Basis capituli rectangular. Scutum comparatively broad with large punctations. (Avian parasite)................++H. HOMDI HOODI Figures 136 and 1357 Palpal dorsal basal margin angular, forming a more or less well devel. oped, caudally—directed spur or recurved point. (Mammal parasites)’. 5ooo.afeseeecccssecees ce ce Trochanter I with a strong ventral spur. Each coxa with a spur reaching next coxa. Tarsi short, stout, ab ruptly tapered. (Common on ground. Squire Nagoya pire. Mossad mies cls de civccecsioesscesipecsceessctle HOULL Figures 140 and IZ1 trochanter i wichoul: Ventral! Spulsre «ciciclecicicle cicicle|eiclclce eo e\eieo ce) - 339 - 5. Coxae without overlapping, pointed spurs and with few hairs all shorter than coxal length. Palpal outline deeply concave laterally and with much reduced basal angle dorsally and ventrally; segment 3 ventrally with wide, bluntly rounded, short retrograde spur. (Rare parasite of HY PAKES|)Iso calc o.v.oeiiicisiele.cisie oicis sis'e sisie'lvieje sleisiavslieisicrnicion Me DEMURE: Figures 132 and 133 Coxae with a short, pointed spur on at least two pairs. Palpal outline laterally straight or very slightly concave (deeply concave only in ex. ceptional, small specimens), and usually with large pointed basal angle or spur dorsally and ventral ly; segment 3 ventrally with narrow, pointed, fairly long retrograde spur. (Commonly on carnivores, rare on Others.antmal’s)) io. cde ci slactieva tee ore eielnieloie1s ale avarsjorsis eraeresiessreeene © Scutum narrowly elongate (approximately twice as long as wide); punctations numerous, small, usually discrete; length from 2.3 m. to 3.8 mm., width from 1.2 m. to 1.9 m. Basis capi. tuli laterally usually diverging only slightly, cornua strong, slongate. Palpal lateral margins straight or slightly concave. Coxae each with a distinct, overlapping spur. Tarsi II to IV elongate, gradually tapering. (Common on canines; less common on other carnivores).....esccececceccecseseeeeeHs LEACHIT LEACHIT Figures 144, 145, and 140 = 340% Scutum wider (approximately 1.6 times as long as wide); punctations few to moderate, medium to large, shallow, nondiscrete; size smaller, length from 1.3 mm. to 2.2 mm., width from 0.8 mm. to 1.2 mm. Basis capituli laterally widely diverging, cornua usually short and wide. Palpal lat— eral margins usually concave. Coxae with a small but definite spur. Tarsi usually short, may be more or less humped and abruptly tapering. (Com mon on small carnivores; rare on Butellapes)) meals csieicic ce ielesicletine es osc cis icles H. LEACHII MUHSAMI Figures 150 an FEMALES Palpal segment 3 with basal margin strongly angled or spurred dorsally. Scutal width and length approximately equal. (Equatoria and Bahr El Ghazal; LAC ONMON) hata cle cistecieteecloetleereeicees cistic crs os boise Se ot moeicic cee Palpal segment 3 with basal margin dorsally forming a straight line WA TOMS pe OF LAN lola cisies s cieisic's c's c1s slo's slelais clelcicies cic wine Palpal segment 3 dorsally with basal margin sharply angled. Basis capi- tuli with strong cornua. Scutal length and width equal or very slight— ly longer than wide. Coxa I with inner basal angle produced to a small spur.............H. ACICULIFm Figures 130 an Palpal segment 3 dorsally with a retrograde spur. Basis capituli with small cornua. Scutum wider than long. Coxa I with small spur overlapping outer basal margin; no LNNET DAS iS PUT sioiers eleieiaicic)eielelo sleieie eciseeeicceccwetls PARMATA Figures 156 and 157 - 341 - 3. Palpal basal margin straight dor. sally, without angle or spur. Basis capituli almost straight laterally, cornua almost obsolete. Scutal length and width approx. imately equal. (Avian parasite)...........s...-H. HOODI HOODI Figures 138 and 159 Palpal basal margin angled or spurred dorsally. Basis capituli diverging laterally, cornua moderate to strong. (Mammal parasites)........... ace oie larerehore vs c1seic/suslavess stesiers Riese ee Palpi ventrally with a comparatively long, narrow spur basally, margin dorsally sharply pointed sublaterally; segment 3 ventrally with narrow elon gate, pointed spur. Coxae each with a strong spur and a few, short hairs; trochanter I ventrally with a strong spurlike ridge (that does not overlap trochantal margin); tarsi abruptly tapering, short, stout, somewhat humped, claw and tarsal length approximately equal. Scutum only slightly longer than wide, broadly rounded posteriorly; punctations few, shallow, mostly ante rior, (Ground_squirrel parasite)... 0s seseses co dontie HOU Figures 142 and 123 Combinations of characters differ as below. (Not normally ground-squirrel parasites) .avsescecesaees wines 3 shale sioieleleo oleleisielsloicisls’ loiclelalorcleisiels 5 a Wie Ds Coxae without overlapping spurs. Palpi without basal spur dorsally or ventrally; basal margin acutely angled sublaterally; segment 3 ventrally with very short, broad spur. Basis capituli with small cornua. Scutal length and width approximately equal but outline abruptly converging posteriorly; punctations scattered, shallow, moderate numbers of varying size. Tarsi tapering distally. (Hyreax parasites) Tare). ccccwsisisiviciciesicles/e «icreeieciietetles DEQUAERTI Figures 1327 and 135 Coxae with an overlapping spur on at least two pairs. Palpi usually with a definite basal spur dorsally or ventrally or both; segment 3 ventrally with an elongate, pointed, narrow ret— rograde spur; lateral outline straight or slightly curved; less commonly con cave. Basis capituli with strong cornua. (Common carnivore parasites; seldom-on Geher. animals) Ne ciecete-ecreisiere evolo ciclelahofitale rele aie'ers.s olele Siermtatieilewe a's 6O Scutum elongate, 1.25 longer than wide, gradually narrowing posteriorly; punc- tations comparatively small, numerous, discrete. Palpal spurs usually all large and definite, lateral margins straight or slightly curved. Tarsi elongate and tapering. (Commonly From Canines). cect cas ao cele s.ccisiane soc sleet p LEACHED LEACHTL Figures 146, 147, and 129 Scutum shorter, length and width approx imately equal, broadly rounded posterior-— ly; punctations comparatively moderate in size and numbers, shallow. Palpal basal spurs frequently reduced, lateral margins more or less concave. Tarsi short, usual_ ly abruptly tapered, sometimes humped. (Usually from smaller carnivores, sometimes on larger carnivores and ruminants; not COMMON ON CANINES) ..eccscscccsccceecssseccoctle LEACHII MUHSAMI Figures and 153 - 343 - “nr ee >a x of 3 nal L «sae eel f € f é = =3 2 131 Figures 128 and 129, <, dorsal and ventral views Figures 130 and 131, 9, dorsal and ventral views HAEMAPHYSALIS ACICULIFER an specimens PLATE AL - 344 - HAEMAPHYSALIS ACICULIFER Warburton, 1913. (Figures 128 to 131) THE SHARP.SPURRED HAREMAPHYSALID Bh! (Gp CE EQUATORIA PROVINCE RECORDS al Lotti Forest Praomys tullbergi sudanensis Apr 1 Torit Lemiscomys striatus massaicus Dec iL Zee Tort Ourebia ourebi aequatoria Apr il Torit Alcelaphus buselaphus roosevelt Apr 2 Atiambo (Alungwe ) Alcelaphus buselaphus subspp. Jun (SGC) ("several") Muragatika Kobus aeteeee subspp. Aug (Weber 1948) Jul il Nagichot domestic cattle Lotti Forest is at 4500 feet elevation and Nagichot is at 6500 feet elevation. Weber's (1948) report is the only previous one of this species from the Sudan. DISTRIBUTION IN THE SUDAN Bahr El Ghazal; All from tiang, Damaliscus korri tiang, 1953. 1c’, 269, 15 miles north of om, March, &. 1. M. Reid laste (with adults of Amblyomma variegatum). ct’, 7o9, Galual_Nyang Forest, July, N. A. Hancock Tegit. Material in HH collection. DISTRIBUTION H. aciculifer ranges across Africa in scattered highlands and forests from Bahr El Ghazal Province south to Mozambique. It has been transported on cattle to the Union of South Africa, where it has become established locally (Theiler 1945C). WEST AFRICA: NIGERIA (Unsworth 1949,1952. Gambles 1951). GOLD COAST (Simpson 1914. Nuttall and Warburton 1915). - 345 - CENTRAL AFRICA: CAMEROONS (Unsworth 1952). FRENCH EQUATORIAL AFRICA (Rousselot 1951; not repeated 1953B). BELGIAN CONGO (Tonelli- Rondelli 1930A. Bequaert 1931. Theiler and Robinson 1954). NOTE: According to Theiler (correspondence) , the record for Ruanda.Urundi by Santos Dias (1954D)is in error. EAST AFRICA: SUDAN (Weber 1948. Hoogstraal 195/B). KENYA (Lewis 19310 ,1932A,1934. Theiler 1945C). UGANDA (War- burton 1913. Tonelli-Rondelli 1930A. Mettam 1932. Theiler 1945C. Wilson 1950A,C). TANGANYIKA (J. B. Walker, unpublished; see HOSTS below). SOUTHERN AFRICA: MOZAMBIQUE (Santos Dias 1953B). UNION OF SOUTH AFRICA (Bedford 1932B,1934. R. du Toit 1942B,1947A. Intro. duced from East Africa: Theiler 1945C). NOTE: H. aciculifer does not occur on Madagascar (Hoogstraal 1953E). HOSTS Antelopes are the most common hosts of adult H. aciculifer. Other wild animals and domestic animals are rarely attacked. Rodents, small antelopes, and possibly hares appear to be the chief hosts of the immature stages. Domestic cattle have been reported as hosts in Uganda (Wilson 1950A,C), Kenya (Lewis 1934 and Theiler 1945C), and the Union of South Africa (Theiler 1945C). Lewis (1932A) noted a single spec. imen from a domestic goat in a Somali village between Karati Forest and Naivasha. Rousselot (1951) reported a female from a domestic dog in French Equatorial Africa. Antelopes hosts of adult ticks are the Uganda kob (Warburton 1913), reedbuck (Simpson 1914, Nuttall and Warburton 1915, Bed. ford 1932B, Theiler 1945C), bushbuck (Lewis 1931C ,1932A, Mettam 1932, Theiler 1945C), waterbuck (Lewis 1931C,1932A, Mettam 1932), various duikers (Mettam 1932, Lewis 1932A), Thomson's gazelle (Lewis 1934), oribi and Roosevelt's hartebeest (Equatoria Province records above), and tiang (Bahr El Ghazal Province record above). - 346 - Miscellaneous hosts of adults are buffalo (Lewis 1931C), serval cat (Bedford 1936, Theiler 1945C), mongoose (Theiler 1945C), and wild cat (Felis lybica group) (Hoogstraal, Kenya collecting). The probability that Mettam's (1932) records of this species from unidentified birds refer to H. hoodi hoodi should be considered. Nymphs have been found on the bushbuck, waterbuck, duiker, buffalo, warthog, and hare according to Lewis (1932A), but these records need checking for accuracy of identification. Lewis (1932A) reared H. aciculifer on hares. Examination of 49 Thom son's gazelles in Tanganyika yielded only a single nymph and no further specimens of this tick were found on many other game animals examined there (J. B. Walker, unpublished). Our Equatoria Province collections contain nymphs from a striped grassmouse, Lemniscomys striatus massaicus, and from an oribi, Ourebia ourebi aequatoria. A larva from a forest rat, Praeomys tullbergi sudanensis, in Lotti Forest (listed above) is apparently the only record extant for this stage in nature. BIOLOGY Life Cycle Lewis (1932A) reared the three-host H. aciculifer on laboratory hares in a minimum of 107 days. His data, are as follows: PERIOD DAYS Preoviposition 97099. tozicce) Oviposition to hatching 20 (22°C. to 25°.) Larval prefeeding period Uh Larva feeds 5) Premolting period 22 (21°C 5 to 2476.) Nymphal prefeeding period tl Nymph feeds 2 Premolting period 22 (21-6 ato. 25oG.) Adult prefeeding period il Adult (female) feeds fh 107 - 347 - Ecology In Kenya, H. aciculifer is found usually in forested areas at about 7500 feet elevation (Lewis 1932A). The writer's experience in Kenya and Sudan confirm that this is mostly a highland species (4500 to 8000 feet elevation) but that it is also present at lower altitudes (Torit, 2000 feet elevation). Records tend to indicate that where this species occurs at lower altitudes it is in more humid habitats but this subject requires further field study. Theiler (1945C) discussed the distribution of H. aciculifer on cattle in localized areas of northern and eastern Transvaal where it sometimes occurs on neighboring farms and at other times on distant isolated farms. The scattered distribution is believed to be due largely to the incidence of cattle importation from East Africa and to the ability of introduced ticks to maintain themselves in new areas. In South Africa, H. aciculifer survives in regions with from fifteen to fifty inches of annual rainfall but especially where thirty or more inches fall each year. These include subtropical evergreen and deciduous tree and thorn forest areas, open park. land areas in highlands, subtropical parkland areas, tall grass areas, and rarely short grasslands adjacent to highveld. H. aciculifer ranges from lowlands to highveld at 4500 feet elevation and 18 present only where winters are not severe, though it may survive where occasional light frosts occur. DISEASE RELATIONS Unstudied. IDENTIFICATION Males; Measure from 1.8 m. to 1.9 m. long, and from 1.1 m. to 1.3 mm. wide. They may be recognized among the African fauna by the long, needlelike spur of coxa IV. The basis capituli is rectangular with well developed cornua; the palpal outline (Fis- ures 128 and 129) is unique in the African fauna. The smooth scutum has small, shallow punctations; the lateral grooves may reach only the scutal midlength or they may be much longer. Soho = Certain variations in material at hand suggest the need for more specimens from various parts of Africa for further morpho logical study. Females: Engorged specimens measure from 6.0 mm. to 6.5 mm. long, and from 4.0 mm. to 4.5 mm. wide; unengorged, they are about 2.5 mm. long and 1.5 mm. wide. The scutal length equals or only slightly exceeds the width (0.8 mm. to 0.9 m. long, 0.7 m, to 0.9 mm. wide); the posterior margin is broadly rounded; cervical grooves are long, deep, and converging; punctations are small and inconspicuous. Palpal features are most distinctive (Figures 130 and 131). There is no dorsal projection on palpal segment 3. Coxae I to IV each have a small posterior spur and, in addition, coxa I has a small outer posterior spur; the spur of IV is short and wide thus differing greatly from that of the male. Palpal and scutal characters easily separate females from all other African species. The larva and nymph have been described and illustrated by Theiler (19450). - 349 - i 133 135 Figures 132 and 133, co, dorsal and ventral views Figures 134 and 135, 9, dorsal and ventral views HAEMAPHYSALIS BHQUAERTI Sudan paratypes PLATE XLI 5 0 - HAEMAPHYSALIS BEQUAERTI Hoogstraal, 1956(A). (Figures 132 to 135) THE EAST AFRICAN HYRAX TICK iN) On ct BQUATORIA PROVINCE RECORDS IMR Bde Imurok Heterohyrax brucei hoogstraali Feb (2) if Lui Procavia habessinica statin? May These records of H. bequaerti, from Torit District on the east bank of the Nile and from the far southwestern corner of the Sudan, are the only ones from this country. DISTRIBUTION H. bequaerti of Kenya and the Sudan is the most northern rep. resentative of three African hyrax-parasitizing ticks. The other two are H. orientalis Nuttall and Warburton, 1915 (= H. zambeziae Santos Dias, 195Z) of Nyasaland and Mozambique, and H. cooley Bedford, 1929, of the Union of South Africa. For further details, see Hoogstraal (1956A). EAST AFRICA: SUDAN (As Haemaphysalis sp. nov.: Hoogstraal 1954B. As H. bequaerti sp. nov.: Hoogstraal 1956A). UGANDA and KENYA (Hoogstraal 1956A). HOSTS Hyraxes: Heterohyrax brucei hoogstraali, Procavia habessinica slatini, and P. capensis meneliki (Hoogstraal 1950A). BIOLOGY H. bequaerti in all its stages is apparently strictly host— specific on hyraxes. A rather large number of hyraxes examined - 351 - in southern Sudan, Kenya, Yemen, Sinai, and the Eastern Desert of Egypt yielded no haemaphysalids other than the ones listed above. It would appear that this tick spends rather little time feeding and that, except possibly locally, hyrax-parasitizing haemaphysalids are rare in nature. DISEASE RELATIONS Unstudied. It is of interest to conjecture that this tick might be the vector of the piroplasm Echinozoon hoogstraali Garn ham, 1951, found in the blood of the Equatoria Province hosts. IDENTIFICATION Males: This is a small species, varying from 1.66 m. to 1.88 mm. in overall length and from 0.99 m. to 1.22 m, in width. It superficially resembles the ubiquitous H. leachii but may be easily differentiated from the subspecies muhsaml and even more easily from the subspecies leachii by the short, broad, bluntly rounded ventral spur of palpal segment 3, the greater lateral concavity of the palpi, the reduction of basal palpal spurs and of coxal spurs, the few, short hairs on the coxae, and other characters. The scutum has moderately numerous punctations, which are shallow, coarse, nondiscrete, mostly large, and widely scattered over the surface; cervical grooves faint to obsolete; lateral grooves enclosing first and second pairs of festoons (extension beside second festoon may be faint or obsolete), extending to anterior fourth of scutum. The coxae are only weakly armed with slight ridges and bear at most six small hairs; the tarsi taper gradually but may be more abruptly tapered in small specimens. The basis capituli diverges widely anteriorly and has bluntly pointed cornua about one fourth as long as the basis capituli. The palpi are short and salient with a weakly produced baso- lateral angle and a concave lateral margin; the ventral spur of segment 3 is short, wide and bluntly rounded. Females; In this sex, the capitulum is like that of the male except for the more elongate palpi and generally smaller cornua. = 2c The scutum is slightly longer than wide and posteriorly is rather abruptly narrowed and pointed; the cervical grooves extend to the scutal midlength; punctations are indistinct, shallow, large and medium size, few in number. Other characters recall those of the male. As in the male, the short, wide, bluntly rounded ventral spur of palpal segment 3 is a most important character in separating this species from H. leachii subspp., as are the short hairs of the coxae and the scutal shape. The larva and nymph have been described by Hoogstraal (1956A). - 353 - Figures 136 and 137, o, dorsal and ventral views Figures 138 and 139, 9, dorsal and ventral views HAEMAPHYSALIS HOODI HOODI ae meres an specimens PLATE XLII - 354 = HAEMAPHYSALIS HOODI HOODI Warburton and Nuttall, 1909. (Figures 136 to 139) THE AFRICAN AVIAN HAEMAPHYSALID LyrtNrepoctee BQUATORIA PROVINCE RECORDS 1 Torit Francolinus clappertoni gedgii Dec 2 Torit Sphenorhynchus abdimii Jan In Torit District, we closely examined over 200 each francolins and Uganda tufted guineafowl (Numida meleagris major), and many other birds, without finding itional specimens of this tick, It would appear, therefore, that H. hoodi hoodi is uncommon in the savannah of eastern Sudan. —~ DISTRIBUTION IN THE SUDAN Bahr El Ghazal: One 9 from tchagra shrike, Tchagra senegalo remigialis, Guar, Gogrial Subdistrict, February, 1353 dune Mohamed 1 Sayed legit (HH eorieceions: This and the Torit collections are the only Sudanese records of this avian parasite. DISTRIBUTION The African avian haemaphysalid ranges through tropical Africa and into southern Africa, but is possibly more common in western Africa and Uganda than elsewhere. A related subspecies, madagas— cariensis Colas-Belcour and Millot, 1948, occurs on Madagascar and other closely related species form a tight complex in the Oriental Region (Hoogstraal 1953E). WEST AFRICA: GAMBIA (Warburton and Nuttall 1909). GOLD COAST (Nuttall and Warburton 1915). FRENCH WEST AFRICA (Villiers 1955). SIERRA LEONE (Simpson's 1913 record of H. leachii from a bush shrike possibly refers to H. hoodi hoodi. Nuttall and War- burton 1915). PORTUGUESE GUINEA (Tendeiro 1047,1948,1951C,D, 1952A ,C Ary = 355 = CENTRAL AFRICA: CAMEROONS (Rageau 1953A,B. Numerous spec- imens seen by HH). FRENCH RQUATORIAL AFRICA (Specimens from Djambala, Moyen Congo; CNHM). BELGIAN CONGO: Bequaert (1931) states that while this spe— cies = not been found in the Congo, it can be expected to occur here. EAST AFRICA: SUDAN (Hoogstraal 1954B). UGANDA (Neave 1912. Nuttall and Warburton 1915. Mettam 1932. Lucas 1954. See HOSTS below). KENYA (Hoogstraal 195,C). SOUTHERN AFRICA: NYASALAND (Neave 1912. Nuttall and Warbur- ton 1915. Wilson 1950B). MOZAMBIQUE (As H. africana: Howard 1909A. Nuttall and Warburton 1915. Santos Dias 1952D,1953B, 19540; see HOSTS below. Hoogstraal 1954C. Theiler, correspondence; see HOSTS below). UNION OF SOUTH AFRICA (Bedford and Hewitt 1925. Bedford 1932B. Theiler, correspondence; see HOSTS below). HOSTS H. hoodi hoodi parasitizes birds exclusively, chiefly those kinds that are habitual ground feeders. See BIOLOGY below. "Fowls" (i.e. ?domestic chickens) (Warburton and Nuttall 1909). Domestic chickens (Tendeiro 1947. Lucas 1954). Burchell's coucal, Centropus senegalensis burchelli (Howard 1909). Western blue-headed coucal, C. monachus occidentalis (Ra geau 1953B. Others seen by HH). Senegal coucal, C. senegalensis (Tendeiro 1948, Villiers 1955, and Nuttall and Warburton Tole Guinea fowl, Numidia meleapris, plaintain eater, Gymoschizorhis leopoldi, and "partridge™ (Watiall and Warburton rey. Redwing Starling, Onychognathus (= Amydrus) morio (Bedford and Hewitt 1925). East African blue-eared nae Lamprocolius chloropterus elisabeth (Santos Dias 1952D). Clapper lark, Mirafa fischeri zombae (French Equatorial Africa specimens noted above). Falcon (Theiler, wnpublished). Tchagra shrike (Hoogstraal 1954B,C, Sudan record above). Double-spurred francolin, Francolinus bicalcaratus (Cameroons, J. Mouchet legit, HH det). opurfowl or francolin (Francolinus spp., Pternistis sp.) (Santos Dias 1953D, 19540, Hoogstraal 1952B,C, and sudan record above). White—browed =O scrub robin, hropygia leucophrys limpopoensis (Santos Dias 1954D). Abdim's stork (Hoogstraal 19525, an record above). Uganda hosts of specimens identified for the Museum of Com. parative Zoology are: grey hornbill, Lophoceros n. nasutus; Grant's crested francolin, F. sephaena grantil; yellow-beaked francolin, F. icterorhynchus; Abyssinian gonolak, Laniarius erythrogaster, and several individuals of both kinds of guinea fowl already noted from Equatoria Province, Sudan. Additional, recently obtained host data (Theiler, correspond- ence) is as follows: Centropus superciliosus from Uganda; "par- tridges" from East London, eastern Cape, and southern Transvaal, South Africa; Turdoides jardinei and Orthochagra senegal from Maringua, Mozambique; and Falco biramicus from Pietermaritzburg, Natal. The possibility that the record of a nymphal H. leachii muhsami from a tchagra shrike in Mozambique (Santos Dias 1954C ) refers actually to H. hoodi hoodi should be considered. The subject of parasitism of birds by ticks has been reviewed briefly by Schulze (19328). BIOLOGY Aside from indications that H. hoodi hoodi feeds exclusively on birds, chiefly on those that feed on the ground, and that all of its stages occur on a single host, little else is known of their biology. If domestic chickens were frequently attacked, more reports probably would have appeared in the literature. In Portugese Guinea, however, Tendeiro (1947) reports this parasite to be common on domestic chickens and in Entebbe, Uganda (Lucas 1954), a flock of chickens was found so heavily infested that a number of hosts died or were badly debilitated. The distribution of H. hoodi hoodi presumably is much more continuous in tropical Africa than present meagre records indicate. Phylogenetically, H. hoodi and related species, all of which close— ly resemble it, represents an old, quite static lineage. In Africa, - 357 - Madagascar, Asia, and outlying islands these ticks parasitize only birds. Related species infest primitive mammals such as hedgehogs (insectivores), and also reptiles. DISEASE RELATIONS DOMESTIC CHICKENS. Fatal anaemia has been reported. IDENTIFICATION Males. Palpi basally are widely salient, straight and lacking dorsal or ventral spurs; laterally they are sharply and narrowly re— curved basally and thence taper gradually to a narrow apex; segment 3 approximates segment 2 in length and medially bears a notably small and wide spur that is usually medially directed. The rectan. gular basis capituli has small but distinct cornua. The scutum is beset with a moderate number of fairly large, shallow punctations; lateral grooves include the first festoon and extend to the anterior third of the scutum; cervical grooves are shallow, concave, and extend more or less to the apical level of the lateral grooves. Coxae bear a small posterior spur, that of III may be obsolete and that of IV may be smaller than illustrated (Figure 137). Tarsi are moderately short and abruptly tapered; they bear a very small ver tral apical hook. Size varies from 1.3 mm. to 2.0 mm. long and from 1.0 m. to 1.4 mm. wide. Female palpi are like those of the male except that they are slightly less salient basally and more conical and elongate. The basis capituli is rectangular with very small cornua and a slightly concave basal margin; the porose areas are shallow and indistinct, The scutum is broadly oval, slightly longer than wide, and gradual. ly converging posteriorly; punctations are evenly scattered and rather large; cervical grooves are slightly concave and may reach the posterior third of the scutum. Tarsi taper somewhat more gradually than in males. The body becomes considerably extended when engorged. The larvae and nymph have been described and illustrated by Nuttall and Warburton 15). = 356 -— i42 140 143 141 Figures 140 and 141, @, dorsal and ventral views Figures 142 and 143, o, dorsal and ventral views imens HAHMAPHYSALIS HOUYI Sudan spec PLATE XLIIT - 359 - HAEMAPHYSALIS HOUYI Nuttall and Warburton, 1915. (Figures 140 to 143) THE WEST AFRICAN GROUND-SQUIRREL TICK ie eG? EQUATORIA PROVINCE RECORDS al Kapoeta Euxerus erythropus leucoumbrinus Dec 3 Torit Euxerus erythropus Teucoumbrinus Jan (2) LS Torit Euxerus erycnropus Teucoumbrinus Feb hropus 2 Torit Euxerus eryt Teucoumbrinus Mar 2) 26, AZ Torit Buxerus erythropus Teucoumprinus Dec (4) aL 3 —S aS all Latome Euxerus e Opus Teucoumbrinus Apr (SVS) al Yei Euxerus erythropus ?lacustris Apr DISTRIBUTION IN THE SUDAN Bahr El Ghazal: All from Galual_Nyang Forest, from five spec- jmens of Buxerus erythropus subspp., in 1953, by H. Hoogstraal: 65%, lg, 17 February; Too, 509, 4 nymphs, 19 February; AR, 16 February. Upper Nile: Bor, ex "Xerus rutilus", <0, 21 May 1909, H. H. King Tegit (S.c.C.). (This host name is a misidentification for Euxerus erythropus subspp.). Blue Nile: As o of H. calcarata: Roseires, from ground squirrel (Neumann 1910A); also records two oU that are probably H. houyi; Ch. Alluaud legit; cf. Hoogstraal (1955D). Kamisa, Binder River, 1c, lo, W. P. Lowe legit [PM (NH)_7. DISTRIBUTION H. houyi is a ground-squirrel parasite extending in a belt across the widest part of Africa from the Atlantic Ocean through French West Africa, Cameroons, the Sudan, and Uganda to the north western corner of Kenya, west of the Rift Valley. It is closely related to H. calcarata that parasitizes a different genus of cround squirrels in bast Africa east of the Rift Valley (Hoogstraal 1955D). See also HOSTS and BIOLOGY below. = 260% WEST AFRICA: FRENCH WEST AFRICA (Rousselot 1951,1953B. Hoogstraal 1955D. Villiers 1955). CENTRAL AFRICA: FRENCH EQUATORIAL AFRICA (Bate, "New Cameroons") (Nuttall and Warburton 1915. Hoogstraal 1955D). EAST AFRICA: SUDAN (As 9 of H. calcarata: Neumann 1910A. As H. houyi: Hoogstraal 1954B,1955D). SS” UGANDA and KENYA (Hoogstraal 1955D). HOSTS Ground—squirrels, Euxerus erythropus subspp. (All authors). King's specimen from "Xerus rutilus" at Bor (SGC) is based on a misidentification of the host. &. erythropus is the common ground. squirrel of West Africa, and of Northcentral and East Africa west of the Rift Valley. East of the Rift Valley it is replaced by Xerus rutilus subspp., parasitized by H. calcarata Neumann, 1902*. In Kenya, xerus is confined to hot lowlands and Euxerus to higher, arable mountains from 2000 feet to 6000 feet elevation, but mostly above 3000 feet. If, as now seems apparent, it is true that these two ticks are so host specific, this would seem to be a bolstering argument against lumping these two squirrel genera in one genus, as some mammalogists advocate (Hoogstraal 1955D). BIOLOGY Aside from the fact that all stages may be found on a single ground-squirrel, little is known concerning the biology of H. houyi. This tick and its host inhabit savannah country with few or scattered trees, and upland grasslands. Along the southern border of the squirrel's range it extends into forested districts, but only in tongues of psrassland with scattered trees between thicker forest. As already stated under HOSTS, in Kenya, where the two host genera and the two related tick species occur near each other, the host of H. houyi is confined mostly to arable up. lands and that of H. calcarata inhabits warmer and more arid low- lands. SR SE ET tS *The record of H. calcarata from Dahomey (Villiers 1955) undoubtedly is based on misidentification. = Jot In the Galual_-Nyang forest area of Bahr El Ghazal, each of five host specimens examined was infested. In Torit District of Equatoria, a third of the 27 hosts examined yielded specimens of H. houyi. DISEASE RELATIONS Unstudied but potentially important. IDENTIFICATION The following characters easily distinguish males among the Sudan haemaphysalid fauna: strong ventral spur on trochanter I; all coxae with distinct spurs; tarsi short, robust, and abruptly tapered; palpi widely expanded basally, without a developed dorsal spur basally, with basal spur ventrally and spur from segment 3 ventrally; basis capituli strongly diverging anteriorly and with moderate cornua; dentition 4/4; scutum with long, deep lateral grooves enclosing first festoon, and few, scattered, shallow, inconspicuous punctations of mixed sizes; size ranges from an overall length of 1.71 mm. to 2.15 mm. and width of 0.99 mn. to 1.20 mn. Females are also easily recognized by the raised spurlike, nonprojecting ventral ridge of trochanter I, coxae and tarsi almost exactly like those of male; palpi essentially like those of male but larger and more elongate, basis capituli short, wide, and with prominent cornua and anteriorly diverging lateral mar. gins; dentition 4/4; scutum only very slightly longer than wide and broadly rounded posteriorly, with few, shallow, scattered punctations of various sizes mostly on anterior half. The size is somewhat greater than that of males. = 362= Figures 144 and 145, &, from domestic dog (Kajo Kaji) Figures 146 and 147, 9, from domestic dog (Kajo Kaji) Figures 148 and 149, o and 9, from civet (Obbo) HAEMAPHYSALIS LEACHII LEACHII an Specimens PLATE XLIV =) 903 = HAEMAPHYSALIS LEACHII LEACHII (Audouin, 1827). L N* o 4 5 2 2 12 5) 2 als A 2 il: Abe dial ava 5) 15 5) 6 8 5 4 5 4 (Figures 144 to 149) THE YELLOW DOG-TICK BQUATORIA PROVINCE RECORDS Torit Crocidura nyansae toritensis Lotti Forest Praomys Eilibergi Sudanensis Torit Kapoeta Juba Nimule Torit Yei Obbo Torit Torit Torit Torit Torit Yubo Torit Gilo Katire Katire Torit Torit Kajo Kaji Beringi, Yei River Bundle, Kheirallah Tatera benvenuta benvenuta Arvicanthis niloticus Jebelae ae Burrows of A. i: jebelae Acomys hystrella US eumia a. albicauda Tchneuma a. 1eauda Civettictus civetta congica Civettictus civetta congica : eS Civettictus civetta congica Canis aureus Soudanicus Canis aureus soudanicus ee : te Ae soudanicus anis mesomelas elgonae Panthera Leo Leo the EA hy pated Domestic dogs Domestic dogs Domestic dogs Domestic dogs Domestic dogs Domestic dogs Domestic dog Domestic dog Feb Apr (2) Dec Apr Dec (4) Mar Apr Apr Apr Feb Jun Apr (2) Nov Dec (2) Mar Mar Dec (4) Jan (2) Oct (2) Jan (2) Dec ie Dec (2) Feb (SGC) Mar (SGC) *The subspecies of these nymphs is not entirely certain. - 364 - DISTRIBUTION IN THE SUDAN King (1926) listed Equatoria, Bahr El Ghazal, Upper Nile, Blue Nile, Kordofan, Khartoum, and Kassala Provinces, and noted that although H. leachii (subspecies not differentiated) has a wide range in the Sudan, it is a relatively rare species here. We now know that H. 1. leachii occurs in every Province of the Sudan. In most areas it Is probably fairly common, though usually only on medium-size carnivores, especially jackals, foxes, and domestic dogs. The following is Sudanese material seen: Bahr El Ghazal: Wau (Domestic cat; SVS. Domestic dogs; HH). Galual-Nyang Forest (Domestic dog; HH). = Upper Nile: Akobo Post (Lion; SGC). Sobat (Domestic dog; . Blue Nile: Magangani (Caracal c. nubicus; MOZ). Kordofan: Delami (Domesticated wild cat; SGC). Umm Dona, (Mustelid; Sac). Darfur; Fasher (Domestic dogs; SVS). Khartoum: Khartoum, near (Vulpes a. aegyptiaca; HH). (Domestic dogs; Balfour 1911F). Kassala: Port Sudan (Domestic dogs; HH). Northern: Wadi Halfa and Atbara (Vulpes a. segyptiaca; HH). DISTRIBUTION Haemaphysalis leachii leachii is a ubiquitous tick of tropical and southern Africa. In Egypt, 1t occurs in and at the edge of the Nile Valley and Delta almost to the Mediterranean coast. It has been reported to range along the Maditerranean littoral at least as far west as Algeria but these records require careful checking for accuracy of identification. H. 1. leachii is fairly Sooo common in the mountains of the Yemen in southern Arabia (Sanborn and Hoogstraal 1953; Hoogstraal, ms.), which is an outlying part of the Ethiopian Faunal Region. It is not known from Madagascar (Hoogstraal 1953E) and I have not seen it in Turkey. In Euro H. leachii has been said to occur in Yugoslavia (Oswald 1938) ard in Greece (Oswald 1938, Pandazis 1947) but these records are considered questionable. A German specimen found on a migrant stork from Africa was mentioned by Schulze (1937A). Soviet records (Olenev 1928) refer to different spe- cies and Pomerantzev (1950) does not consider it to be a member of the Russian fauna. The exact relationships of Oriental Faunal Region forms ascribed to this species by Nuttall and Warburton (1915) are at present under study. Apparently most, except the subspecies indica Warburton, 1910, represent different species. Numerous species, obviously derived from an H. leachii prototype, range throughout Asia and its nearby islands, the Near East, and the Madagascan archipelago. It appears from remarks by Dumbleton (1953), that this tick has not been found in New Zealand since the original record by G. E. Mason (1921), that Mason's ticks may have been a different species. Mason's distributional concepts for this species most probably require revision. Note In the following list, all available references to "H. leachii® are noted, though it is usually impossible to deter- mine whether authors are referring to the subspecies leachii or muhsami. I have seen actual specimens of tne subspecies leachii from all geographical areas and from almost all political territories listed below. The following records are those for continental Africa and Arabia. As stated above, it is uncertain whether the subspecies leachii occurs outside of the Ethiopian Faunal Region and parts of the Mediterranean Subregion of the Palearctic Region. Analysis of studies of the distribution of this tick will be presented subsequently. = o00= NORTH AFRICA: EGYPT (Savignyi 1826. Audouin 1827. Neumann 1911.” Mason 1916). The following North African records require checking for accuracy of identification: LIBYA: -Tonelli-Rondelli (1926B). Franchini (1927,1929A,E). TUNISIA: Colas-Belcour and Rageau (1951). See Hoogstraal (1955B). ALGERIA: Neumann (1897) from a "nightingale" and from grass. Some of the H. leachii reported by Neumann (1897) were later (1905) described by him as H. numi- diana, which is a synonym of H. erinacei Pavesi, 1884.7 WEST AFRICA: NIGERIA (Simpson 1912A,B. Nuttall and Warbur- ton 1915. Johnston 1916. Pearse 1929). TOGO (Neumann 1901,1911). GOLD COAST (Simpson 1914. Nuttall and Warburton 1915. Beal 1920). FRENCH WEST AFRICA (Rousselot 1951,1953B. Villiers 1955). PORTU- GESE GUINEA (Tendeiro 1948,1951A,D,1952A,B,C ,D,F,1953,1954). SIERRA LEONE (Simpson 1913. Nuttall and Warburton 1915. Entomological Reports 1916). GAMBIA (Simpson 1911. Nuttall and Warburton 1915). CENTRAL AFRICA; CAMEROONS (Neumann 1901,1911. Ziemann 1912A. Nuttall and Warburton 1915. Rageau 1951,1953A,B. Rousselot 1951, 1953B. Dezest 1953). FRENCH BQUATORIAL AFRICA (Nuttall and War. burton 1915. Fiasson 1943B. Rousselot 1951,1953B. Giroud 1951. Giroud and LeGac 1952). LIBERIA (Bequaert 1930A). BELGIAN CONGO and RUANDA_URUNDI (Newstead, Dutton, and Todd 1907. Massey 1908. Neumann 1911. Nuttall and Warburton 1915, 1916. Seydel 1925. Schwetz 1927A,B,C. Bequaert 1930A,B,1931. Tonelli-Rondelli 1932E. Wanson, Richard, and Toubac 1947. Fain 1949. Giroud and Jadin 1950,1955., Giroud 1951. Jadin and Giroud 1951. Jadin 1951B. Schoenaers 1951A,B. Rousselot 1931, 1953B. Theiler and Robinson 1954. Santos Dias 1954D. Van Vaerenbergh 1954). EAST AFRICA: SUDAN (Balfour 1911F. King 1911,1926. Nuttall and Warburton 1915. Hoogstraal 1954B). ETHIOPIA (Neumann 1902B,1922. Nuttall and Warburton 1915. Tonelli-Rondelli 1930A. Stella 1938A,1939A,1940. Charters 1948. D'Ignazio and Mira 1949). ERITREA (Nuttall and Warburton 1915. Tonelli-Rondelli 1930A. Stella 1938A,1939A,1940). ITALIAN SOMALI. LAND (Paoli 1916. Franchini 192%. Tonelli-Rondelli 1935. Niro 1935. Stella 1940). FRENCH SOMALILAND (Stella 1940). Ol a= KENYA (Neave 1912. Nuttall and Warburton 1915. Neumann 1922. Anderson 1924A,B. Lewis 1931A4,B,C ,1932B,1934,1939A. Roberts and Tonking 1933. Kauntze 1934. Roberts 1935. Loveridge 19364. Dick and Lewis 1947. Weber 1948. Heisch 1950B. Binns 1951,1952. Wiley 1953. Weyer 1955). UGANDA (Neave 1912. Nuttall and Warburton 1915. Neumann 1922. Bequaert 1930B. Loveridge 1936. Carmichael 1942. As H. leachii humerosoides: Theiler 1943B. Wilson 1950). TANGANYIKA (Neumann 1907C,1910B. Neave 1912. Nuttall and Warbur- ton 1915. Morstatt 1913. Loveridge 19234. Bequaert 1930A. Allen and Loveridge 1933). SOUTHERN AFRICA: ANGOLA (Neumann 1901. Gamble 1914. Nuttall and Warburton TOL5. Sousa Dias 1950. Santos Dias 19500. Theiler and Robinson 1954). MOZAMBIQUE (Howard 1908. Nuttall and Warbur- ton 1915. De Meillon 1942. As H. leachii humerosoides: Theiler 1943B. Santos Dias 1952H,1953A,B,C,1054H,1055A. Theiler and Robinson 1953A). NORTHERN RHODESIA (Neave 1912. Nuttall and Warburton 1915. Morris 1933 ,1934,1935 ,1936,1937,1938,1939,1940. As H. leachii humerosoides: Le Roux 1947. Theiler and Robinson 1954). SOUTHERN ; ch 1903. Edmonds and Bevan 1914. Nuttall and Warbur. ton 1915. Jack 1921,1928,1937,1942). NYASALAND (Old 1909. Neave 1912. Nuttall and Warburton 1915. De Meza 1918. Wilson 1950B). BASUTOLAND (Scarce: Theiler and Robinson 1953A). SWAZILAND (Theiler and Robinson 1953A). SOUTHWEST AFRICA (Tromsdorff 1914. Sigwart 1915. Absent in Southwest Africa: Theiler and Robinson 1953A). UNION OF SOUTH AFRICA (Neumann 1897,1911. Lounsbury 1901,1902A,1904A. Howard 1908,1909A. Galli-Valerio 1909. Spei- ser 1909. DOnitz 1910B. Moore 1912. Van Saceghem 1914. Nuttall and Warburton 1915. Bedford 1920,1926,1927,1931B,1932B,1936. A. Theiler 1921. Cowdry 1925C ,1926A,1927. Curson 1928. Cooley 1929, 1934. Bedford and Graf 1934,1939. Pijper and Crocker 1938. J.H.S. Gear 1938. Brumpt 1938). J. Gear 1939,1954. Gear and Douthwaite 1938. Mason and Alexander 1939. Gear and De Meillon 1939,1941. De Meillon 1942. du Toit 1942B,1947A. Theiler 1943B. Cluver 1944. Neitz and Steyn 1947. Theiler and Robinson 1953A). OUTLYING ISLANDS: ZANZIBAR (Neave 1912. Aders 1917). Not known from Madagascan archipelago (Hoogstraal 1953E). - 368 . ARABIA: YEMEN (Sanborn and Hoogstraal 1953. Hoogstraal, ms.). HOSTS H. leachii leachii, in the adult stage commonly parasitizes domestic dogs. ts local incidence on dogs varies from greater to less than that of the kennel tick, R. s. sanguineus. Locally, R. simus simus is sometimes also a common parasite of dogs. On wild animals, H. 1. leachii is frequently numerous on the Canidae (foxes, hunting dogs, and jackals). It is, in comparison with the subspecies muhsami, rare on the various families of smaller carnivores such as the viverrids, which are tropical Africa's most common carnivores. On larger Felidae, lions, leopards, cheetahs and the like, the subspecies leachii may occur in either larger and smaller numbers than muhsami but present data do not suggest that the large cats are hosts of preference. Records of H. 1. leachii from smaller cats and from domestic cats are rare indeed. Domestic animals, other than dogs, are parasitized only exceptionally. Under a very few local conditions cattle may be attacked. Possibly tribal customs, in which man, cattle, and dogs sleep in the same hut or compound, account for these instances. Larvae and nymphs usually parasitize common field rodents, especially Arvicanthis and Mastomys, in their nests. They are said also to feed on domestic dogs (see BIOLOGY below). Rarely one finds a few nymphs, along with considerably larger numbers of adults, on wild canines. This would indicate that, under the influence of some yet unknown factors, nymphs have left rodent burrows to feed elsewhere, or that an apparently small propor. tion of the nymphal population does not feed on rodents. Inasmuch as it is impossible to distinguish which of the two subspecies of H. leachii most authors are referring to, the only data that may be used in this section are from the present observations and those of the very few recent students who have differentiated their material. A more exact study of available data will be presented in a subsequent report. = Oo BIOLOGY Life Cycle Nuttall's (1913B) and Nuttall and Warburton's (1915) summary of Nuttall's and of Lounsbury's (1901,19024,1904A) observations on rearing H. leachii (most likely H. 1. leachii) are essentially as follows: This tick requires three hosts upon which to feed during its larval, nymphal, and adult stages. About a week after molting each stage readily attaches to the host, which under experimental conditions may be a number of different animals, jackal, dog, ferret, hedgehog, goat, or rabbits It appears to be immaterial upon which of these hosts the ticks feed. Larvae and nymphs feed for three to seven days (two to three days; Lounsbury), occasionally longer. Females attach for eight to sixteen days. Males may remain upon the host for many weeks. Air temperature, within the limits observed (9°C. to 23°C.), appears to exert little or no influence upon the time ticks re. main upon the host, "the warmth from the animal being doubtless sufficient to keep the ticks active". The time required for metamorphosis is influenced by temper ature. Larvae hatch after 26 to 37 days at 20°C. or after 58 to eighty days at 12°C. to 13°C. Nymphs emerge, as a rule, after thirty to forty days. Adults emerge after fifteen or sixteen days at 24°C. to 26°C., but require up to seventy days at 14°. The unfed tick survives for long periods under favorable conditions. In small corked bottles maintained at about 12°., larvae are still active after 169 days, nymphs after 52 days, and adults after about 210 days. Males and females placed simultaneously upon the host scatter but in two or three days both sexes are found attached in close proximity to each other. Copulation occurs upon the host (HH observation). Lounsbury saw marked males detach and reattach close to females. A male may mate with more than one female. After a replete female abandons the host, the interval before egglaying commences is markedly influenced by temperature. Fe. males held at 23°C. begin oviposition after three to five days; at 16°C. to 21°C. after fourteen to eighteen days; at lower tem - 370 — peratures after 24 to 60 days. Whereas an occasional female dies as soon as oviposition is completed, others may survive for a few days or, exceptionally, for a month. One tick deposits from 2400 to 4300 eggs. In nature, the yellow dog—-tick doubtless may produce two generations a year. Lounsbury reared three generations a year in an incubator. Taking average figures for ticks raised under favorable conditions, the cycle may be completed in 123 days, as follows: PERIOD DAYS Preoviposition TINe3°CS) Oviposition to hatching 30 (20°C...) Larval prefeeding period 7 Larva feeds 5 Premolting period Zi CL7"C 5) Nymphal prefeeding period 7 Nymph feeds 5 Premolting period ea 22°C%) Adult prefeeding period 7 Adult (female) feeds We 23 Ecology The distribution of H. 1. leachii has been determined for South Africa by Theiler and Robinson (1953A). The most important factor in limiting this tick's spread there is increasing aridity. Twenty inches of annual rainfall, irrespective of vegetation type, appears to be the critical level there. However, in northern Sudan, where rainfall is absent or considerably less than ten inches annually, H. 1. leachii is still fairly common on foxes. In the Nile Valley of Egypt, where rainfall is nil to ex ceedingly low, the tick thrives. But, it should be stressed, the microhabitats of its larval and nymphal host, the grass rat, Arvi- canthis n. niloticus, are more humid than elsewhere, situated as - 371 - they are beside irrigated, cultivated fields or in dykes. We have never found these ticks in burrows in the desert, even on the Mediterranean littoral where burrows are frequently patently damp. Foxes that pick up newly-molted adult ticks, probably from vegeta tion near grassrat burrows when they forage in cultivated areas at night, retreat to very dry desert caves and dens to rest by day. In these situations, evaporation from the host skin may be the factor that allows the parasite's survival. A comparative study of the survival of Northeast African and South African populations under local conditions of humidity and temperature should be of considerable interest. Theiler and Robinson (1953A) have also found that H. 1. leachii does not occur in those parts of South Africa with over sixty days of heavy frost per annum. It is generally absent from the arid Karroo except where grasses are present. Altitude does not effect the distribution of the yellow dog-tick within the limits of crit- ical frost days noted above. Variability of incidence in various zones of South Africa is also discussed. According to Lewis (1939A), H. leachii (probably including both subspecies ~ HH) occurs in all districts and altitudes of Kenya but seems to prefer the shelter of dense shrub and grassy woodlands. In some areas it is more common on dogs than is R. s. sanguineus ° Theiler and Robinson (loc. cit.) state that the immature stages of H. 1. leachii may feed on dogs. Our experience in East Africa, Egypt, and Arabia indicates that nymphs are very rarely found on roaming wild carnivores such as mongooses, civets, and jackals, but that larvae and nymphs frequent rodent burrows. Onderstepoort records (Theiler, correspondence) show one hundred collections of nymphs from murid rodents, one from cattle, one from shrews, one from Felidae, one from hares, eight from elephant shrews, three from mustelids, five from springhaas, four from squirrels, three from mongooses, and one from civet. It is obvious that many factors governing the life cycle and possible variability in host preference of immature stages remain to be determined from field studies. Roberts (1935) found larvae and nymphs in the Nairobi area common on various field rodents and in their nests. These are - 372 - the same as those listed for R. s. simus (page 743). The writer's experience in other parts of Kenya in general confirms Roberts? findings. However, Roberts observed that nests of Mastomys (= Mus) coucha near the surface of the ground rather than deeper nests are preferred by H. leachii, but I do have numerous records from deep. er nests of grassrats. This factor also requires further study (See R. s. simus, page 746). It is of some interest to note that all specimens that have been reared in our laboratories from nymphs from rodent nests in Kenya, the Sudan, and Egypt have been sub. species leachii. The chalcid wasp parasite Hunterellus hookeri has been bred from nymphs in South Africa (Cooley 1929,195Z). This subject is further discussed under R. s. sanguineus (page 710). DISEASE RELATIONS MAN: Boutonneuse fever (Rickettsia conorii). Experimental evidence indicates efficiency as a vector of Rocky Mountain spotted fever (Rickettsia rickettsii). MAN AND ANIMALS: Q fever (Coxiella burnetii). DOMESTIC DOGS: Canine babesiosis (Babesia canis). ?7DOMESTIC CATS: Feline babesiosis (Nuttallia felis). ?JACKALS: Canine babesiosis (B. canis). REMARKS A gynandromorph of H. leachii (probably subspecies leachii) has been described and illustrated by Santos Dias (19530). The misshapen specimen of H. leachii described and illustrated by Nuttall (1914A), and widely quoted by subsequent authors, refers to the Asiatic subspecies indica Warburton, 1910. Very slightly misshapen specimens, due to injury, of both African subspecies have been described and illustrated by Santos Dias (1955A). The measurements and increase in relative size from stage to stage have been studied by Campana-Rouget (1954), apparently from data in Nuttall's Monograph. - 373 - It should be noted that some gradation appears between the two African subspecies of H. leachii and that a few specimens do not conform strictly to the criteria for one or the other form. The third subspecies, indica Warburton, 1910 of southern Asia, is more like the subspecies muhsami than like the subspecies leachii, and is distinguishable from both by minor but apparently constant and valid characters. Dr. G. Theiler and the writer for several years have been collaborating on a morphological study of considerable series of this species from a variety of hosts and localities. The results, with complete data, will be presented in a separate report. The variety humerosoides, common on canines, informally proposed by Theiler (19238) for the large, narrow, elongate form with extreme ventral projection of spurs, appears from this study to be an extreme body form of the somewhat variable H. 1. leachii and not a separate morphological or biological subspecies. In numerous long series of specimens from single hosts, gradations from this to less extremely narrow and elongate forms occur. There are, however, suggestions that the extreme form is a reflection of particular host factors, and application of some of the more com plex aspects of newer taxonomic concepts may eventually justify the name humerosoides. IDENT IFICATION Males. This long, narrow tick has tarsi II to IV gradually tapering; punctations numerous, mostly small, and discrete; palpi obtusely angled and widely triangular, widest at level of basal third, with lateral margin straight or very slightly convex but almost never concave; basally both dorsally and ventrally forming a conspicuous and usually strong spur just laterad of the point of insertion; palpal segment 3 with a retrograde spur that is long and tapering; basis capituli with lateral margins varying from almost parallel to somewhat divergent anteriorly and with cornua that are usually large and pointed; coxae always with a distinct basal spur overlapping the basal margin and with a number of long, conspicuous hairs. This combination of characters must be considered in separating males from those of other species and from the subspecies muhsami. - 374 - The scutum varies from about 2.3 mm. to 3.8 mm. long and from 1.2 mm. to 1.9 m. wide; average specimens are about 2.6 mm. long by 1.3 mm. wide. This length-width ratio is important in comparing this subspecies with muhsami, though a few intergrade specimens, with respect to this feature, do occur. The punctations, always numerous and mostly comparatively small, are usually discrete; they cover the entire dorsum including lateral areas and festoons but frequently are reduced in the narrow, elongate area correspond— ing to the posterior median groove of rhipicephalids. The long, narrow lateral groove encloses the first one or two pairs of fes. toons; the closely approximated, arched cervical grooves usually extend to the anterior level of the lateral grooves. The scutal surface is more or less arched. The palpi are notable for their wide, obtusely angled form. The lateral margin, either straight or very slightly convex in outline, distinguishes this subspecies from muhsami, but, rarely, a similar form occurs on ticks with the short, broad scutal type of muhsami. The recurved basal margin is typically broken both dorsally and ventrally by a strong spur just laterad of the point of insertion; while this spur is usually accentuated in large, narrow, elongate specimens it is surprisingly reduced in some in. dividuals of this type. The ventral retrograde spur of palpal segment 3 notably is consistently strong, overlapping the base of segment 3, and narrow and tapering. Segment 3 is about half as long as segment 2. The basis capituli, typically, is elongate with strong, tapered cornua and with lateral margins slightly divergent anteriorly, but the length-width ratio and size and shape of the cornua is surprisingly variable, even in specimens in which the general appearance would otherwise lead one to expect that these features would be typical, and the degree of divergence of the lateral margins is also somewhat variable. The hypostome has 4/4 or 5/5 dentition. The coxae are notable for the basal spur that overlaps the basal margin and for the presence of twelve to twenty long hairs on each (hairs may be broken or rubbed off in old or carelessly collected or preserved material). The size and position of these spurs always approximate those illustrated herein and are important in distinguishing this species from some others. In newly molted or fresh specimens, the numerous long hairs are a very character- istic feature of this species. The elongate tarsi taper gradually - 375 - apically and bear a small pad and claw; the claw curves distad of the apex of the pad. Females. This sex closely recalls the male and while it is equally aS variable it appears to be less frequently confusing with the subspecies muhsami. The elongate scutum, from one fourth to one third longer than wide, posteriorly tapers gradually to a more or less narrow point. Scutal punctations are mich like those of the male, and while they are frequently somewhat larger and less numerous than those of the male they are distinguishable from the consistently large and sparse punctations of muhsami. The cervical grooves gradually converge to the scutal midlength and thence diverge towards the posterolateral margins but do not reach these margins. The palpal outline is like that of the male except that it is more elongate, the length of segment 3 more nearly equalling that of segment 2 than it does in the male; and the basal spur ventrally is absent or extremely reduced in the form of a bluntly rounded projection. The lateral margin, which as in the male is typically straight or slightly convex, is actually more readily and definite- ly usable as a diagnostic character because of its greater length; while this margin is very slightly concave in some specimens these are unusual. The basis capituli is definitely wider and shorter than that of the male and bears shorter cornua. Coxal and tarsal characters are like those of the male; in spite of some variation they are not likely to be confused with most specimens of the related subspecies. Va considering the female coxal spur as minute and the tarsus as "stout", Nuttall and Warburton (1915) mst have been referring to specimens of muhsami./ The body form of unengorged females is typically elongate and comparatively narrow, as in males, though the overall size is some what larger. Engorged females may become so large in the latter hours of feeding that superficially they resemble typical boophilid females. The larvae and nymph of this species, but not definitely refer. able to this subspecies, have been described by Nuttall and Warbur- ton (1915). = BSS Figures 150 and 151, o&, dorsal and ventral views Figures 152 and 153, 9, dorsal and ventral views HAEMAPHYSALIS LEACHII MUHSAMI Sudan Specimens from White-tailed Mongoose PLATE XLV = 377 = HAEMAPHYSALIS LEACHII MUHSAMI Santos Dias, 1954(E) (= H. LEACHII INDICA (in Africa) or H, LEACHIT near INDICA of authors). (Figures 150 to 153) THE YELLOW SMALL-CARNIVORE TICK i. Ni =Oe cc; FQUATORIA PROVINCE RECORDS Oss Crocidura nyansae toritensis Feb ale Torit Atelerix pruneri owen. Feb 2 Tarangore MAtelerix pruneri owen Jun Zee ea Loree Mellivora capensis abyssinica Jan 1 lorie Canis ee elgonae Dec 3-1. » Torit Canis aureus soudanicus Apr (2) 5 Obbo Civettictis civetta congica Apr 2. Jee Lorit Civettictis civetta congica Feb 9 Torit Civettictis civetta congica Jun (2) £ 9 Toriit Civettictis civetta congica Jul 1 eZee Lor Genetta ti eri na aequatorialis Feb 2 15 Kapoeta Herpestes sanguineus sanguineus Apr Leo torit Teleomie aTelcouta seer oe Jan LS Lorik Tchneumia icauda albicauda Mar 2: LOSS: — Torit Tchneumia icauda albicauda Apr (3) LV aay 329 ver Tchneumia albicauda albicauda Apr (2) al Torit elis ca ugandae Nov al Kapoeta Lepus capensis subsp. Apr ak Ikoto e capensis crawshayi Feb aL Ikoto Rhyncho are guenthert smithii Dec al Torit omestic dog a aes DISTRIBUTION IN THE SUDAN Bahr El Ghazal: 12%, 499, black-legged mongoose, Galual-Nyang Forest, 27 May 1055, E. T.M. Reid legit. «6, same host and collec. tor, Yirol, 22 January 1954. ‘5, Atelerix eri oweni, Galual. Nyang Forest, 24 February, 1953, H. Hoogstr egit. Tc’, lo, - 378 . Atelerix cael oweni, Majan Yom, 2 May 1953, E. T. M. Reid legit. Trodent burrowing in peat mound", Waynjok, N. W. Goerials April, 1953, W. Dees legit. ¢ tien, Damaliscus korri tiang, Galual_Nyang Forest, April, : 53, BE. T. M. Reta legit. ao, eee 36 miles south of Yirol, 18 January, 1953, . M. Reid legit. lo, domestic cat, Galual_Nyang Forest, March 1953, an » same host, Wau, October, 1953, both SVS. Blue Nile: ‘5%, 309, "mongoose", Wad Medani, 29 November 1950, D. JT Lewis legit (Scc). Khartoum: 4c’, "fox", Khartoum, 9 January 1918, R. Cottam legit (SC). DISTRIBUTION The subspecies muhsami occurs in all areas of the Ethiopian Faunal Region, including the mountains of the Yemen in south. western Arabia. We have not seen it in Egypt. The data will be published subsequently in a series of reports on Africa haema- physalids. HOSTS The subspecies muhsami is especially common on small carni- vores such as mongooses, genets, civets, and wild cats. It seldom attacks wild or domestic canines or wild antelopes. Usually smal- ler numbers are found on mle rats, shrews, hedgehogs and hares; ° the possibility that certain of these may represent separate forms is being studied. Full data will be presented in the report men- tioned under DISTRIBUTION above. BIOLOGY This subject requires study, especially in relation to that of the subspecies leachii. DISEASE RELATIONS Unstudied. os WHeNe REMARKS As already indicated, Theiler and the writer have been studying variation in this species for some years and the final report is nearing completion. Recent material sent by Santos Dias to Theiler for identification, and returned with the note that this was typical of what we were provisionally referring to as "H. near indica®™ pending completion of our studies, was utilized as the type series for the "species" muhsami. These specimens and their description and illustration correspond with what we have been considering as the "H. leachii near indica" of Theiler (1943B). Subsequent Theiler- Santos Dias correspondence, however, indicates that the latter worker considers muhsami as a separate species, separate and distinct from "near indica", Recently (November 1955) Dr. J. Bequaert has kindly sent me his paratype specimens of "H. mbhsami" and study of these confirms the already mentioned Theiler viewpoint. Santos Dias (1954C) reports a nymph of "H. mhsami" from a tchagra shrike in Mozambique. The likelihood that this is actual ly a specimen of H. hoodi hoodi should be considered. IDENT IFICATION H. leachii muhsami in some instances intergrades with H. l. leachii and these specimens may be difficult or impossible To — separate to subspecies. Criteria for separating this species from others are established in the key and under IDENTIFICATION of H. leachii and in the latter section the characters differen. tiating the two subspecies are also noted. Results of a long- term study of this subject will be presented subsequently. Here, only a brief resume of the characters separating H. leachii mihsami from the nominal subspecies is provided. ~ Males. These ticks are smaller and their scutal outline is wider than that of the subspecies leachii. Scutal punctations are only moderate in number and are generally fairly large and shallow. In outline, the palpal outlines of the two are quite similar except that the lateral margin of mbhsami is slightly concave though in exceptional specimens it may be straight and even more uncommonly it may be very slightly convex. The ventral retrograde spur of palpal segment 3 notably is like that of leachii, =) o50r= but in muhsami the basal spurs may be more reduced. The basis capitull Ls short and wide, with lateral margins widely diverging anteriorly, and the cornua are usually smaller and weaker than those of leachii. The coxal spurs are comparable with those of leachii, an important criterion for separating this subspecies from some other equally small, not otherwise greatly differing species in Africa. The scutal size varies from 1.3 mm. to 2.2 mm. long and from 0.8 m. to 1.2 m. wide. A majority of specimens fall within the lower size range and are easily recognizable. The few larger specimens may be typical or they may approach the form of the subspecies leachii in shape of palpal lateral margin or in development of palpail spurs. The smaller and more compact size and shape of mbhsami is almost always reflected in stouter and more abruptly tapering tarsi. Females. Like males, this sex is smaller, rounder, and more compact than females of leachii. The scutal length is about equal to or only very slightly greater than the width; the posterior margin is more broadly rounded; the punctations are rather large and moderate in numbers; and the cervical grooves are more con cave and more distant from each other. The palpal outline usual. ly has the lateral margin definitely concave; the ventral basal spur is absent but the dorsal basal spur is quite variable; the ventral retrograde spur of segment 3 is like that of the male and of the subspecies leachii. The basis capituli in all avail. able specimens is definitely short and wide with lateral margins distinctly diverging anteriorly and the cornua are usually broad. ly tapered and short. Coxal spurs correspond to those of the male. Tarsi tend to be shorter, stouter, and more abruptly tapered than those of female leachii. = 351. = 155 iS7 Figures 154 and 155, oc’, dorsal and ventral views Figures 156 and 157, o, dorsal and ventral views HAFMAPHYSALIS PARMATA Rio Muni specimens PLATE XLVI = 20L t= HAFMAPHYSALIS PARMATA Neumann, 1905. (Figures 154 to 157) THE WEST AFRICAN ANTELOPE HAEMAPHYSALID 1 Fy: eee ale fo EQUATORIA PROVINCE RECORDS ik Noli Hills Cephalophus caerulus musculoides Mar (SGC) ik Nagichot domestic cattle Jul 2 Gilo domestic cattle Dec BIRD 1 6 Lotti Forest Guttera edouardi sethsmithi Apr These localities, all in the central area of the east bank of Equatoria Province, lie between 4500 feet and 6500 feet elevation. H. ta has not been found in other Provinces of the Sudan. The fort Trtrs specimen is somewhat atypical, see REMARKS below. DISTRIBUTION 2 2m is a quite common Central and West African tick that ranges ins er numbers into the forested highlands of eastern Africa. It is especially numerous in the Cameroons. WEST AFRICA: GOLD COAST (Nuttall and Warburton 1915). FRENCH WEST AFRICA (Villiers 1955). SIERRA LEONE (Simpson 1913. Nuttall and Warburton 1915). NIGERIA (Ziemann 1905. Neumann 1911. Simpson 1912B. Nuttall and Warburton 1915). CENTRAL AFRICA: CAMEROONS (Neumann 1905,1911. Ziemann 1905, 19124. Nuttall and Warburton 1915. Rageau 1951,1953A,B. Hoogstraal 1954C). RIO MUNI (Numerous specimens in HH collection from north. central part of state; K. C. Brown legit; gift of Colonel R. Traub). FRENCH EQUATORIAL AFRICA (Fiasson 1 - Rousselot 1951,1953B. Hoogstraal 19540). BELGIAN CONGO and RUANDA-URUNDI (Nuttall and Warburton 1915,1916. Bequaert 1930A,B,1931. Schoensaers 1951A,B. Van Vaerenbergh 1954. Santos Dias 1954D. See HOSTS below). - 383 = EAST AFRICA: SUDAN (Hoogstraal 19543). KENYA (Neave 1912. Anderson 19244. Neumann 1913. Nuttall and Warburton 1915. Lewis 1931A,C. As H. calcarata:; Lewis 19318. As H. bispinosa: Lewis 1934. Loveridge 1956A. Hoogstraal Tea UGANDA (Nuttall and Warburton 1915. Mettam 1932. Theiler 19450). Z SOUTHERN AFRICA: MOZAMBIQUE: Santos Dias (1954F); not typical specimens 11 description and illustrations are correct. UNION OF SOUTH AFRICA: Theiler (1945C) states that this species actually has not been found in the Union of South Africa, and (correspondence) that Curson!s (1928) and Bedford's (1932B) records from Zululand are misidentifications./ NOTE: Records from Sumatra (Galli-Valerio 1909B) undoubtedly are based on erroneous identification. HOSTS The chief hosts of adults are antelopes; any domestic animal may be attacked. Immature stages parasitize carnivores and ante— lopes and larvae have been recorded from forest birds. Adults Domestic animals; Cattle (Neumann 1905,1911, Ziemann 1905, 1912A, Nuttall and Warburton 1915,1916, Mettam 1932, Schoenaers 1951A,B, Rageau 1953B, Hoogstraal 1954B, Sudan records above). Sheep | are 1905,1911, Ziemann 1905, Mettam 1932, Rageau 1953B). Goats (Neumann 1905,1911, Ziemann 1905,1912A, Mettam 1932, Rageau 1953B). Dogs (Ziemann 1912A, Mettam 1932, Rousselot 1951, Rageau 1953B). Pigs (Rageau 1953B). (?Domestic) Pigs (Neumann 1905,1911, Ziemann 1905). Antelopes: Hartebeest (Nuttall and Warburton 1915. Mettam 1932). Sishiaek (Simpson 1913, Lewis 1931C, Nuttall and Warburton 1915, Bequaert 1931, Mettam 1932, Hoogstraal 1954C). Harnessed antelope (Nuttall and Warburton 1915). Royal antelope, black duiker, bay duiker (Villiers 1955). Bushbuck and Maxwell's duiker (Cameroons, J. Mouchet legit, HH det.). Impala and Harvey's duiker - 384 - (Hoogstraal 1954C). Blue duiker (Mettam 1932, Hoogstraal 1954B, Sudan record above). Forest or red duiker (Mettam 1932). Duiker and "forest antelopes" (Rio Muni specimens noted above). Okapi (Belgian Congo specimens, MCZ, HH identified). Other wild animals: Water chevrotain (Bequaert 1931 and Onderstepoort collection). Buffalo (Nuttall and Warburton 1915, Mettam 1932). Bushpig (apparently from Ziemann's (1905) remarks for "pigs": Nuttall and Warburton 1915). Immature Stages All the following records are for nymphs wnless larvae are also noted. Antelopes: Harnessed antelope (Nuttall and Warburton 1915). "Duiker® (Fiasson 1943B, Rousselot 1951). Harvey's duiker (Hoog. straal 1954C). "Forest antelopes" (Rio Muni specimens noted above). Nymphs and larvae from bushbuck (Theiler 1945C) and from duiker (Theiler, unpublished). Larvae from "duiker™ (Rio Muni specimens noted above). Carnivores: Genet and mongoose (Hoogstraal 1954C). Domestic dog (Rio Muni specimens noted above). Civet, and larvae and nymphs from genet (Cameroons, J. Mouchet legit, HH det.). Other mammals; Black and rufous elephant shrew, Rhinonax petersi, from Tanganyika (Theiler, unpublished). Birds: Larvae and nymphs from forest guineafowl (Hoogstraal 1954B; Sudan record above). BIOLOGY Available data indicate that H. parmata inhabits humid, for ested regions of West Africa but that more easterly populations find optimum conditions for survival chiefly in forest and uplands. Neumann's (1913) and Lewis! (19310) Kenya reports were from areas between 1000 feet and 8000 feet altitude. In Ruanda-Urundi, this tick is found up to about 5600 feet elevation (Schoenaers 1951B). - 385 - Sudan records and several others in my collections are all from altitudes above 3000 feet. In the Sudan and frequently elsewhere, these hills are more humid than the surrounding plains. DISEASE RELATIONS Unstudied. REMARKS The only other tick recorded from Africa that has palpal characters more or less similar to those of H. parmata is H. bispinosa Neumann, 1897, an Asiatic species that 1s said to be found rarely on domestic animals in Kenya. Males of H. bispinosa can be distinguished by their more narrow and elongate scutum, long lateral grooves, and abrupt tapering of tarsus IV. Females of H. bispinosa have a scutal outline that is slightly longer than aoe converging cervical grooves, and a shorter, wider basis capituli. In addition, the distal tapering of tarsus IV is more abrupt. Nuttall and Warburton (1915) recorded a few specimens of H. bispinosa from Kenya, but Lewis! specimens under this name ee cnelis H. parmata (Hoogstraal 1954C). With respect to the tapering of tarsus IV, the Noli Hills female specimen from Equatoria Province is like H. bispinosa. In all other characters, however, it appears to equal fr rmata and it is therefore assigned to the latter species, Sara with some hesitation. Students of Haemaphysalis ticks believe that such tarsal characters are constant within a species, but because of the dearth of comparative material it is impossible to arrive at a satisfactory conclusion concerning this variation. The material described by Santos Dias (1954F) from Mozambique appears to differ somewhat from that known from the rest of Africa. In a formidable discussion, Schulze (1938A, figure 31C) has illustrated the palpi of H. parmata as data for his theories con cerning generic and specific indicators resulting from pressure of the body within the developing nymph of ticks. =50)— IDENTIFICATION Males are easily recognized by the pointed dorsal projection from the basal margin of palpal segment 3; peculiarly shaped palpi; short and broad scutum; very short cervical grooves; lateral grooves that reach only midlength of scutum; numerous, medium size, deep scutal punctations; coxae with short but distinct basal spurs, and tarsus IV gradually tapering. Palpal charac- ters alone are enough to quickly separate H. parmata from other African species. Males are very small; they measure from 1.3 mm. to 1.8 mm. long and from 0.75 m. to 1.1 mm. wide. Females have the same distinctive palpal features as do males. The subcircular scutum measures from 0.64 mm. to 0.70 mn. long, and from 0.75 mm. to 0.90 mm. wide; it has broad, shallow, parallel cervical grooves extending to its midlength and medium size punctations that are more shallow than those of the male. Coxa I has a rather wide, short posterior spur but other coxal spurs are replaced by broad posterior ridges. Female palpal characters are as distinctive as those of the male among the African fauna. Theiler (1945C) has redescribed and illustrated both sexes and the immature stages of H. parmata. Dr. Theiler identified the larvae and nymphs from the forest guineafowl from the Sudan. =), Soe Ey Voce BOM ahteA INTRODUCT ION The genus Hyalomma is a complex of a few species exhibiting an almost endless variety of facies. Its original center of dis persal was probably Iran or southern Russia. Genetic instability may in part account for the wide morphological differences found in many specimens. Environmental vicissitudes are undoubtedly important additional factors in modifying size, color, and over. all appearance in this genus. These are tough, hardy ticks that survive under conditions in which all other species are uncommon or entirely absent; they may even thrive in such environs. They inhabit country where humidity is frequently low, seasonal climatic conditions are extreme, favorable niches for development away from the host are rare, smaller animals for immature-stage feeding are sparse, and larger-size hosts are frequently poorly nourished and wander widely among inhospitable situations. Owing to their medical and economic importance and the pressing need to clarify the relationships of all presently recognized spe— cies in the genus Hyalomma, the plan of this section has been mod- ified to include a key to all species and illustrations of non. Sudanese species. Further research will somewhat modify present concepts but this compilation of information will provide a firmer foundation for subsequent revision than is now available without considerable background study. The presently recognized species of continental Africa are, however, fairly well stabilized and the disconcerting prospect of further nomenclatorial changes and addition of new species applies chiefly to populations from the Near East to the Far East. NOMENCLATORIAL BACKGROUND HISTORY It is hardly surprising that criteria for identification of Hyalomma species have long been in a chaotic state. The thir- teen species described by Koch (1844), when he erected the genus, = 988 = in addition to three previously described species, remained mostly unrecognized by subsequent workers. The genus was reduced to four species, including a single new one, and four subspecies by New mann (1911). H. aegyptium was used as a "catchall" name by most persons until The repo During the early twentieth century, British workers in Africa, depending on Nuttall and Warburton at Cambridge for identification of their collections, developed a group of names that are herein referred to those in contemporary usage after having studied the Nuttall collection in British Museum (Natural History). Between 1919 and 1950, Schulze and a few of his students and followers seized upon the apparently unlimited opportunities for providing dozens of species names for variants in this genus. Scarcely a single one of the some eighty species and subspecies proposed by Schulze and colleagues has withstood the test of comparison with reared progeny from a single female tick. After having studied parts of Schulze's collection, now housed in Rocky Mountain Laboratory, one can understand, from the small series and poor labelling, how misconceptions regarding species identity developed among persons eager to tag each variation with a spe— cies name. Schulze even went so far as to name the progeny of a single female as different species (H. delpyi Schulze and Gossel, 1936) (Delpy 1946A). RECENT REVISIONAL AND SUPPORTING STUDIES During the last twenty years a certain amount of cosmos has begun to evolve from this nomenclatorial chaos, although it is obvious that additional modifications in species concepts and names are yet to come. The careful, tedious, and time-consuming pioneer work of Delpy, who secured specimens from many areas where hyalommas occur and reared the progeny from single fe males, enabled him to determine the range of variation within a single species and to show that characters proposed for many so-called species were due merely to mltiformity of appearance within a few species. In a few instances, however, Delpy im cluded species that we now know to be distinct genetic entities worthy of species rank. - 389. Shortly afterwards, Adler and Feldman-Muhsam commenced rear. ing Palestinian species in the same manner as Delpy. They corro- borated Delpy's species definitions but not his species names. In their 1948 paper these authors provided a potash clearing method for females by which they established constant species characters for the unmated female genital aperture. Delpy expanded this finding to mated females, thus making it of greater value for identification of field-collected material. Neverthe. less, some questionable specimens inevitably crop up in routine collections. Unfortunately, as stated above, Delpy and Adler and Feldman. Muhsam arrived at different conclusions regarding which name from the scores available should be applied to individual species. Re cently Feldman-Muhsam (1954), after study of Koch's (1844) type specimens for several species in the genus, has corroborated some of Delpy's earlier decisions and proposed a few changes. Although Koch!s material is badly damaged and its labels have been inexcus— ably tampered with, these studies probably represent the final word on these species; therefore this terminology is accepted with certain reservations as noted in the appropriate places. Delpy's chief morphological and taxonomic contributions to Hyalomma have been his notes on the genus (1936,1946A), descrip. tion of H. schulzei (1937A), description of the immature stages of H. dromedari1 (1937B), generic revision by experimental methods (1927D and 19Z9R, especially the latter), and a synoptic list and discussion (1949B), Feataes studies on bovine theilerosis and tick transmission (19370 ,1946B,1947A,1949% and 1950). Adler and Feldman-Muhsam presented their chief overall findings in their 1948 paper; subsequent reports by the latter author are listed in the bibliography. Whenever possible, Delpy's (1949B) synonymy has been followed in the present work. Some changes have been necessary, however, on the basis of the kind of proof that Delpy himself advocated: rearing of progeny from single, known females. A few other changes have been necessary due to Feldman.Muhsam's study of Koch's types. It is impossible to decide whether Delpy or Pomerantzev (1950 should be followed for the synonymy of certain Russian species. Pomerantzev's ideas, whenever they differ, have been included as notes under the names indicated by Delpy. - 390 - It has been attempted herein to indicate present and previous nomenclatorial concepts of these species as clearly as possible, especially for experimental workers and reviewers. Non-taxonomists, who consider themselves "practical workers™, will undoubtedly be annoyed by the remaining confusion. The end is now in sight, and within a very few years will undoubtedly be reached. A little more patience will be rewarded by better understanding of what has been an especially difficult complex of variable species in previously poorly explored parts of the world. HYALOMMA DISTRIBUTION IN AFRICA SRS pe Two species, H. truncatum and H. rufipes, are common in drier areas throughout the Ethiopian Faunal Region (Figure 1). Two others, H. albiparmatum and H. impressum, are restricted to equa torial regions of Africa; these four species appear to have e. volved in Africa from Near Eastern stock. Only H. rufipes ex tends beyond the confines of the Ethiopian Region. Two other species (H. detritum and H. marginatum) range into North Africa from the Near Bast and have tenuous, scattered footholds in the transitional zones just south of the sreat deserts alons the northern periphery of the Ethiopian Region. Another Near Eastern species, H. impeltatum, appears to be extending its range a little more aggressively into East and West Africa. The last species known from continental Africa, H. turanicum, has established it- self in the South African Karroo after having been introduced on sheep from the Near or Middle Hast. In East Africa, the arid lowlands along the Red Sea and the Indian Ocean carry a number of Near Eastern and North African species southwards into the Somalilands and parts of Kenya to_ wards and even slightly south of the equator. For instance, H. dromedarii is known from the coastal lowlands of Kenya (Walker, unpublished) and H. impeltatum occurs in scattered foci in Kenya and Tanganyika. There is little question that other species do exist in nature but their identity can be established only by breeding experiments. The presence of a possibly undescribed species similar to H. drome— = 391, darii in the French Somaliland fauna has also been noted (Hoogstraal T953D). Note the following incorrect Ethiopian Region records: marginatum (= H. savi savignyi), reported by Rousselot (1948) from fee West africa, was =o subsequently confirmed (Rousselot 1953B). H. detritum reported from French Cameroons (Rageau 1951) was subsequently (1553) assigned to H. truncatum GH. transiens) by the same author. References to "H. savignyi" from Portugese Guinea (Tendeiro 1949,19520) actually apply * H. truncatum. In northern and central Sudan, eight species are established though seldom in a continuous range. Of these, only four are com. mon. In most Near Eastern and North African areas about the same proportion of common and rare species occur. Yet with even so few species among which to choose the student frequently encounters difficulty in positive identification of all material in most large collections. Some specimens are so variable and intermediate that they defy assignment to a definite species. Unfortunately, previous workers have not provided pertinent details over extremes of variation among species that they have reared. Attempts to properly identify Sudan material for this report have necessitated so much study of material from other parts of the world that pub. lication has been long delayed. With regard to the paucity of specimens of some species col. lected in northern Sudan, it should be emphasized, from our ex. perience with vertebrate and invertebrate animals in arid and semiarid areas of Africa, Arabia, and the Near East, that not infrequently small relict populations of animals are found in. explicably surviving in barely marginal habitats. This appears to be true of H. detritum, H. marginatum, H. truncatum, and H. impressum in northern Sudan. The northcentral areas of the Sudan, by reason of their proximity and similarity to the Mediterranean subregion and their tenuous routes of entry from Arabia, West Africa, and via Sees the Nile, are inhabited by more species of Hyalomma ticks than apparently any other area of the Ethiopian Faunal Region. The fact that some of these species appear to be represented in the Sudan only by small populations, either as a result of chance introduction or as survival or relicts, has been noted above. The Asiatic species that do not reach the Sudan are H. hussaini of India (page 520), H. schulzei, an Iranian camel par- asite that reaches the Sinai Peninsula between Asia and Africa (page 525), H. aegyptium, the tortoise parasite that extends from southern Russia westward through much of the Mediterranean basin (page 514), and H. turanicum of southern Russia and Iran that has been introduced into the South African Karroo (page 528). As stated above, the original center of distribution of hyalommas appears to have been in southern Russia or Iran. Delpy and Adler and Feldman-Muhsam have provided few de— tails about the geographical source and range of species that they treat, and there is still considerable question in the minds of specialists and reviewers as to the distribution of Hyalomma species. This section has therefore been given special attention in the following text. Synonyms, listed by country of origin of specimen material whenever it can be determined, are based on Delpy!s (1949B) lists, which give every evidence of being carefully and judiciously assembled. These references do not include the entire literature, except I trust for Africa, but are furnished for what they are worth in elucidating the distribution of Hyalomma species and indicating the major stu. dies of each species in different parts of the world. HOSTS AND BIOLOGY Biological data for Hyalomma ticks derive chiefly from veterinarians! observations on those infesting domestic animals and on laboratory experiments. From field work and from a few other sources we have gained a somewhat different impression of Hyalomma biology, especially relating to host preferences of eS immature stages. In this respect, special attention is called to the HOSTS and BIOLOGY sections in the following text, especially for H. excavatum. The natural life cycle of Hyalomma - 393 - ticks may be altered by the size, numbers, and density of available hosts. Further research on this subject is strongly indicated. In the introduction to this section it has been stressed that many Hyalomma populations survive in inclement environments and are greatly affected by extremes in temperature, humidity, and condition of host nourishment, as well as by the wide wandering of their hosts over thinly populated, inhospitable xeric areas. Much more collecting, observing, and careful identification is necessary before the ecology of most species in this genus can be adequately determined. The value of innumerable published reports on the biology of the genus is vitiated by the inaccuracies in identification. Extraordinary survival factors play a large part in permitting these ticks to exist and even thrive where few or none others live. The life cycle of hyalommas may be greatly lengthened in un. favorable climatic conditions, or shortened under optimum condi- tions. Nuttall (1915) kept adult specimens alive without food for approximately two years and observed copulation and feeding after this period of starvation. Nuttall (1920) also found the capacity for regeneration of lost appendages and injured mouth. parts to be greater in Hyalomma ticks than in most others. A certain amount of ebridiwation is possible though curiously mis. formed individuals may result; these and other greatly misformed specimens that have still survived are reviewed by Pervomaisky (1950B,1954). Special attention is called to the discussion below of the two-host, summer-feeding H. detritum, and its biological race H. scupense which is a single-host, winter-feeding form with slight morphological differences in most of its range (page DISEASE RELATIONS Adult Hyalomma ticks, except H. S25 are today chief. ly parasites of domestic animals wherever they are found, and, as such, are of considerable economic importance. Hyalommas - 394 - appear to be unusually efficient vectors of a variety of disease. causing organisms. In their immature stages, they often feed on birds, rodents, and hares that are important reservoirs of patho. gens, especially viruses and rickettsiae. Few ticks have been incriminated as reservoirs and vectors of pathogenic viruses, but several species of hyalommas are known to be hosts and vectors of the viruses causing several distinctive acute infectious hemorrhagic fevers of human beings in the Soviet Union. Unpublished studies by Daubney (conversation) indicate that one of these same species may transmit in nature the virus causing a Near Eastern encephalomyelitis of equines. These same tick species occur in North Africa and northern Sudan. Other species cause paralysis of man and animals, apparently as a result of toxins injected into the host while the tick is feeding. The association of Hyalomma ticks with a number of other human and veterinary diseases is noted in the following text. Many Hyalomma species, in our experience, attach readily to man and teat on him. The "cursorial ticks" of North African and Arabian deserts, as first described by Mann (1915), are several species of hyalommas that come rushing from beneath every shrub when persons or animals stop nearby. These are almost invariably unfed adults, of uniform size, shape, color and general appearance, that have molted from the nymphal stage in rodent burrows beneath shrubs. Although few of these highly agitated young adults actual. ly attach to man, some do. Confusion in nomenclature has limited the value of many ear. lier studies on biology and disease-transmission in this group, for it is often impossible to be certain which species the writer used in his work. Considerable study on this genus has been and is being done in Russia, and it is frequently difficult for re- viewers to determine exactly the species being reported and to satisfactorily evaluate the reports. In addition, it should be indicated that the range of Hyalomma ticks covers, in large part, a vastly undeveloped part of the world in which little serious scientific research has been accomplished. Before many years have elapsed, enough evidence probably will have been presented to indicate that Hyalomma ticks are economically among the most important of ani ectoparasites to be found any. where in the world. - 395 - IDENTIFICATION Use of the following key should never be attempted without reference to the section on identification for each species men. tioned in the text. In the identification section, an attempt has been made to present lucidly all important characters of typical specimens and to indicate the range of variability seen in each species. I am most grateful to Mr. Makram N. Kaiser, Chief Technician in the Department of Medical Zoology at U.S. Naval Medical Research Unit, who has served as a "sounding board" and has given invaluable assistance in grouping the very large numbers of specimens that have been studied and often re. studied for this section. Special acknowledgement should also be made of the services of Dr. L. P. Delpy, who initially ider tified many of our early collections of Hyalomma and of Mr. Glen Kohls who has spent several days eee over specimens in the Schulze collection, now deposited in the Rocky Mountain Laboratory at Hamilton, Montana. Persons attempting to identify field-collected material of Hyalomma should recognize that a certain proportion of spec- imens in many series will defy final determination of species. These had best be called "Hyalomma species" and sent to a capable specialist in the group or put aside for further study as addi_ tional information becomes available. - 39 - KEY TO SUDAN SPECIES OF HYALOMMA (TYPICAL SPECIMENS ONLY) MALES 1. Center of subanal shields character. istically exterior of the axis of the adanal STON S Hiarecove eo evorarore even 0 cia elocotavciererave.e\esayenese ele osele @/e1ee Center of subanal shields character. istically in line with the axis of the adanal SBHAVONG Sie race o'oree evelave ee iaraceveveleverevelalersiaiele a eielere leia"sleie.) 2. Medium size ticks (scutum about 3.8 x 3.0 m.). Lateral grooves extending anteriorly at least to midlength of the scutum. Scutum largely covered by medium size punctations. (Common on cattle in central Provinces)....e.se..sseeeHe wee Figures I Large ticks (scutum about 4.5 x 3.2 mm.). Lateral grooves limited to posterior third of scutum. Scutum with few large, scattered, irregular punctations (sometimes also with small ones). (Common where camels occur in Sudan). .ccccecccccscncnccectocccescesls ee Figures 1 *In H. excavatum, and sometimes in other species that have en. gorged on Large animals such as camels, and have considerably stretched their integument, the subanal shields may be laterally displaced. Such excavatum specimens would appear to be H. dromedarii, except for smaller size, fewer and smaller scutal punctations, and differences in the caudal area of the scutum. See also identification of H. impeltatum for superficial varia. tion among unfed males, which may cause them to be suggestive of H. marginatum. - 397 - Sle 4. 5e Lateral grooves not extending beyond the posterior third of the scutum. Scutum with few punctations except in the caudal area which is depressed between two lateral ridges and some. times very shagreened. Small ticks, often frail, maximum overall length usually less than 5 mm. (Fairly common on cattle and especially on horses in central Provinces; also occurs in Northern Province} woccncccecccaccccccccsctls EXOAVALUM Figures 165 an Lateral grooves extending beyond the midlength of the scutum (may be obs— cured in very heavily punctate spe cies; examine by oblique orientation) ....c.ccscsecescscccencess Scutum smooth, bright*, with very few, large, shallow, scattered, punctations; posteromedian and paramedian grooves well marked. Legs usually not ringed. (In northcentral Provinces; rare)....sssseeeeeeeeeeeHe DETRITUM Figures 158 and 159 Scutum densely punctate, or with dense, contiguous punctations posteriorly...ccccccccccceccsccccceceeed Scutum densely, entirely, and almost always uniformly covered by puncta. tions often obscuring the lateral GEGOWES oicyorctovclororarevatosoraisio! stoke ahaloxoiaiaierarsisisvolaiareiolaielere\eishsialalniainioiniaiate sare Scutum irregularly punctate, or punctate only POSCELLOPLY . ) Genital apron a large, wide shield, bulging in profile. Circumspiracular Grea AMuUsuAlL Lys P1OSOs ssisa'elc cies cicieie clateieicicicie cise vicccctle RUPIEIss Figures 164 and 185 Genital apron a wide triangle with a narrow transverse anterior ridge (bulging in profile) and a posterior button (sloping or depressed in profile). Circumspiracular area Not vexcepttonal] yu pUlOse ss «is\e ocle s cisisisieie cimisisiceeoc He IMPRESOUM Figures 176 and 177 - 401 - 6. tio Genital apron a transverse oval, posteriorly deeply concave in profile. Scutal punctation usually small and regular, TALFELY COATSC. cccccccccccvcccccccccccccccccccccels LRUNCATUM Figures 1 Genital apron strongly bulging (convex) in POL 11 Cc cicieic.c e\e'c cleieieleis.e cioieinloisieieleistelu/ersiele alvin oiaieles Genital apron large, a widely trans verse oval or triangle. Scutal punc- tations shallow, large ones restricted to anterior half, small ones widely SCACLEPEAs cicocsccccccccocceccesccesssscoscccoell. MAG INALUM Figures 180 and 181 Genital apron an elongate triangle flanked by two lobes, giving it a trilobed appearance. Scutal puncta tions usually numerous, regularly spaced, and medium-size posteriorly and centrally with two rows of larger and deeper punctations among them; with larger punctations anteriorly and in scapular fields. ...ccccccccccccccccccccets. IMPELTATUM Figures 172 and 173 = 402 Figures 158 and 159, @, dorsal and ventral views Figures 160 and 161, 9, dorsal and ventral views A to F, 9 genital area outline and profile. A to C, partly engorged. D, fully engorged. E and F, unengorged. HYALOMMA DETRITUM Sudan Speci mens PLATE XLVII - 403 - HYALOMMA DETRITUM Schulze, 1919 (Including H. SCUPENSE Schulze, 1919) (Figures 158 to 161) THE SHINY HYALOMMA NOTE: The some twenty synonyms of H. detritum, as presented by Delpy (19498), are listed in the section on DISIRIBUTION below according to the political areas from which the specimens of each originate. It is impossible to believe that H. detritum had not been described as a species by some author somewhere prior to 1919, yet no contemporary specialist has come forth with a previous name for this species. Steps should be taken to stabilize the name H. detritum in order to prevent the further confusion that is bound to arise when an earlier name for this tick is inevitably dis. covered. Special attention is called to the "biological race", H. scupense, discussed below. DISTRIBUTION IN THE SUDAN H. detritum occurs rarely in northern parts of the central Provinces of the Sudan. Additional collecting is necessary to determine its exact distribution here. It would be of some interest to know whether H. detritum has invaded the Sudan from the Red Sea coast or via the Nile Valley. Kassala: Port Sudan (cattle; SVS). It is not known whether hosts of these specimens were local or transient animals. Kordofan: Four males have been collected from Kordofan cattle at the Khartoum Quarantine station (January and February) (HH). Khartoum: See Kordofan above./ 4 LOA = DISTRIBUTION H. detritum is an Asiatic tick that ranges from Manchuria through China and India, much of southern Russia, southeastern Europe, and the Middle East; into Asia Minor, the Near East, the Mediterranean littoral of Africa westwards to Algeria; and into northcentral Sudan, where it occurs only in small, localized populations. It also occurs in Spain, probably having been intro duced from northwestern Africa. In Egypt and apparently in Libya, H. detritum is consider. ably less common than it is to the west in Algeria and in Near Eastern countries. This distributional pattern is common for animal groups that have invaded North Africa from the east. NOTE: Specimens referred to this species by Rageau (1951) from the Cameroons were later determined by him (1953) as H. truncatum (= H. transiens). NORTH AFRICA: *ALGERIA (All as H. mauritanicum or as H. mauritanicum annulatum: Senevet 19228,C,19224,19-5,1928A,1929B, 1937. Senevet and Rossi 1924. Sergent, Donatien, Parrot, Lesto. quard, and Plantureux 1926,1927A,B,C,D,E. Sergent, Donatien, Parrot, and Lestoquard 1928A,B,C ,1931A,B,C ,D,E,F ,1932A,B,1933A,B, 1935A,B,1936A,B,C,1945. A. Sergent 1930. Sergent and Poncet 1937,1940,1941. Sergent, Donatien, and Parrot 1945. E. Sergent 1948. Blanc and Brunneau 1949. d'Arces 1952. As H. detritum mauritanicum: Schulze 1930. Kratz 1940). in MOROCCO (As H. mauritanicum: Desportes 1938). TUNISIA (As H. detritum: Senevet 1937. Colas-Belcour and Rageau 1951). LIBYA (As H. mauritanicum: Franchini 1927,1929A,E. Hoogstraal, ms.). EGYPT (Present but rare: Hoogstraal, ae EAST AFRICA: SUDAN (As H. detritum: Hoogstraal 1954B). Note: H. mauritanicum has been reported from Somalia, with. out precise locality data, by Niro (1935), but this record has not been subsequently repeated by Italian workers. *Algerian specimens of H. mauritanicum, kindly presented by Dr. E. Sergent and Dr. Senevet, conform to H. detritun. = (Os = NEAR EAST: PALESTINE (All as H. detritum: Adler end Feldman Muhsam 19Z6,1948. Feldman Muhsam 1948. Adler 1952). SYRIA (As H. detritum damascenium: Schulze and Schlottke 1930. Schulze 1930. Kratz 1920. As H. ?mauritanicum: Pigoury 1937). TURKEY (As H. steineri steineri: “Schulze 1930D. Kratz 1940. As H. detritum: Mimioglu 195. Kurtpinar 1954. Hoogstraal, ms. As Hf mauritanicum: Yasarol 1954). IRAN (As H. detritum: Delpy 930B, L040). ap: ig ae danicum: Pavlov 1947). ¥*YUGO s either H. scupense or H. savignyi by different local workers according to oe 1937. As HW. scupense and H. detritum dardanicum: Oswald 1937,193&A, 3 CT1SOL TTS 31940. As HW. detritum: Angelovsky 1954). GREBCE (As H. scupense; Knuth, Behn, and Schulze 1918. Schulze 1919,1930. ance and Schlottke 1930. Kratz 1940. Pandazis 1947. As H. detritum scupense: Delpy 1946. As H. detritum dardanicum: Schulze and acrTottke 1930. Schulze 15350. Kratz 10L0. Pandazis 1947). EUROPE: *BULGARIA (As H. Heyeout and as H. detritum dar. SPAIN (As H. steineri codinai: Schulze 1936p. Kratz 1940. As H. mauritanicum: Gil Collado 1948A. Miranda-Rntrenas 1954). RUSSIA: As H. detritum: Olenev 1929B,1931A. Pavlovsky 1940. Kurchatov 1941. Pavlovsky, Galuzo, and Lototsky 1941. Galuzo 1941,1943,1944. Lototsky and Pokrovsky 1946. Tselish cheva 1953. Viazkova and Bernadskaia 1954. Petrisheva 1955. Zhmaeva, Pchelkina, Mishchenko, and Karulin 1955. As H. detritum detritum: Olenev 1929A,1931C. Schulze and Schlottke 1930. Schulze 1930. As H. detritum rubrum: Schulze 1930. Olenev 1931A,C ,1934. Pomerantzev 1954. Galuzo 1935. Galuzo and Bespalov 1935. Kratz 1940. *The hosts of immature stages listed by authors in these countries indicate that they are quite possibly dealing with a different species of tick. = 4067— As H. detritum pavlovskyi: Olenev 19294. As H. transcaucasicum: Olenev 1934 (the synonymy of this name appears to have been overlooked by subsequent workers). As both H. detritum and H. scupense: Pomerantzev 1937. Pomerantzev, Matikashvily, and Lototsky 1940. Markov, Gilden. blat, Kurchatov, and Petunin 1948. Note: Pomerantzev 1950, in his work on Soviet ticks, considers these two as distinct species. See BIOLOGY and IDENTIFICATION below. As H. scupense: Olenev 1934. Nikolsky 1948. Petunin 1948. Pomerantzev . Alfeev 1951. Shatas 1952. Melnikova 1953. Rementsova 1953. Shatas and Bustrova 1954. Note: Pomerantzev 1950 considers H. volgense and H. uralense to be synonyms of i. scupense, while Delpy places them under either H. detritum or A. excavatum (see two paragraphs below). Note the references To WH. scupense" in the section on European distribution above. See also paragraph above and below. H. verae Olenev, 1931B, is also placed in synonymy under H. scupense by Pomerantzev 1950; Delpy did not consider H. verae In his re T is of synonyms. As H. volgense and/or H. uralense: Schulze and Schlottke 1930. Sei SSnTS5O. Olenev 1929K,19314,C ,1934. Zasukhin 1932, 1935. Borzenkov and Donskov 1934. Zolotarev 1934. Galuzo 1935. Kochetkov 1935. Artjukh 1936. Kurchatov 1940B. Markov, Abramov, and Dzasokhov 1940. Enigk 1947. See paragraph on H. scupense above and paragraph below. Delpy (1949B) was not certain whether H. uralense Schulze and Schlottke, 1930, and H. volgense Schulze and Schlottke, 1930, are synonyms of H. detritum or of H. excavatum, and stated that Russian workers may have included both species under these two names. In Schulze's collection, now in Rocky Mountain Laboratory, there are 20 and 209 from Ukrainia, identified by Schulze as H. volgense. These are typical H. detritum. The same institution Possesses and 399 from Crimea, determined by Schulze first as H. ?marginatum, and later crossed out and identified by him as H. - 407 - uralense. The males are, all but one, H. detritum*; the exception appears to be H. marginatum; the females are in poor condition. It is reasonable to assume that what is now considered to be H. detri_ tum (or, in part, "H. scupense") was treated by Schulze in per ‘as H. uralense and in part as H. volgense, though, he overlooked other Species in the same collection and referred to them by the same name. As H. tunesiacum pavlovskyi: Schulze and Schlottke 1930. Described and SMe as H. detritum pavlovskyi by Kratz 1940. According to Galuzo 1935, the H. asiaticum of Olenev is H. detritum (= H. detritum rubrum). H. aslaticum is usually con Sldered as a synonym of i dromedarii. MIDDLE EAST: INDIA (As H. aegyptium ferozedini and as H. a. isaaci: Sharif 1928. As H. sharifi: Schulze and Schlottke 1930.” Schulze and Gossel 1936. “Kratz 1940. As H. isaaci: Kratz 1940). PORTUGESE INDIA (As H. detritum: Santos Dias 19543). FAR EAST: CHINA including MONGOLIA (As H. detritum albi- ictum: Schulze 1919,1930. Schulze and Schlottke 1930. “Yama. shita 1939. Kratz 1940. As H. detritum perstrigatum: Schulze 1930. Schulze and Schlottke 1930. fee and tone 1933. Olenev 1934. Kratz 1940). HOSTS Domestic cattle and horses are the most common hosts of H. detritum, all stages of which feed on the same kind of animal. Sheep and goats are sometimes attacked. For the Soviet Union, Pomerantzev (1950) lists cattle, horses, donkeys, pigs, camels, sheep, and hares; and, for nymphs, especially cattle and horses. Man is apparently commonly attacked under local conditions. Oswald's (1939) and Pavlov's (1947) remarks for parasitism by immature stages in Yugoslavia and Bulgaria of various birds and *At the time of checking this material, I did not realize the significance of "H. scupense". It cannot, therefore, be said that these specimens = not resemble the latter form. = 408 = lizards, and of hares and dogs are either incorrect or refer to a different species of tick, most probably H. marginatum.7 A. Sergent (1930) noted a nymph of H. detritum parasitizing another nymph of the same species. Hosts of specimens in British Museun (Natural History) are domestic cattle and camels (Palestine), domestic buffalo and pony (India), hare (India), and deer (Romania). "H, scupense” attacks the same hosts as H. detritum. It has also been round on the Persian or goitred gazelle, Gazella subeutterosa, and on the red deer, Cervus elaphus bactrianus (Pomerantzev 1950). In the Crimean Nation orest (MeLnikova 1953), "H. scupense” is common on red deer and occurs in smaller numbers on roe deer. It is present but not common on hares but absent on squirrels and jays. Wild foxes may also be attacked. Domestic cattle are heavily infested, collections from single animals in various localities in and near the forest averaging from 78 to 756 ticks, with individual maxima ranging from 350 to 5000 ticks per animal. Domestic pigs in the same forest averaged 21 ticks per host. Zolotarev (1934) listed this tick (as H. volgense) from camels. BIOLOGY Introduction Because of the interesting biological and taxonomic prin. ciples involved, separate reviews of life cycle and ecology are devoted to H. detritum and "H. scupense". H. detritum is a two- host species whose adults feed in oe Summer and whose nymphs undergo an extensive winter diapause; this feature is common throughout the range of H. detritum though the overwintering habits in Algeria and (usually) in Russia differ markedly; it should be determined what factors account for this variation in habits. H. scupense is confined to parts of the Soviet Union and possibly to Greece and Yugoslavia; it is said to be distinguishable from H. detritum by slight morphological differences through parts of its geographic range (see IDENTIFICATION below), and is a one-host — (Hole) — tick whose adults feed in the winter and early spring. Pomerantzev (1950) considers "H. scupense™ to be a "biological race" of H. detritum. Life Cycle of H. gotritum The life cycle of H. detritum (= H. mauritanicum) in Algeria has been studied by Sergent, Donatien, Parrot, and Lestoquard (1931B and subsequent vorks). Larvae hatch in the late autumn and feed and molt to nymphs on the same host, remaining attached for approximately sixteen days. Nymphs hibernate for approximate. ly eight months either in groups in cracks and crevices of farm yard walls about six feet above the ground, or under boulders. They are never found in fields without trees and boulders. Hi bernation sites are generally those with a warm, sunny exposure, and those nymphs in the warmest places molt earliest in the year (June). Some days after molting, young adults start out in search of a host; they leave their hibernation place at night and travel towards stables, sometimes covering as much as thirty yards a week to reach cattle or horses. Adults commence attaching to the host in mid June. They mate and feed there, females dropping off after ten to twelve days. Feeding females are most common in July and August; afterwards fewer are found on animals. Those that feed later do not oviposit the same season. Eggs are laid on the ground near animals and hatch in about six weeks. Females die after laying eggs. There appears to be quite a little variation from this typical life cycle. Males remain on the host for a much longer time than fe. males and may move from one host to the other. Nymphs some. times move from one host to another. Note that according to the Algerian reports mentioned above, nymphs hibernate in cracks or crevices of farmyard walls but that almost all Soviet workers, mentioned below, find nymphs alone or with larvae on cattle during the winter. This raises the question whether the Russian and Algerian ticks are actually the same spe- cies, and, if so, whether climatic or other factors modify their choice of niches for hibernating in these far-flung areas. a eOr— Galuzo (1943) made a special point of the fact that H. detritum is not found in cracks in the walls in southern Kazakstan. As stated four paragraphs below, under certain local conditions in Russia, nymphs may also overwinter off the host. No explanation for these differences in overwintering habits has been found in the Soviet literature. In southern Kazakstan, Galuzo (1941,1944) reported H. detritum as a two-host tick with unfed nymphs hibernating on cattle during the winter. They engorge and drop from the host in the spring (end of February through April) and molt to adults anywhere from May to August, but mostly in June. Adults disappear in September. Eggs are laid in shaded places in moist meadows, waterside vegeta tion, or under grass. Larvae aestivate in cracks in the soil or on the surface of the ground until October. Then they ascend prass, attach to grazing cattle, feed, and molt to nymphs on the host. Few nymphs feed and drop from the host before winter. In Tadzikistan, where H. detritum is most common in irrigated valleys, it has one generation a year (as apparently everywhere else where it Ronee and the seasonal distribution of feeding of the immature and adult stages is like that in Khazakstan (Pavlovsky, Galuzo, and Lototsky 1941). According to Pomerantzev (1950), when H. detritum is reared in the laboratory nymphs drop from the host any time between October and April. Yet they all undergo the typical winter diapause and molt to adults from May to July, mostly in June. Thus, nymphs that begin life as larvae in October require eight and a half months to become adults, but those that commence feed ing as larvae (under experimental conditions) in April require only two months to reach the adult stage. Furthermore, females may fast for six or seven months and feed, between May and August, for from seven to 27 days, average eleven days. After dropping from the host, females commence oviposition in six to 31 days. Eggs number from 5000 to 7000. Larvae appear 25 to sixty days after the eggs have been laid, and may fast for seven or eight months. They (normally) attach to the host about a week after hatching, feed for eleven days, and molt on the host about eleven days later. There is one generation a year. Pomerantzev (1950) notes that in Middle Asia, (overwintering) nymphs may be found under cattle dung (stored for fuel), and also in the walls of stables (Pavlovsky). Metamorphosis from nymph to adult may occur in buildings and stables thus increasing the importance of the species in the spread of disease. Life Cycle of "He scupense" Knuth, Behn, and Schulze (1918, p. 254) first noted, in their studies of tick-borne diseases in Greece, that "H. scupense" is a single-host tick. According to Pomerantzev (1950), "H. scupense™ (which occurs in the Kursk and Saratov areas, lower Volga, feet: Crimea, Caucasus, Kazakh, Tadzhik, and Yugoslavia, Central Asia, and the rayon of Kogen) is a single-host tick. Adults appear in the winter from January to April and rarely until June. Larvae feed in November, nymphs from November to March. In the laboratory (Markov, Abramov, and Dzasokhov 1940, as H. volgense), females begin to oviposit from 45 to 85 days after Teaving the host and continue to lay eggs for thirty days. Larvae hatch from 45 to ninety days after commencing oviposition and en. gorge in six or seven days. They molt to nymphs on the host, com. mence feeding after several days, and feed for four to six days. Nymphs molt to adults, on the same host, ten days afterwards. In nature, unfed larvae begin to attack cattle in October and molt to the nymphal stage during the same month. The maximum number of nymphs are found in December, when young adults begin to appear. Adults move to different parts of the host and pass the winter on the host. Adults commence feeding early in March. Early in April large numbers of adults leave the hosts. Oviposi- tion and development of the eggs occurs during the summer. [Note the long summer period of oviposition and embryological development, whereas the larval and nymphal development is rapid./ Unfed adults and fed nymphs may undergo a winter diapause. Some engorged nymphs that do not molt remain attached to the host until February. From December to March, adults wander over the host's body but do not start feeding till early spring. The cycle re quires one year. + NA < In northern Caucasus, larval "H. scupense" infest cattle in October and overwinter on the animals. roma appear early in spring (Markov, Gildenblat, Kurchatov, and Petunin 1948). These workers confirm the single-host life cycle of this tick as does Alfeev (1951) and Melnikova (1953), in his study of the ticks of the Crimean National Forest. There, adults are found on cattle and wild ungulates from September through April but rare individuals may be collected on deer during the summer (May to June). All stages occur on deer during the winter (November- December) butin January and February only nymphs and adults are found, and by the second half of March all nymphs have molted to adults. In April and the first half of May, only males and greatly engorged females are found on wild ungulates. A % Ecology of H. detritum In eastern Transcaucasia, H. detritum inhabits various types of desert and steppe areas, described by Pomerantzev, Matikashvily, and Lototsky (1940) (see six paragraphs below). In the Arax val. ley of Armenia, Pomerantzev (1934) found this tick only in saline, marshy types of grassland pasture areas, not in Artemesia semi_ desert areas. It appears that H. detritum infests a selected variety of semidesert areas, and also steppe and other grassy areas, but not forest zones. It still requires to be explained why H. detritum, which seems to be ecologically more limited than WH. scupense", should have a so much more extensive geographical range n "H. scupense". Larvae, since they attach to cattle in the cool months of October and November in Tadzhistan and eastern Uzbekistan, select those parts of the host body most exposed to the sun. Summer-feeding adults attach on the shaded undersides of the host (Galuzo 1943). Among the enemies of these ticks are wagtails (Motacilla spp.), one of which may eat as many as a dozen engorged female ticks from a cow. This bird, and the heron, are considered of importance in the control of H. detritum in Spain (Miranda. Entrenas 1954). FSU T EE InEIT Ina HEE MEATUS Gea aE GN RES Jarape a a ere See also life cycle of this form above, and ecology of "H. scupense" below. pay AUS Ecology of "H. scupense" The single-host type of life cycle, the rapid immature—_stage feeding period, the long lag between female feeding and oviposition, and the long period of egg laying of this form have already been commented upon, in addition to the phenomenon of adult feeding during the late winter and early spring months. Pomerantzev (1950) considers that the change in life cycle from a two-host to a single-host type has allowed "H. scupense" to spread further afield geographically than H. detritum. I is assumed that the remark refers to encroachment into otherwise inclement areas, for, so far as now imown, the overall geograph ical range of H. detritum is much more extensive than that of “i. scupense", In their work on the ixodids of Transcaucasia, Pomerantzev, Matikashvily, and Lototsky (1940) list both H. detritum and "H, scupense" as widely spread species in the southern part of the earctic Region. Only H. detritum is listed as a "spe cies typical for the desert and steppe formations of eastern Transcaucasia (including salsola, and wormwood-salsola forma. tions, gramineous-wormwood semideserts, Andropogonetum semi. steppes and the formations of highland nee of southern Transcaucasia)™". "H. scupense™ is included under "species equally occurring both in the forest zone of highlands and in desert and steppe formations". [ Several biological studies on this tick mentioned by Pomerantzev (1950) are unavailable to the writer./ Melnikova's (1953) study of ticks of the Crimean National Forest showed "H. scupense™ to be most common in populated areas adjoining the forest. It is closely associated with the grazing grounds of domestic cattle and wild deer and occurs at all altitudes except that it is much less common in mountain meadows near the peaks (1200 to 1400 meters elevation) and in coniferous forest (500 to 900 meters elevation). It is common in valleys with deciduous and mixed forest (400 to 600 meters), oak forest with many open meadows (600 to 800 meters), and beech forest with small, open fields. REMARKS Miscellaneous observations on H. detritum (= H. mauritanicum) by Sergent, Donatien, Parrot, and Lestoquard (1945) concern para. sitism of an engorged nymph by a nymph of the same species; re. generation of appendages mutilated in the immature stages; mon strosities; progeny of intermated adults from a single egg batch; and negative results of various attempts to crossbreed this spe. cies with H. excavatum. Most of these data derive from publica tions listed under ALGERIA above. Desportes (1938) described a nymph of H. detritum (= H. mauritanicum) with two anuses and a third spiracular plate. Other abnormalities have been noted by G. Senevet (1922A) and by Sergent and Poncet (1941). Abnormal specimens and gynandromorphs have been described by Pavlovsky (1940). According to Schulze (1921), "H. Soupense” may become over — grown by the host skin when it feeds on the scrotum of sheep. In attempts to determine whether feeding by tick larvae may produce host immunity preventing the feeding of subsequent larvae on the same host, Brumpt and Chabaud (1947) found that while pos— sibly some immunity may develop from the bites of Dermacentor ictus this does not hold when larvae of H. detritum are applied ater to the same animal. Skin lesions, a powerful salivary anticoagulin, and an eg¢ toxin that did not, hdwever, produce symptoms of tick paralysis have been studied by Hoeppli and Feng (1933). Schulze (19320) utilized males and females of H. detritum to illustrate his point that on unornamented ticks Tongitudinal furrows are found in constant situations; these indicate the place of attachment of the weakly transversely-striated dorso- ventral muscles, etc. In the same paper, the molting of "H. scupense" from nymph to adult is discussed in relation to — ornamentation of the body surface. A condition similar to polymorphic eczema among mammals has been described and illustrated for Yugoslavian females of = (Als) = "H. scupense" by Oswald (1938A). It is characterized by secre. tion of drops of serous liquid over the entire body surface. In one or two days, papules appear, followed by desquamation and later becoming scabby. The same author (1938B) described a toxin in the eggs of "H. scupense". DISEASE RELATIONS MAN: "H. scupense" (= H. volgense), harbors Pasturella pestis, the causative organism of plague, for several days after feeding on an infected animal and can transmit the bacteria by biting another animal; its feces also contains virulent organisms. "H. scupense" appears to be of negligible importance in the sustaining of certain Soviet foci of tularemia. H. detritum is considered by Soviet workers to be a possible vector of Uzbekistan hemorrhagic fever (virus). In North Africa and in Uzbekistan, H. detritum is a natural reservoir of Q fever (Coxiella burnetii), and the organism is transmitted from stage to stage of the tick and through the eggs to the subsequent generation. CATTLE: H. detritum is an important vector of theileriasis (Theileria dispar and T. annulata). "H. scupense" is also a vector of theileriasis and a carrier of brucellosis (Brucella spp.). HORSES: H. detritum is a vector of equine piroplasmosis (Nuttaliia equi) and possibly of Nuttallia minor. IDENTIFICATION The following remarks are based on typical specimens of H. detritum collected in Morocco, Egypt, the Sudan, Turkey, south. eastern Europe, and Russia. Remarks on variations are based on Iranian specimens determined definitely as H. detritum by Delpy, and on Middle and Far Eastern specimens in the Schulze collection. = 41675 Males: Typical males have the subanal shields directly pos— terior of the adanal shields; the lateral grooves are clear and uniform, reaching almost to the eyes; scutal punctations are either entirely absent, or number up to twelve superficial, um obtrusive punctations either scattered or more or less localized on the scutal surface; the appearance of the scutum is bright and shiny; the posteromedian groove is narrow and long, extending from the central festoon to the scutal midlength; the paramedian grooves are distinct, wider, and somewhat deeper, either as long as or shorter than the posteromedian groove; the area in which these grooves lie is usually depressed. The size is medium to large, the typical color is intensely black; the legs are red dish or yellowish brown and may be somewhat lighter on the an terior surface. The posterior margin of the scutum is quite rectangular, the scutal surface is usually arched. The specimen illustrated (Figure 158) indicates the maximum amount of punctation found in furrows or on the scutal surface; the punctations in the grooves of this specimen are especially numerous. A parma is either present or absent; if present it is small and inconspicuous. Subanal shields vary in size and shape from small to moderate size. The leg segments may have pale rings but these do not con trast greatly with the basic color of the segments. The following variations have been seen; Lateral groove on one side not so distinct as on the other side; or indistinct on both sides beyond the posterior third (these are most confusing specimens), in which case the lateral grooves continue anterior. ly as a row of contiguous punctations. Linear scutal depressions extending anteriorly from the posterior grooves frequently mar the scutal gloss. The scutal color may be various shades of brown. In greatly engorged specimens, the subanal shields are displaced from their typically posterior position to a lateral position, external of the central axis of the adanal shields; such specimens should not be confused with H. dromedarii or H. impeltatum. - 417 - Females: The genital apron is typically an almost equilateral triangle with slightly convex margins, and the posterior juncture is rounded or narrowly truncate, not narrowl inted. In profile, it is very gradually sloping (Figure OLE) aes unenforged but somewhat more abruptly sloping when engorged; in greatly engorged specimens the central area protrudes (Figure 161,D) sometimes with avery slight concavity centrally. The anterior margin may be straight or slightly concave or slightly convex, and in profile the posterior slope may be more acute than it is in typical fe males; in these cases there may be a very narrow bulge along the anterior margin. The shieldshaped scutum is longer than wide; smooth and shiny; with from four to ten superficial, scattered punctations of large diameter; punctations may be entirely lack. ing or a very few, shallow, minute punctations may be present. The scutal surface in greatly engorged specimens may be slightly rugose. The cervical grooves are long and wide, and reach the posterior scutal margin. Note: According to Pomerantzev (1950), H. detritum is largest in Middle Asia and gradually diminishes in size towards the west (Algeria). Color differences of various parts of the body do not support division of H. detritum into ceorraphical races. = The larva and nymph of H. detritum have been described and compared with those = other Russian species by Bernadaskaia (1939C). Feldman-Muhsam (1948) also described these stages and compared them with those of Palestinian species, Identification of "He scupense" The following remarks concerning the biological race "H. scupense" are from Pomerantzev (1950); H. scupense is similar to H. detritum and difficult to distinguish. It is smaller, has shorter legs, and has a more distinct caudal field in the male as well as greater scutal rugosity. The dorsal projection of the spiracular plate is broader and shorter. The capitulum is smaller and the porose areas of the female are more rounded than in H. detritum. The maximum size of the spiracular plate occurs in specimens from - 416 . Central Asia, where the species is almost indistinsuishable from H. detritun. "It may be best to regard "H. scupense" as a winter, Single host form of H. detritum".. Schulze collection material labelled H. scupense conforms to the above remarks but is not extensive enough to be of great value. It is difficult to decide what taxonomic disposition should be made of biological entities with slight morphological varia tions, such as "H. scupense" appears to be. It appears that the observations on this form have been carefully done, but the bio logical and taxonomic principles involved are unusual if not unique in the study of ticks. Pomerantzev's approach appears to have considerable merit. afte) Figures 162 and 163, dorsal and ventral views Figures 164 and 165, dorsal and ventral views A, Q, genital area. B to D, 9, genital area outline and profile. B, unengorged. C, partly engorged. D, fully engorged. HYALOMMA DROMEDARIT Egyptian Specimens PLATE XLVIII - 420. HYALOMMA TDROMEDARII Koch, 1844. (Figures 162 to 165) THE CAMEL HYALOMMA NOTE: Numerous early literature references to "H. aegyptium" apply in whole or part to H. dromedarii, but without study = indi_ vidual author's material it iis impossible to state the exact spe cies. In the pre-Delpy period, the name H. dromedarii was fre. quently used for Hyalomna ticks from camels, but the possibility that several species may have been included under this name indi- cates that caution should be exercised before it is assumed that all early references to Hyalomma ticks on camels refer to H. dromedarii. The various "subspecies" of H. dromedarii, now con. sidered invalid, and the few known synonyms of this species are noted below in the section on distribution. No type specimens conforming to present concepts for this species are available (see Feldman-Muhsam 1954) but since species criteria are now well defined, substitute type specimens should be selected and so designated in an established collection. DISTRIBUTION IN THE SUDAN Hyalomma dromedarii ranges extensively north of n2eN Ee ata tude but is entirely absent in the south of the Sudan. The camel hyalomma has not been previously recorded from the Sudan, but earlier references to H. aegyptium undoubtedly refer in part to H. dromedarii. Localities from which specimens have been seen (all from camels unless otherwise specified) are: Northern: Ed Damer and Shendi (camels and bulls; SVS). Berber (camels and horses; SVS). Wadi Halfa, Abu Hamed, Atbara, Ed Damer (camels and cattle; HH). Khartoum: Khartoum end Omdurman (common on domestic animals, especially camels; SVS, SGC, HH). Al raseehe: Kassala (camels and cattle; SVS). Port Sudan (cattle and horses; SVS). Tokar (donkeys; Svs). Darfur: Nyala (SVS). Muhagariya (donkeys, camels, horses, and cattle; SVS). Zalingei (camels, goats, and horses; SVS). Kordofan: El Obeid (camels and cattle; SVS). "Northern Kordofan™ (SVS). Umm Indiraba (cattle; svs). "Western Kordofan" (sheep; SVS). Blue Mile: Wad Medani (cattle and camels; HH). Hassa Heissa (Kohls cet., G. B. Thompson, correspondence). DISTRIBUTION H. dromedarii is common wherever camels occur: in southern Russia (Turkemnia, southern Tadzhikistan and Uzbekistan)*, and in the Far, Middle, and Near East. In Africa, it is found in North Africa, in the transitional belt just south of the great northern deserts, in the eastern and coastal lowlands as far south as the Somalilands, and in northeastern Kenya. Small in. troduced populations have been found in the Union of South Africa and in Southwest Africa, but whether they now survive is unknown. In some areas, e.g. Anatolia, the camel hyalomma feeds on other hosts since camels are now considerably less common than hereto fore. The distribution of this species has been mapped by the American Geographical Society (1954). It is of interest to note that within the range of H. dromedarii, from the Eastern Desert of Egypt to Afghanistan, there exists a more localized but highly distinctive parasite of bactrian, or two-humped, camels, H. schulzei Olenev, 1931 (see page 525), which also attacks Gromeda ries. *Pomerantzev (1950) lists the synonymous "H. asiaticum" from des. erts and semideserts of southern and eastern Transcatcasia and a considerable part of Kazkhstan. In the south its distribution extends to the boundaries of the Soviet Union, embracing a con siderable part of Turkmenia, Uzbekstan, and Tadzhiskstan; also Iran and central Asia. = 22 Note: All references below are to "H. dromedarii”®™ unless otherwise specified. OUTLYING ISLANDS: CANARY ISLANTS (As i. d. canariensis;: Schulze ana Schlottke 1930. As He d. Gromedarii: Kratz 1940). NORTH AFRICA: MOROCCO (Lavier 1923. Blanc, Bruneau, Martin, and Maurice 1026. Blanc, Martin, and Bruneau 1949). ALGERIA (As H. aezyptium dromedarii and as H. a. margaropoides: Senevet 19228). TUNTSTA (Pavesi 10e,. Tonelli-RondelTi 1030A. Colas_Belcour 1931. Colas-Belcour and Rageau 1951). LIBYA (Franchini and Cadeddu 1927. Franchini 1927,1928B,1929A,B,E. Stella 1938. Tonelli-Rondelli 19304 ,19324,C ,D,1935. Gaspare 1933. As H. anatolicum zavattarii: Tonelli-Rondelli 1935. Stella 193&. Kratz 190). EGYPT (Koch 13844. Nazaty 1947. Taylor, Mount, Hoogstraal, and Dressler 1952. Wassif 1954). WEST AFRICA: FRENCH WEST AFRICA (Blanc, Martin, and Bruneau 1949. Villiers 1955). EAST AFRICA: SUDAN (Hoogstraal 1954B. The "H. dromedarii £. leptosoma™ attributed to Schulze by Kratz 1940, on the basis of material from Sudanese cattle, represents weak, poorly ceveloped individuals of H. dromedarii, H. excavatum, or H. impeltatum; it is most likely the last-named Species). ETHIOPIA (Stella 1940). ERITREA (As H. tunesiacun ganorai: Tonelli-Rondelli 19320. Stella 1940. Kratz IOJ0. As Hu. a. dromedarii: Tonelli-Rondelli 1930A. As H. dromedarii:” Niro 1955). FRENCH SOMALILAND (Hoogstraal 1953D). ITALIAN SOMALILAND (Pavesi 1834A. Paoli 1916. Tonelli-Rondelli 1926A,1935. Fran chini 1929. Niro 1935. Stella 1939A4,1940). BRITISH SOMALILAND (Specimens from camels at Las Anod in BMNH collections; HH det.). KENYA /Lewis! (1931B,1934) H. dromedarii from the Rift Valley and from the Masai Reserve are misidentifications (material checked by HH). H..dromedarii does, however, occur in the arid Northern Frontier Province (material sent to the writer by Miss J. B. Walker for determination). This species has been reported by Daubney (1937) and Mulligan (1938) without further details except that the material had been identified by Schulze.7 = 428) oO [ TANGANYIKA specimens identified by E. A. Lewis as H. drome darii have been reported by Cornell (1936). This improbable record should be verified by checking the material./ SOUTHERN AFRICA: Although southern Africa is out of the normal range of . cromedarii. Theiler (correspondence) has material of this species from camels introduced into the Vryburg and Gordonia districts on the old camel patrol route with head quarters at Rietfontein on the border of the Union and Southwest Africa. Another collection is from the ears of a camel at Okalan ca in the extreme west of Ovamboland, Southwest Africa. This material was collected between 1939 and 1942, before the dis. continuance of the camel corps. Whether H. dromedarii survives in these localities is not known,/ rz NEAR EAST: PALESTINE (Bodenheimer 1937. Adler and Feldman. Muhsam 1936,19485. Feldman-Muhsam 1948). YEMEN (Sanborn and Hoogstraal 1953. Hoogstraal, ms.). IRAQ (Khayyat and Gilder 1947. Hoogstraal, ms.). TRANSJORDAN, SAUDI ARABIA, and ADEN (Hoogstraal, mS.) TRAN (Delpy 1936,1937B,1946A,1949A,B,C. Telpy and Gouchey 1937. WNemenz 1953. As H. yakimovi persiacum: Olenev 1931A,C. As H. yakimovi: ee aie 193 2a Asin delpyi: Schulze 1936D). TURKEY (Koch 1844. Kratz 1940. Kurt pinar 1954. Mimiogiu 1954. Common throughout central and eastern Anatolia, even where camels have recently become scarce; rare in western Anatolia: Hoogstraal, ms.). RUSSIA: As H. asiaticum or H. a. asiaticum; Galuzo 1935. Pomerantzev 1937. Bororoditsky and Bernadskaia 1936. Bernadskaia 19398,C. Chumakov, Petrova, and Sondak 1945. Pervomaisky 1947. Markov, Gildenblat, Kurchatov, anu Petunin 1946. Olenev 1950. Pomerantzev 1950. Rementsova 1953. Pavlovsky, Pervomaisky, and Chagin 1954. Tselishcheva 1954. Petrisheheva 1955. Zhmaeva, Pchelkina, Mishchenko, and Karulin 1955. Serdyukova 1955. As H. asiaticum caucasium Pomeranizev: in Pomerantzev, Matikashvily, ana Lototsky 1940. Pomerantzev 1950. As H. dromedarii: Yakimov 1923. Olenev 1929B. Pomerantzev 19341950. "Pavlovsky and Pomerantzev 1934. Kurchatov 1941. Pomerantzev, Matikashvily, and Lototsky 1940. Pavlovsky 1940. Chumakov, Petrova, and Sondak 19/5. Alfeev 1943,1951. Pervo- maisky 1949. SS ba eK As H. dromedarii asiaticum: Schulze and Schlottke 1930. Olenev 1929A,193514,C, 1932. Galuzo 1934. Pomerantzev 1950. Sone of these may apply to H. excavatum, this is especially likely in the case of Galuzo 1937. Feldman-iMuhsam 1954 does not accept Delpy's synonymy of this form under H. dromedarii but does not provide reasons or suggest a substitute. As H. aegyptium dromedarii: Yakimov 1917,1922. As "H. tunesiacum amurense Schulze": described and illus. trated by Kratz 1940. Delpy 1949B attributed "H, amurense Olenev, 19310" to H. excavatun. As H. pavlovskyi: Olenev 1931C. Galuzo 1935. As H. yeas enev 1A,C. Galuzo 1935. Pospelova.Shtrom MIDELE EAST: AFGHANISTAN (Hoogstraal, ms.). INDIA and PAKISTAN Zs HW. asiaticum citripes: Schulze (1934X). As H. d. citripes: “Kratz (1940). According to Delpy (19498) these ‘are synonyms of H. dromedarii. Feldman_Muhsam (1954) states, without further explanation, that they are not. Records of H. dromedarii by Sharif (1928) and Sen (1933) are questionable.7 FAR EAST; TIBET (and MONGOLIA): As H. kozlovi: Olenev 1931B7 Kratz 1940. As H. asiaticum kozlovi: Pomerantzev 1950. INDOCHINA: The H. d. indosinensis of Tounanoff (1944) refers to H. marginatum. 7— IMPORTED SPRCIMENS: ARGENTINA (Found on camels from Dakar: Lahille 5 NOTE: Portugese material under this name (da Silva Leitao 1945) 1s assumed (HH) to be misidentified. HOSTS Until recently, only the hosts of adult H. aromedarii were known. These are chiefly camels, but also include cattle and a HR horses anc, to some extent, sheep, coats, and doss. Man is um commonly attacked by adults, but more frequently by larvae and nymphs. H. dromedarii is so intimately associated with camels that it does not occur outside the normal range of these animals. However in parts of Anatolia where camels are now considerably less numerous than heretofore, large numbers of acults have been oe on cattle and horses, fewer on sheep and goats (Hoogstraal, mS.). In nature, remote from large concentrations of domestic ani_ mals, larvae and nymphs feed on small burrowing mammals and hares, rarely on lizards. Adults venture forth in search of larger hosts. Nymphs appear to be more versatile, depending on local situations, and may infest camels, cattle, and horses in large numbers. Factors inducing the selection of hosts by nymphs are in need of study. In the laboratory, Delpy and Gouchey (1937) fed larvae on hares and calves but this stage rarely ensorged on camels or sheep. The same was true for nymphs. Adults attached rapidly to camels and cattle, rarely to sheep. It was often difficult to rear on a calf the Fj generation of a female collected on a camel. Further review of Delpy and Gouchey's report is presented in BIOLOGY below. In Yemen, Southwestern Arabia, numerous larvae and nymphs have been collected from the following hosts (Hoorstraal, ms.): Lepus arabicus arabicus Ehrenberg Lepus arabicus subsp. Rattus ratvtus rattus Linnaeus (rare) Gerbillus cheesmani maritimus Sanborn and Hoosstraal In Ecypt, including Sinai, nymphs have been taken from hedgehogs, hares, rodents, and lizards (identifications based on adults reared from nymphs) (Hoogstraal, ms.). Hemiechinus auritus aegyptius Fischer (common on Mediterranean littoral) Paraechinus aethiopicus dorsalis Anderson ana De Winton ew hosts examined LAS) Lepus capensis sinaiticus Ehrenbers (few hosts examined) Lepus capensis aefyptius Lesmarest (common) Gerbillus (lipodillus) quadrimaculatus Lataste (rare) Gerbillus g. cerbillus Olivier (fairly common) Gerbillus P+ pyramiaun Geoffroy (common) Meriones c. crassus Sundevall (common, also in burrows) Meriones S. Shawi Duvernoy (common, also in burrows) Psammomys 0. obesus Cretzschmar (common, also in burrows) Jaculus j. jaculus Linnaeus (rare) Acanthodactylus boskianus Daud, (Lizard) (uncomnon) According to Pomerantzev (1934) hosts of the immature stages in Armenia include reptiles and wild birds. The fatty sub cutaneous layers of ground squirrels, Citellus sp., used as laboratory hosts (Pospelova-Shtrom 1937), may hinder the attach. ment and feeding of larval H. cromedarii (= H. yakimowi). In Russia (Pomerantzev 1950), adult hosts are camels, horses, and sheep while nymphs feed in large numbers on cattle and camels. Hosts of acults of the synonymous H. asiaticum are camels, cattle, horses, and sheep; rarely donkeys, goats, domestic and wild pigs, heres and hedgehors; sometimes man. Nymphs of this latter form occur on hedgehogs, in burrows of larre "peschanki" and gophers, while larvae are often on hedgehogs. Both immature stages are found (?tocether) on hares, gophers, jerboas, "peschanki", cats anc dogs, and single nymphs are found on cattle and sheep (Bosoroditsky and Bernadskaia 1938). BIOLOGY Life Cycle The question of the normal number of hosts of H. dromecarii is moot. Delpy and Gouchey (1937) consider it as a three_host tick that may utilize only two hosts under unfavorable conditions; i.e. great heat, when larvae molt quickly and reattach as nymphs nearby on the same host in order to avoid desiccation. It would appear that normally, on burrowing mammals, this need would not arise since the hosts venture from their relatively cool tunnels only after the sun goes down (deserts are usually cool at night). SATS Alfeev (1951) reared H. dromedarii as a single host tick on rabbits and noted that adults move to new positions shortly after molting and remain unattached to the host for a day or two afterwards. Field observations suggest that this is normally a two host tick, the change in hosts usually occurring after the nymphal. adult molt, infrequently after the larval_nymphal molt (Hoogstraal, see In Russia the former type appears most common (Pomerantzev 1950). Experimentally, Pospelova-Shtrom (1932) bred H. dromedarii (= H. yakimowi) on one, two, or three hosts. For the single host life cycle, she employed the hedgehog. The highly artificial conditions and exceptional hosts utilized in this experiment pre- clude additional deductions from the results. 7 During the warm season in Iran, the briefest life cycle ob served by Delpy and Gouchey (1937) was 93 days: During cold weather, 280 or more days were necessary. Two generations a year may occur in nature. Variations in life cycle length are due to external or climatic factors affecting oviposition, hatching, and premolting periods; feeding times are similar at all seasons. These findings are diametrically opposed to those of Pospelova. Shtrom (loc. cit.), who concluded, after rearing two generations under different temperature and humidity conditions, that varia. tions in the length of different stages depends more on host body. temperature than on atmospheric differences. The Delpy—Gouchey conclusions are more in line with usual concepts concerning factors affecting tick life cycles. (See page 704 In Egypt, some slight seasonal variation in incidence of adults on camels is noticed (Hoogstraal, ms.). Normal life cycle activity appears to continue the year around, except that it is slower during the winter. For instance, during the summer, nymphs molt to adults from seventeen to 26 days after dropping from the host while in winter this period is extended to from 27 to 48 days. Feeding time for each stage, according to Delpy and Gouchey, is as follows: = [HPI STAGE DAYS EXTREMES Larva 6 eB) ro) ll Nymph 7 patous? Female ial Te tor 5 These authors reported individual females laying from 2000 to 8000 eggs, but Pomerantzev (1950, for the synonymous H. asiaticum) notes as many as 14000 eggs. Males mate with unengorged or with feeding females while on the host. In the absence of males, females either detach pre. maturely from the host or remain fixed for an exceptionally long time — as long as two months. In the Kazalinski district of Russia (Pomerantzev 1950), adults parasitize domestic animals from April to the end of October, but are most common from May through August while larvae and nymphs attack hedgehogs in July and August. Fe. males that become engorged late in autumn undergo a winter diapause and do not oviposit till spring. Ecology Delpy and Gouchey (1937) found H. dromedarii to be very tolerant of low humidity and extremes in temperature (0°C. to 37°C.). Unfed larvae and especially unfed nymphs avoid dry atmosphere more than engorged stages and adults. H. dromedarii is probably the most completely desert-~adapted of all ixodid ticks. In remote Siwa Oasis of Egypt, it, together with R. sanguineus, is the only common ixodid tick. Italian and French observers elsewhere in North Africa and Soviet workers in Russia have made similar observations. The camel hyalomma ap. pears to be equally at home in all desert situations where ani. mals occur, as well as in semidesert and steppe areas. In Yemen and Eritrea, it is also common in mountainous areas to 7000 feet altitude wherever there are people and camels (Hoogstraal, mS.). When traveling over remote parts of the African and Arabian deserts engorged females have been observed desperately crawling = Ae = on sand. The opportunities for dispersal of H. dromedarii over long camel routes may easily be imagined, but a Targe number of ticks undoubtedly perish in either the egg or larval stage along these wide and indefinite trails through barren deserts. Blanc, Martin, and Bruneau (1949) report females on camels that had ar. rived, after a trek of longer than a week, across the Sahara, at Goulimine in southern Morocco from Mauritania, over a thousand kilometers away. In Egypt, similar infestations are found in the Cairo area on camels just arrived from the Sudan (these also bear other Sudanese but not Egyptian species of ticks) and in Siwa Oasis on camels from distant parts of Libya. In Egypt, unfed adults may be taken on the desert of the Mediterranean littoral at any time of the year. Unfed adults either come rushing at any potential host, including man, from under desert shrubs or are collected in rodent burrows before they depart in search of larger hosts. Engorged, ovipositing females may be observed in rodent burrows as well as under des— ert shrubs, in camel yards, and under stones. On the littoral desert unfed adults are seen at any time of the day in all sea sons. REMARKS In attempts to determine whether larvae produce an acquired immunity in the hosts, thus preventing subsequent larvae from engorging on the same animal, Brumpt and Chabaud (1947) fed this stage on rabbits with negative results. Differences in size of each stage and sex have been presented by Campana-Rouget (1954). A capillary tube arrangement, which has proved successful for the artificial feeding of adults of H. dromedarii for physio. logical and disease-transmission studies, has been described by Chabaud (1950A). Parasitic wasps oma) Hunterellus hookeri, have been bred from nymphs of H. dromedarli G LE asiaticum) in Russia by Bernadskaia (1939B). - 430- Integumentary sense organs, which are fixed in number and location and essentially similar in all stages, though more primitive in larvae, have been described and illustrated by Dinnik and Zumpt (1949). These are the organs that Delpy (1938) had previously referred to as spiracles ("stigmates respiratoires"). Pervomaisky (1949) was unable to rear a full Fj generation from parthenogenetic females of H. dromedarii. Abnormal specimens have been described by Pavlovsky (1920) and Alfeev (1948). The feeding of large numbers of this tick (= H. asiaticum) induces inflammation of the host skin that hinders normal engorge- ment, especially of females that are likely to die as a result. When additional species compete for space, "a further antagonistic factor increases the obstacles" (Pavlovsky, Pervomaisky, and Chagin 1954). Warburton and Nuttall (1909, page 71) produced an excellent illustration of H. dromedarii (H. aegyptium) but the legend, inferring South Africa as the source, is misleading. Apparently the only specimen from South Africa is the abnormal one (figure 18), the identity of which is uncertain. "H. asiaticum", which Delpy considers to be a synonym of H. dromedaril, 18 still treated by Soviet workers (Pomerantzev 1950) as a separate species with several subspecies and with a wider geographical range than H. dromedarii. This form is less robust, smaller, and more slender, with a shorter posteromedian groove than in giant males typical of H. dromedarii. The fe males are more elongate than the typically robust female H. dromedarii and the length-width ratio of their palpi and of Their scutum is longer. Such individuals are also encountered in African populations. Delpy (1946) first considered the short, wide palpal shape to be diagnostic but further studies indicated so much variation that "applied strictly to determination of isolated specimens, (this character) would have led to errors. (Also) the ratios that are true for a young, recently molted tick are no longer accurate when this tick has aged and become engorged", The shortness of the posteromedian groove appears concomitant with the general lack of robustness in these smaller and more slender ticks. - 431 - Feldman-Muhsam (1954) states "The type specimens of H. drome darii asiaticum as well as H. asiaticum citripes differ from Hi. dromedarii and are not synonyms of H. SS Set Without further explanation, it is impossible to evaluate this remark. DISEASE RELATIONS MAN; A host and vector of Q fever (Coxiella burnetii). Experimentally, this tick can be infected with the virus of Russian spring-summer encephalitis, with the virus of msquito— borne autumm encephalitis from the Russian Maritime Province, and with the virus of a Japanese mosquito_borne encephalitis. CATTLE: Theileriasis (Theileria spp.). This tick (as H. asiaticum) is apparently not a carrier of brucellosis (Brucella Spp- ee CAMELS: Theileriasis (Theileria camelensis). IDENTIFICATION Males: Typical males are recognized by (1) large size (average length: 5.7 mm., range 5.2 mm. to 7.0 mm.; average width: sep 8 mm., range 3. Oomnssto 5.0 m.), (2) distinct lateral displacement of the eabeaell shields, (3) short, deep lateral grooves limited to the posterior third of the scutum; and (4) few, large shallow punctations variously scattered over the surface and complete or almost complete absence of other puncta tions. Also distinctive is the deep, usually narrow, postero median groove extending from the distinct parma to the scutal midlength. This groove is bounded on either side by converging ridges; laterad of these ridges are deep, wide, often rugose paramedian grooves. The paramedian grooves are of variable length, and posteriorly delineate the two median pairs of fes. toons, which are often massive and which are larger than the more or less distinct three lateral pairs of festoons. Another pair of parallel ridges may be present between the paramedian grooves and the lateral grooves. a ey Variations of the above characters are as follows: Small punctations may be present and rarely may be obtrusive enough to suggest H. impeltatum; also a line of large punctations ex tending eae from the apex of the lateral grooves may sug- gest H. impeltatum. In engorged specimens, at least, differences in the posterior area of the scutum separate such specimens. In some specimens a dense field of small punctations may be present posteriorly, suggestive of H. excavatum. The subanal shields are sometimes divided into two pairs, but very few such specimens have been seen during the present study. In unengorged males, the subanal shields are closer to the central axis of the anal shields than otherwise, but their center is still exterior of the central axis of the adanals. Color varies from yellowish brown through dark brown to brownish black. The legs are usually paler than the scutum and the segments may be ringed by a paler band. Females: Typical females may be recognized by (1) the nar. rowly elongate and triangular genital aperture, which in profile slopes gradually, (2) the comparatively wide scutal outline (lengthwidth ratio about equal, (3) few, large, scattered punc- tations and frequent rugosity of the scutal surface, and (1) large size. The palpi are usually twice as long as their com bined width. The color varies from yellowish brown to black; the legs are usually paler than the body and may be concolorous or ringed. The genital aperture is most distinctive and hardly variable. In newly molted females its outline is slightly concave sub apically, but in engorged specimens the lateral margins are usually straight. The narrowly triangular outline is apparent. ly never lost. A narrow, raised integumental fold surrounds the apron, The scutal outline is widest just anterior of the eyes. The wide, deep, long cervical grooves are usually rugose as is also the scutal surface of most engorged specimens. From ten to twenty (average thirteen) large punctations are scattered over the central field and an equal number occur in each scapular field. Average newly molted specimens measure about 5.5 mm. long by 3.2 mm. wide, but engorged individuals may be 30.0 mm. long and 15.0 mm. wide. Scutal length is about 3.2 m., width about 2.3 m. - 433 - The larva and nymph (as H. asiaticum) have been described and compared with those of other species by Bernadskaia (193%) and (as H. dromedarii) by Feldman-Muhsam (1948). - 434 - Figures 166 to 167, &, dorsal and ventral views Figures 168 to 169, 9, dorsal and ventral views A, 9 genital area. B to D, 9 genital area outline and profile. B, unengorged. C, partly engorged. D, fully engorged. HYALOMMA EXCAVATUM Egyptian Specimens PLATE XIL - 435 - HYALOMMA EXCAVATUM Koch, 1844. (Figures 166 to 169) THE SMALL HY ALOMMA NOTE: Schulze and his co-workers employed the name H. savignyi (Gervais 1844) for this tick, and following their usual practice appended a variety of subspecific names to it. References to H. depressum, H. lusitanicum subspp., H. pusillus subspp., and Loy Si icephaloides also apply to H. excavatum. The name H. anatolicuw, as used by Russian and French workers, applies to H. excavatum. For a list of synonyms, see Delpy (1949B, pp. 475-577). Adler and Feldman-Muhsam (1948) and Feldman-Muhsam, following Schulze'ts lead, applied the name H. savignyi in studies of H. excavatum. After examining Koch's type specimens, Feldman-Muhsam (1954) agreed with Delpy that H. excavatum is the proper name for this tick. This is accepted as a final decision. DISTRIBUTION IN THE SUDAN H. excavatum is moderately common in northern Sudan and in the northern parts of central Sudan. No records from Darfur Prov. ince are available. The small hyalomma is generally considerably less numerous in the Sudan than in Egypt. It shows a strong predilection for horses, even in areas where other domestic animals are present in large numbers. The following are data for material seen: Northern: Shendi (bulls, donkeys, and horses; SVS). Wadi Halfa, Abu Hamed, Atbara, and Shendi (camels, cattle, horses, donkeys, goats, and sheep; HH). Khartoum; Khartoum, Omdurman, and Shambat (camels, horses, donkeys, cattle, goats, sheep, and dogs, SVS; Gordon College collection; HH). - 436 - Kassala; Kassala (goats, horses, cattle, and camels; SVS). Port Sudan (cattle, donkeys, and horses; SVS). Sinkat (horses; SVS). Tokar (cattle, horses, and donkeys; SVS). Darfur: No records. Kordofan: El Obeid (horses and sheep; SVS). Blue Nile; Wad Medani (camels and horses; HH). Hassa Heissa (camels; G. B. Thompson, correspondence). DISTRIBUTION H. excavatum is common throughout northern Africa, and ranges through the Near East, Asia Minor and southern Russia to India. It is abundant locally in southern Europe, but is particularly numerous in Egypt, Palestine, Asia Minor, and southern Russia. In Africa, H. excavatum ranges along the northern and north. eastern littoral, a continuous belt characterized by less than ten inches of rainfall per annum. Its southeastern limit is Somalia. The Sudan has been invaded by this parasite possibly both via the Nile from Egypt and via the Red Sea coast. The distribution of H. excavatum has been mapped by the American Geographical Society (1954). All references below are to H. excavatum unless otherwise noted. ATLANTIC OCEAN: CANARY ISLANDS (Nuttall lot 3226 in EMNH; H.H. det. As 38 depressum: Schulze 1919. As H. lusitanicum cicatricosum: Schutze and Schlottke 1930. Kratz : NORTH AFRICA: MOROCCO (Blanc, Bruneau, Martin, and Maurice 1948. Blane and Bruneau 1953,1954,1955. As H. lusitanicum: Blanc and Bruneau 1949). ALGERIA (As H. 1. lusitanicum: Senevet 1922B,1925,1928A,B, 1937. Senevet and Rossi 1925. Kratz 1940. dtarces 1952. As - 437 - H. lusitanicum berberum; Senevet 1922B. As H. lusitanicum ‘algericum: Senevet 928A. As H. savignyi: Sergent and Poncet TS37-T0L0,1913. Sergent and co-workers also employed the name H. lusitanicum at one time or another. As H. lusitanicum algericum: Schulze and Schlottke 1930. As H. aegyptium im pressum: Senevet 1922B). a TUNISIA (As H. tunesiacum: Schulze and Schlottke 1930. As H. tunesiacum tunesiacum: Kratz 1940. As H. lusitanicum depressum: ochulze and Schlottke 1930. Kratz 1920. As H. usitanicum algericum: Colas-Belcour 1931. As H. excavatum: Colas-belcour and Rageau 1951). LIBYA (As H. depressum; Franchini 1927,1929E. As H. tunesiacum franchinii: Tonelli-Rondelli 1932C. As H. anatolicum: Tonelli-Rondelii 1932D. As H. fezzanensis: Tonelli=Rondelli 1935. Stella 19380. Kratz 1940. Wumerous specimens in Hoogstraal collection). RGYPT (As H. excavatum sp. nov.: Koch 1844. As H. rhipi- cephaloides: Neumann 1901,1911. Schulze 1919,1921,1936F. Kr&tz TSt0- As H. pusillum alexandrinum; Schulze 1919. As H. savi al savignyi: Kratz . As H. excavatum: Daubney and Said opal SRenaees in part confused with H. impeltatum. Taylor, Mount, Hoogstraal, and Dressler 1952). 7~ EAST AFRICA: SUDAN (Taylor, Mount, Hoogstraal, and Dressler 1952. Hoogstraal 1954B). ERITREA (Specimens from several localities in HH collection. Apparently not reported under any known synonym by Italian workers; material probably differently identified by them). FRENCH SCMALI- LAND (Hoogstraal 1953D). ITALIAN SOMALILAND (As H. lusitanicum: Franchini 1927,19290,E. Niro 1935. As H. somalicum: Tonelli. Rondelli 1935. Stella 1939A,1940. Kratz 19J0). ?KENYA: H. anatolicum, a synonym of H. excavatum has been reported in various notes by Lewis, and by Daubney (1937), Mulli- gan (1938), and Yalvac (1939). This name derives from material identified by Schulze. Although it would not be surprising to find isolated populations in northeastern Kenya, recent workers have not encountered it and earlier specimens referred to this species are not available./ - 436 = NEAR EAST; PALESTINE (As H. anatolicum: Bodenheimer 1937. Kratz 1940. As H. rhipicephaloides Neumann 1901,1911. Schulze 1921,1936F. Kratz Sp s H. savignyi: Bodenheimer 1937. Adler and Feldman-Muhsam 1946,19Z8. Pol eae sen 1947,1948, 1949,1950,1951A. As H. excavatum: FeldmanMuhsam 1954. As H. tunesiacum: Bodenheimer 1937). ” SYRIA and LEBANON (Hoogstraal, ms.). IRAQ (As H. aegyptium mesopotamium: Schulze 1919. Schulze and Schlottke 1930 he Tr s He Savignyl mesopotamium: Kratz 1940. As H. excavatum:; Hubbard 1955. Hoogstraal, ms.). “ARABIA" (As H. pusillum: Schulze 1919). TRUCIAL OMAN, YEMEN, ADEN, SAUDI T oogstraal, mss.). IRAN (Delpy 1946B,1949C,1952). AFGHANISTAN (Anastos 1954. Hoogstraal, ms.). TURKEY (As H. illus: Vogel 1927. As H. excavatum: Kurtpinar 1954. Mimioglu 1954. One of the most common ticks on the Anatolian steppes: Hoogstraal, ms. As H. lusitanicum: Yasarol 1954). CYPRUS / The H. savignyi exsul of Schulze and Schlottke (1930) , ee ecia bat py (19Z95B) to H. marginatum (= H. savignyi of Delpy), appears rather to be H excavatum; see pp. Pa 535 of Kratz (1940) 7. EUROPE: PORTUGAL (As H. lusitanicum: Koch 1844. Schulze 1919. Kratz 1940). ———————— SPAIN (As H. depressum; Schulze 1919. Gil Collado 1948A. As H. excavatum: De Prada, Gay, and Llorente 1950. De Prada, Gil Collado, and Mingo Alsina 1951. As H. lusitanicum: Gil Collado 1936,1948A. Kratz 1940. As He Tusitanicum algericum: Jordano Barea 1951. NOTE: H. depressum is considered to be a synonym of H. excavatum, but the species called H. depressum by Gil Collado 19Z5A is one that cannot readily be determined). FRANCE (As H. excavatum: Brumpt and Chabaud 1947. Brumpt 1949. Buttner 1949. Colas-Belcour and Rageau 1951. Chabaud and Choquet 1953). ITALY (As H. lusitanicum: Schulze 1936. Tonelli-Rondelli 1938. Kratz 1940). GREBCE (As H. anatolicum: Kratz 1940. Enigk 1947. Pandazis 1947). - 439 - RUSSIA: As H. anatolicum; Pomerantzev, Matikashvily, and Lototsky 1940. Galuzo 1944. Blagoveshchensky and Serdyukova 1946. Lototsky and Pokrovsky 1946. Pervomaisky 1954. Pav- lovsky, Pervomaisky, and Chagin 1954. Viazkova and Bernadskaia 1954. Gajdusek 1956. As H. anatolicum anatolicum: Serdyukova 1946A,B. Markov, Gildenblat, Kurchatov, and Petunin 1948. Pomerantzev 1950. Pervomaisky 1950A. Gajdusek 1953. Tselishcheva 1953. As H. anatolicum excavatum: Serdyukova 1941. Pervomaisky 1949,1950A. Pomerantzev 1950. Petrisheheva 1955. As H. amurense: Olenev 1931A,C. As H. asiaticum caucasicum: Pomerantzev, Matikashvily, and Lototsky As H. excavatum:; Blagoveshchensky and Serdyukova 1946. Zhmaeva, Pchelkina, Mishchenko, and Karulin 1955. As H. turkmeniense; Olenev 1931A,C. Kornienko-Koneva and Shmulreva 1022. Chumakov, Petrova, and Sondak 1945. Pomerantzev 1946. Markov, Gildenblat, Kurchatov, and Petunin 1948. As H. tunisiacum turkmeniense: Kratz 1940. Delpy (1949B) considered H. turkmeniense as questionably a synonym of H. excavatum; Pomerantzev (1950) synonymizes it under iH. excavatum (= i. anatolicum). ?As H. savignyi armeniorum: Olenev 1929A. Schulze and Schlottke 1930. Lototsky and Popov 1934. As H, armeniorum: Kratz 1940.7 & ?As H. savignyi: Zolotarev 1934. Galuzo 1935,1941,1944. Bern skaie ISCAS. Pavlovsky 1940. Zotova and Bolditzina 1943. Galuzo, Bolditzina, and Kaitmazova 1944. For a discussion of Delpy's remarks concerning Soviet confusion between H. excavatum and H. marginatum (= H. savignyi i) see page 470. yf As H. rhipicephaloides: Yakimov 1922,1923. Olenev 1929B. = 440 = MIDDLE EAST: INDIA /As H. kumari; ‘Sharif (1928,1930). Delpy (19Z9B) considers H. kumari as a synonym of H. excavatun, but it seems best to reserve Judgement on this matter for the moment. Numerous specimens typical of H. excavatum are present in BMNH collections, H.H. det. As H. savignyi; Dasgupta (1955) and Dasgupta and Ray (1955); the possibility that these refer to H. marginatum should be considered7. PORTUGESE INDIA (Santos Dias 19525). HOSTS H. excavatum is a parasite of cattle, horses, donkeys, camels, buffaloes, sheep, goats, and swine. It also attacks man and dogs. Hares appear to be especially important wild hosts. Nymphs are variable in occurrence on cattle, but nymphs and larvae are often found on calves. Nymphs and larvae frequently attack rodents, and normally do so on the desert. They also feed on man, hares, lizards, and birds. All stages of H. excavatum have been observed feeding on hares in a forest near Casablanca where other wild and domestic animals are absent (Blanc and Bruneau 1953,1954,1955). In Ana tolia, numerous adults have been reared from nymphs taken from hares (Hoogstraal, ms.). A larva has been reported from a hare in Iraq (Hubbard 1955). In Yemen these animals are heavily infested by immature stages (Hoogstraal, ms.). British Museum ee, History) collections contain specimens from Indian hares Nuttall lot 3423; H.H. det.). Wherever hares and H. excavatum occur together the association appears to be an important one. The complete absence of any specimens of H. excavatum on more than five hundred hedgehogs collected throughout Egypt, is noteworthy (Hoogstraal, ms.). Hedgehogs were, however, used as laboratory hosts of immature stages by Feldman-Muhsam (1948). Delpy (1949C) considers birds, especially nestlings, impor- tant hosts of nymphs. A male in British Museum (Natural History) collections has been reared from a redstart, P. phoenicurus (= Ruticilla pluvenicurus) at Amara on the Tigris River (Nuttall lot 32,0; FH. ° CUe)eo Sab eK Single instances of attack of human beings have been reported from France (Buttner 1949) and Iraq (Hubbard 1955; whether actual. ly feeding not stated). During field work for the present study, feeding specimens of H. excavatum have been taken from personnel in Egypt, Turkey, and Yemen (Moogstraal, ms.). In Uzbekistan, this tick (= H. anatolicum) often attaches to man (Gajdusek 1953). Apparently the only larger wild animals yet recorded as hosts of the adult stage are gazelles in French Somaliland (Hoogstraal 1953E). Biological observations in Egypt thus far have been confined to searching for naturally infested wild animals in the field, keeping them alive in the laboratory, and allowing ticks that drop from them to molt to the next stage. Adults reared from nymphs taken from wild animals have been from the following hosts: Lizard Acanthodactylus boskianus (fairly common) Lesser Egyptian gerbil Gerbillus g. gerbillus (common ) Greater Egyptian gerbil CGerbillus p. amiaum (common) Fat sandrat Psammomys 0. obesus are common) Sundevallts jird Meriones c. crassus fairly common) Shaw's jird Meriones Ss. shaw (fairly common) Spiny nouse Acomys spp. eeu) Lesser Egyptian jerboa aculus j. jaculus (uncommon Hares Lepus Capensis subspp. (common) Colas.Belcour and Rageau (1951) report adults in Tunisia from burrows of gerbils, jirds, and fat sandrats and nymphs from jirds. They also found H. excavaium in burrows and on other rodents in France. Adults of H. excavatum in rodent burrows are always newly molted, remaining there before they venture forth to seek a larger host (Hoogstraal, ms.). There is no evidence to consider gerbils as common hosts of adults, as stated on the map of the American Geographical Society (1954); see also Erratum sheet). BIOLOGY Life Cycle The several investigators who have reared H. excavatum in the laboratory (Delpy 1952 in Iran; Daubney and Said 1951 in Egypt; = hie Feldman-Muhsam 1948 in Palestine; Brumpt and Chabaud 1947 in France, and Serdyukova 1946A in Reais) confirm that this is normally a three-host species. In Tadzhikistan, however, Lotot— sky and Pokrovsky (1946) consider H. excavatum (= H. anatolicum) to be a two-host tick. Feldman-Muhsam observed that Some larvae may remain on the host through the nymphal stage, but Delpy (19460) stated that if they do so, they first detach and wander away, for example to the ear, and reattach only after molting. Daubney and Said observed a single larva molting while still attached. On desert rodents in Egypt nymphal H. excavatum have on several occasions been found attached to the host and par- tially enclosed by the larval exuvia. Possibly in these situa tions, where hosts are scarce, the typical life cycle is more commonly somewhat altered. On Egyptian deserts, the molt from nymphal to adult stage typically occurs in rodent burrows. Remarks that desert rodents dislodge most ticks attached to them by rubbing, shaking, or eating are contrary to frequent experience in Egypt. The effect of a small size host on the life cycle of H. excavatum has perhaps best been described by Serdyukova (1946a, as FH. anatolicum) (from abstract in Review of Applied Entomology): "Larvae from a single egg-batch engorged on a rabbit, which is an unusual host for this tick. Some detached after engorgement, others molted on the ani-_ mal. Some of the resulting nymphs wandered on the rabbit without feeding but others engorged and then dropped off. Larvae placed on the ears of a calf all detached after engorging, and no engorged or molting larvae or larval exuvia were observed on calves in the field. Ticks collected in a calf shed included freshly engorged and molting larvae and unfed nymphs. It is concluded, therefore, that H. excavatum (= H. anatolicum) develops as a three-host tick on its normal host, but that an unusual host may alter this behavior. The cycle of ixodid development has prob- ably altered as a result of evolutionary processes. The type of development that occurs on the usual host should be considered as normal, and deviations from it on unusual hosts as atavistic." hs) = It is significant that the Russian worker considers the rabbit to be an atypical host of immature stages of H. excavatum. This is far from true in North Africa and Arabia, where hares and other smaller animals are frequently parasitized (rabbits do not occur here). Most probably smaller animals are also parasitized in Russia but workers there, who have been occupied chiefly with veterinary problems, have failed to investigate this possibility. Indeed, it seems that a diametrically opposed, theoretical con clusion might be drawn; Under primitive conditions, in areas lacking large numbers of domestic animals, H. éxcavatum spends part or all of its life cycle on small animals, usually no larger than hares. In these situations it undergoes a one or two-host type of life cycle. However, when herds of larger domestic animals are present in the range of this tick, the adults and sometimes the immature stages may be confined to these animals. On these larger ani- mals, H. excavatum undergoes a three-host type of life cycle, which is an atypical one for the species, in fluenced as it is by the availability of hosts due to human activities. This matter is obviously in need of further investigation. With regard to the seasonal cycle of H. excavatum, the aforementioned authors working in Iran, Egypt, and Palestine, as well as Serdyukova (1946B) working in the semi-deserts of Tadzhikstan, agree that engorged nymphs and young adults hi- bernate in cracks and crevices of buildings during the winter; the Soviet observer states that larvae may also overwinter under these conditions. In Russia, hibernating ticks were taken in, among, under, and in association with wooden fixtures of animal enclosures, but not under cakes of dung plastered on loose walls. When these structures were removed the incidence of ticks found in these yards the following summer was only a fraction of what it had been the previous year. In the deserts of Egypt engorged nymphs and unfed adults overwinter in rodent burrows. In spring, ticks that have hibernated venture forth to feed. Under experimental conditions, if they are removed to a warmer oy Nii place during the winter nymphs molt and adults feed. In Egypt, wild nymphs do not normally molt until almost summertime, pos— sibly because of cold nights during early spring. By late March, however, females begin to feed, and at this time they may oviposit very shortly after dropping from the host —~ as soon as seven days afterwards. The incubation period may be as short as 27 days, so that in May new larvae and last year's nymphs can be found feeding. Under the Russian conditions already mentioned, oviposition does not commence until July or August and it appears that there is only a single generation a year in those climes. In Tadzhikistan (Lototsky and Pokrovsky 1946), adults of H. excavatum (= H. anatolicum) infest cattle from the end of February to November, and Larvae and nymphs from July through September; all stages are most numerous early in August. Larvae feed for from four to six days (32°C., 75% R.H.) according to Feldman-Muhsam, but the Russian observer reported only two to four days in nature. Larvae molted six days after leaving the host (30°C.), after four or five days at 38°C., and after thirty to fifty days at 17.5°C. (after six to twelve days in Russian observations). Nymphs fed a week later and remained on the host from nine to twelve days (four to six days in Russia) but the time between dropping and molting to the adult stage varied greatly (twelve to twenty days in Russia). At 35°C., the molt to females occurred in from eleven to 24 days and to males in from twelve to 35 days. At 30°C. both sexes appeared between eleven and 56 days after nymphs dropped from their host. The duration of one generation at 32°C. was estimated at 116 days, but it was concluded that under field conditions some of these stages may be shorter and there may be three generations a year. Parthenogenesis in H. excavatum has been observed by Pervo— maisky (1949) who found That a few F1 females could be reared from eggs laid by females in the absence of males. Ecology Feldman-Muhsam (1947) observed that unfed larvae and nymphs are much more sensitive to humidity than to temperature. At any 5 PANGS ¥ temperature survival of unfed immature stages increases with higher humidity. In laboratory tests, larval longevity varied between two and 241 days (mean 1.25 and 162.6 days), nymphal longevity between ten and 246 days (mean 6.8 and 149.2 days). The author considered that under undisturbed natural conditions survival would have been longer than in these tests in which daily counts were made. Nymphs are less susceptible than larvae to low humidity. The length of nymphal life increases directly with humidity and inversely with temperature. Other life history and hibernation studies by Feldman-Muhsam (1949) are of interest, but since they apply to frigid Palestinian winter conditions that do not occur in the area under consideration this work is not presently per- tinent. Serdyukova (1946A) observed ticks detaching from their hosts at night; larvae and nymphs mostly between nine and eleven o'clock in the evening. A number of larvae and nymphs placed on a calf in the morning became fully engorged during the day but ‘remained on the calf when darkness set in. This behavior, probably an adaptation to local climatic conditions, protects the ticks from exposure to the direct rays of the sun, which are fatal to them. They are apparently inactive during the hot part of the day, and attach to hosts at night as has been recorded for other Hyalomma species under desert conditions. In an isolated plot only slightly over five percent of a counted number of unfed ticks attached to calves between noon and sunset. H. excavatum is obviously a tick of xeric regions not neces. sarily associated with domestic animals although populations are considerably larger where these animals occur. Experience in the comparatively lightly-vegetated desert areas of the Mediterranean littoral of Egypt, where it infests rodents in their burrows, indicate this tick's habits away from large, dense flocks of domestic animals. In Transcaucasia, H. excavatum (= H. anatolicum) is typical of various desert, semidesert, and steppe formations, but does not occur in forested zones (Pomerantzev, Matikashvily, and Lototsky 1940). This agrees with observations in Anatolia (Hoog- straal, ms.). With respect to altitudinal distribution in Trans. - 446 - caucasia, these authors say H. excavatum falls in group in which the "upper limit of distribution ..... 1S ..... inversely pro- portionate to the moisture of the climate and to the amount of rainfall and directly proportionate to the height of the snow line during the summer period". In Armenia, H. excavatum (= H. LATE armenorium) is found in pasture at 6500 feet elevation an gher (Lototsky and Popov 1934). During the present study it has been found at similar heights in Sinai, Yemen, Anatolia, and Eritrea. Feeding Sites and Reactions Adults feed on cattle chiefly on the scrotum and perineum and in the inguinal and axillary areas. Nymphs generally feed on the neck, chiefly along its crest. Larvae are not commonly found on Egyptian cattle. Note: The following section is ancillary to further remarks on the “subgenus Hyalommina" (page In Egypt (Hoogstraal, ms.), larvae and nymphs of H. excavatum are frequently found completely overgrown by rodent host skin. This phenomenon is especially common among young jirds, Meriones shawi shawi Duvernoy, and sometimes on young fat sandrats, Psammo-— S 0. obesus, on the Mediterranean littoral. During springtime most every nestling jird in the vicinity of Mersa Matruh is infested in this manner. The ticks can be detected by lumps under the skin, most frequently around the neck, axillary areas, shoul. ders, and flanks. Some of these rodents have as many as 22 im mature ticks under the skin. The host skin may partially or completely enclose the ticks*, which are almost always misshapen when removed. Some nymphs extricated from under the skin of jirds have molted to adults in our laboratories. They have in. variably been tiny, weak, misshapen, poorly developed, pale spec. imens, which, if identified according to criteria offered by Schulze (1919), would be H. rhipicephaloides Neumann, 1901, in the subgenus Hyalommina. ~ *Nuttall (1914B) has quite accurately described this processes as an oedematous swelling of the host skin, as a result of irritation when the long mouthparts of Ixodes (and Hyalomma) ticks reach firm subcutaneous connective tissue; the oedematous swelling may gradual. ly engulf the feeding tick. - 447 - Schulze (1921) recognized this growth phenomenon for mate rial of "H. nn picephaleiies from a "steinbock" (probably meaning an ibex) near the De ea in Palestine, although he continued to apply a species name to these runts. Yet, Schulze and Kratz have presumed to refer to this as a "half endoparasitic type of para. sitism" by ticks. Large number of ticks were found in pale red dish cysts in the ibex's subcutaneous tissue, especially in the axillae. Holes in the host skin could not be detected. In several instances we have found cast larval skins en casing subdermal nymphs. Young jirds raised to adulthood in the laboratory yielded dead H. excavatum in the middle of the summer when the hosts were sacrificed, and nymphal skins were found around them. We have no evidence that a Hyalomma tick overgrown by the host's skin can force its way out.through the skin. Pavlovsky, Pervomaisky, and Chagin (1954), in preliminary studies, have also indicated that when H. excavatum (= H. anato_ licum) feeds in large numbers on a restricted area of the host, poorly developed specimens result. Females especially do not fully engorge and may even die due to inflammation of the host skin. When other species also compete for a restricted area of the host skin, an additional antagonistic factor increases the chances of abnormal development or death. Prolonged infestation on rabbits by H. excavatum under lab. oratory conditions does not confer host immmity, preventing en gorgement by subsequent larvae, against this species or against Dermacentor pictus (Chabaud 1950A, Brumpt and Chabaud 1947). REMARKS A capillary tube arrangement, which has proved successful for the artificial feeding of adults of H. excavatum for physio. logical and disease.transmission studies, has been described by Chabaud (1950A). Schulze (1932C) illustrated the leg segments (of "H. anato- licum) to support theories of tick ornamentation. In The same = LO es work, he illustrated a cross section of the scutum (of "H. lusita. nicun" ) and compared it with that of A. cohaerens. Gynandromorphs and malformed specimens of this species have been described and illustrated (as H. savi : Pavlovsky Sey by Pervomaisky (1950). A gynandromorph of a: excavatum (= H. savignyi) has been described by Feldman-Muhsam (1950) but ~ Campana-Rouget (1950) considers this to be an "intersexue" (see also Chabaud and Choquet 1953). An abnormal male (as H. kumari) has been illustrated by Sharif (1940). The rate of growth and comparative differences among those morphological parts that are either similar or different between the two sexes of this tick have been studied by Chabaud and Cho. quet (1953). Cuticle growth has been mentioned by Lees (1952, as H. savignyi). Campaniform sense organs have been briefly described (Das— gupta 1955). Specimens from Kenya, identified as H. anatolicum, were used by Yalvac (1939) to describe features of development of the adult stage in nymphs. DISEASE RELATIONS MAN: The virus of Uzbekistan hemorrhagic fever has been isolated from H. excavatum (= H. anatolicum) in Soviet Central Asia, where this tick appears to be at least an important natural reservoir if not a vector. Experimental work with H. excavatum (= H. turkmeniense) indicates, for the virus of Russian spring.summer encephalitis, transmission by bite and transovarial transmission, and the same for the virus of Japa nese (mosquito-borne) endephalitis except that transmission by biting was not obtained. These viruses, as well as that of Russian (mosquito-borne) encephalitis, persist for many months in infected ticks. H. excavatum is commonly found infected with the rickettsiae of Q fever (Coxiella burnetii) in North Africa, southern Europe, - 449 - and Uzbekistan. Its importance as a vector of this disease to man requires investigation. If, as appears likely, it is true that the "H. savignyi" of Soviet workers with ticks and brucellosis applies actu Viton Hs excavatum, it should be noted that hereditary transmission and subsequent infection of the host by the bite of this tick is claimed. The spirochetes of certain Russian relapsing fevers do not survive in this species (as H. anatolicum excavatum) for even a day. ay CATTLE: An important vector of theileriasis (Theileria annulata). BIRDS: Benign piroplasmosis of nestlings. IDENTIFICATION Males: Typical males are very distinctive but in almost any field collection a large number of atypical specimens may be found. Characteristically, the center of the subanal shields is posterior of the central axis of the adanal shields. This holds true for all flat, unengorged and slightly engorged indii viduals. Males that have not flattened after molting and before being preserved, and engorged males, especially those that have fed on large animals such as camels, almost always have the subanal shields borne on an udder—like swelling and laterally displaced as in H. dromedarii. (The subanal shields are al ways small and elongate, frequently minute or even hardly dis. tinguishable). Such specimens can be distinguished by smaller size and by the characteristic strong depression of the posterior part of the scutum between two smooth lateral ridges; this de pression is almost always densely punctate. (Some atypical H. dromedarii tend towards a resemblance of this last character). The lateral grooves are very short, restricted to the posterior third of the scutum. A line of punctations frequently continues anteriorly from the lateral grooves; these may be groove—like enough to confuse keying the specimen. The scutum, away from a oO the depressed caudal area, has rare, widely scattered, medium-size punctations, or none, but atypical very small and superficial punc- tations may rarely confuse this pattern. A pale parma is frequent— ly present; the festoons are greatly variable in distinctness. The scutum is usually strongly convex, and all specimens are definitely small in size for Hyalomma ticks (scutal length no more than 4.18 mm., rarely over 3.75 mm.; width no more than 2.19 m., rarely more than 2.19 mm.). Certain populations that key to H. excavatum but measure above the upper level of this range represent distinct spe— cies of uncertain identity (see pages 880 to 886). Females: The knob-like genital apron is more or less (but al. ways definitely) bulging in profile; it may be circular, elongate— ly triangular, or widely triangular (but if so always distinctly mich smaller than in H. marginatum or similar species) in outline; the circular outline is Most characteristic and distinctive; the elongately triangular outline is fairly common and usually fairly distinctive; the widely triangular outline is not common but is apt to be confusing. The scutum is extremely variable in color and in length-width ratio, but it has very few large punctations scattered in the central field, a few more in the scapular areas, and some— times some to many very fine, superficial punctations over much of its surface. The scutal surface of engorged specimens frequently becomes extremely rugose. Typical engorged females are comparative— ly small and narrowly elongate but quite thick dorsoventrally thus presenting a narrowly rectangular appearance. The larva and nymph (as H. series) have been described and compared With those of other species by Bernadskaia (1939) and by Feldman-Muhsam (1948). - 451 - Figures 170 and 171, o, dorsal and ventral views Figures 172 and 173, 9, dorsal and ventral views A, Q genital area. B to D, 9 genital area, outline and profile. A and B, unengorged. C, partly engorged. D, fully engorged. HYALOMMA IMPELTATUM Egyptian Specimens PLATE L eS Gy HYALOMMA IMPELTATUM Schulze and Schlottke, 1930 (= H. BRUMPTI Delpy, 1946A). (Figures 170 to 173) KRATZ'S HYALOMMA NOTE: The name impeltatum, proposed by Schulze and Schlottke (1930) in a brief key to Hyalomma ticks, was said to apply to a subspecies of H. savignyi. Subsequent workers were unable to recognize this tick on the basis of the brief data provided. The original material was redescribed and illustrated by Kratz (1940). Kratz also indicated that Tonelli-Rondelli's (1932C) H. eS! from Eritrea is probably a synonym. Delpy overlooked bo : erythraeum and Kratz's description of impeltatum and (1946A) escri H. brumpti as a new species from Cameroons. It now appears that Tagecam rumpti is a synonym of H. impeltatum. The status of H. erythraeum is still moot. DISTRIBUTION IN THE SUDAN In the Sudan, H. impeltatum is restricted to the drier cen tral Provinces. It may occur Locally in Northern Province, but no records are available. Kassala: Kassala and Sinkat (cattle; SVS). Port Sudan (donkeys and cattle; SVS). Kordofan; El Obeid and Umm Inderaba (cattle; SVS). “Northern Kordofan™ (camels; SVS). See also Khartoum below. Darfur: Sibdo (horses; SVS). Muhagariya (camels, cattle, horses, and donkeys; SVS). See also Khartoum below. Khartoum; A large number of specimens taken from cattle from Kordoran and Darfur Provinces at the Khartoum Quarantine Station, where they were enroute to Egypt (HH) .7 - 453 - Males of H. impeltatum are frequently found in rather small numbers on Sudanese cattle at the Cairo abattoir. This species is present near the Sudan frontier in the southeastern corner of Egypt (HH). DISTRIBUTION H. impeltatum is thus far known to range from Iran and Arabia to fe and Libya; into the Sudan, Eritrea, French Somalit land, and restricted areas of Kenya and Tanganyika; and westwards in localized areas of central, western, and northwestern Africa. Within this area, H. impeltatum appears to be present usually in widely scattered foci. However, it is anticipated that, as the identity of this species becomes better known, certain gaps in our knowledge of its distribution will be closed. The absence of reports of this tick by Palestinian workers causes one to wonder under which name it is being identified there. Recent finding of small numbers of H. impeltatum on wild and domestic animals in single localities of Kenya and Tanganyika, a unique distributional pattern for a Near Eastern Hyalomma, sus. gests that this species is slowly extending its range into East Africa. NEAR EAST; IRAN (As H. brumpti: Delpy 1949C). TURKEY (Specimens from Istanbul abattoir, on sheep said to be from Bedlise: Hoogstraal, ms.). IRAQ (Specimens from a number of localities in HH collections; others from Amara in BMNH; H.H. det.). PALESTINE (Material from Gaza in BMNH; H.H. det.). YEMEN (Fairly common: Hoogstraal, ms.). NORTH AFRICA: EGYPT (As H. brumpti: Hoogstraal 1954A. One of the most numerous of local ticks; frequently taken on cattle from the Sudan at the Cairo abattoir: Hoogstraal, ms.). LIBYA (Numerous specimens from many localities in HH collection. There is a suspicion that the H. marginatum balcanicum of Tonelli- Rondelli 1930A from Tripoli and Bengasi may refer to He impeltatum). MOROCCO (Material from Mogador in BMNH; HH det.). - 454 = WEST AFRICA: “WEST AFRICA“ (As H. savignyi impeltatum: Schulze and Schlottke 1930). RIO DE ORO (SPANIGH GAARA): (As H. impeltatum: Kratz 1940). FRENCH WEST AFRICA (As H. brumpti: Rousselot 1948,1951,1953B, and Villiers 1955. Material from Mauritania in BMNH collections; H.H. det.). NIGERIA (ds H. brumpti: Unsworth 1952. Material from Kano in BMNH collec_ tions; H.H. det.). IVORY COAST (As H. brumpti: Rousselot 1948. Record not repeated by Rousselot TOS but specimens checked by Theiler). CENTRAL AFRICA: CAMEROONS (All as H. brumpti: Delpy 1946a, 19494,B. Rageau 1951,1953). EAST AFRICA: SUDAN (As H. brumpti: Hoogstraal 1954B. As He impeltatum: Feldman_Muhsam 1952). ERITREA (Specimens from near Karkobat in HH collection. ?As H. erythraeum: Tonelli-Rondelli 1932C; Niro 1935; and Stella 1939A,1940; see NOTE above). ITALIAN SOMALILAND (?As H. erythraeu: Tonelli-Rondelli 1935 and Stella 1940; see NOTE above). ae SOMALILAND (Material in HH collection}. NOTE: H. marginatum balcanicum of Tonelli-Rondelli (1930A) from Eritrea, Somalia, and Ethiopia may refer to H. impeltatum; this name was also used by Stella (1938A,1939A,19%0) for meter material from Italian Somaliland. KENYA (Material in BMNH from sheep at Laisamis, Northern Frontier District; H.H. det.; collected by E. A. Lewis who had determined it as H. impressum albiparmatum. Numerous specimens in BMNH collections from cattle a agadi and Shombole). TANGANYIKA (Miss J. B. Walker has sent a large series of typical specimens for identification; material from a rhino. ceros and wildebeest at Mto-wa-Mbu, a few miles north of Lake Manyara in northeastern Tanganyika, April 1952, A. C. Brooks legit). - 455 - HOSTS All domestic animals are attacked by adults of this tick. Available data does not indicate that among these animals H. impeltatum shows any marked host predilection. In Egypt a number of specimens have been taken feeding on personnel during field trips. Wild animals known to be infested are gazelles in Egypt (HH), wild pig in Eritrea (HH), rhinoceros and wildebeest in Tanganyika (Walker records above), and caracal in French West Africa (Villiers 1955). Hosts of immature stages are rodents, hares, birds, and man. At Amara, on the Tigris River in Iraq, Lt. R. A. Buxton reared adults from nymphs taken from hares and from a redstart, P. phoenicurus (= Ruticilla pluvenicurus) (Nuttall lots 3239 and 3240 in BMNH). In Egypt we have reared many adults from nymphs that have dropped from both the lesser and the sreater Egyptian gerbils, Gerbillus g. gerbillus and G. p. pyramidum, and fewer from the following animals: Lesser Egyptian jerboa, Jaculus j- jaculus; fat sandrat, Psammomys o. obesus; Egyptian hare, Lepus capensis aegypticus; and man. Although Rousselot (1948) reared this species, he furnished no data on the hosts of the immature stages either in the labo ratory or in the field. BIOLOGY Life Cycle Rousselot (1948) claimed that H. impeltatum (= H. brumpti) is a three-host species that in eo West Africa Laboratory completed its life cycle in about three months. Results of studies in NAMRU_3 (Cairo) laboratories will be presented when completed. - 456 - Ecology Although H. impeltatum is a tick of arid and semiarid regions, small populations aes exist in certain African savannah areas. Biological and ecological characteristics and limitations of this species are still poorly known. As noted above, immature stages are found in common association with desert and desert edge rodents in Egypt. In Nigeria, where H. impeltatum is almost entirely confined to the more arid northern provinces, it is sometimes the most common tick collected on cattle and appears to require a drier climate than do H. truncatum (= H. transiens), H. impressum, and H. ru. ores (Unsworth 1952). Adults are found aon the anus and ers and in the axillary regions of their hosts. DISEASE RELATIONS Apparently this tick is not a vector of Theileria annulata of cattle. Note, from Egyptian records, that nymphs and adults are known to feed on man in nature. REMARKS The comparative size of each stage and sex has been noted by Campana_Rouget (1954). The remarks below are based in part on specimens originally identified (as H. Ss by Dr. L. P. Delpy, on his remarks (correspondence) on s material, and on our further observa tions of additional collections consisting of some 2000 spec. imens. IDENTIFICATION Males. In TYPICAL specimens, (1) the exterior position of the comparatively large subanal shields, (2) the lateral grooves that extend anteriorly at least to the scutal midlength, and (3) - 457 - the numerous, moderate size (few large), shallow, scutal puncta. tions that are uniformly and widely distributed over most of the scutal surface, is a combination of characters easily separating males from those of all other species. Variation in each of these characters are as follows: (1) In specimens that have fed, the subanal shields are always situated well exterior of the axis of the adanal shields; they are usually borne on a slightly rounded, elevated protrusion of the ventral integument, and usually extend posterior beyond the body margin. However, in unfed specimens, where the sub. anal shields are still closely appressed to the ventral integuw. ment, these shields may appear to be in line with the central axis of the adanal shields. Close observation reveals that the base of the subanal shields is in an exterior position but that the unique tilting of the subanal shields in a medially-directed position gives the first impression that they are situated direct- ly posterior of the adanal shields. In fed males, the subanal shields are usually vertical and parallel. (2) The lateral grooves are usually well delineated and extend from the festoons in a progressively more shallow line almost to the eyes. In some spec. imens, they are more or less obscured, at the level of the scutal midlength and anteriorly, by scutal punctations; questionable spec. imens should be tilted towards the source of the light. In other individuals, the anterior extension of the lateral grooves con. sists chiefly of a distinct row of more or less contiguous puncta. tions; such specimens may be confused with H. dromedarii, and, if the subanal shields are still closely appressed to the ventral integument, possibly even with H. excavatum. (3) Punctations are usually very slightly larger, deeper, and more dense posteriorly than elsewhere on the scutum. Punctations over the scutum are typically dense but not contiguous, regular, medium-size with a few scattered larger, deeper ones among them, and fairly shallow. The number and placement of these punctations is subject to con siderable variation; in some specimens the central scutal area may be almost devoid of obvious punctations; this is especially true in engorged individuals. Other characters are as follows: The area just anterior of the festoons is almost always slightly depressed and contains a long, narrow posteromedian groove, and a pair of shorter, wider, SP ADO = and deeper paramedian grooves. A parma, the color of which may be lighter or darker than the rest of the scutum, may be present, or may appear as a normal median festoon. Two definite pairs of festoons and two more or less fused pairs lie on either side of the parma or median festoon. Delpy states that the scutum is flat, actually it is usually more or less arched, especially in males that have fed. The scutal color varies from dark brown to black; excep. tional specimens, usually very small ones, may be lighter. The leg segments are usually pale anteriorly and posteriorly and darker centrally, but they may be entirely pale yellowish. Female; The scutum posteriorly and centrally has numerous rather regularly spaced, moderate size, noncontiguous punctations. Scattered among them are several larger and deeper punctations in two parallel rows centrally. The moderate size punctations are usually mostly discrete, but exceptions to this are common. Anteriorly and in the scapular areas, punctations are large and deep; in the lateral fields punctations are absent or present. The deeply depressed cervical grooves are more or less rugose, and the punctations in them are more or less contiguous. The scutum of engorged specimens frequently has less distinct punc— tations and grooves. The scutum is generally dark brown in color. It is definitely longer than wide, but the ratio may be reduced in some newly molted, misshapen, or greatly engorged specimens. The genital area is distinctive. The central genital apron is an elongate triangle much like that of H. dromedarii but shorter, wider, and usually not quite so narrowly pointed apical. ly. In profile, it definitely bulges anteriorly and is depressed posteriorly. An important principal additional feature is that, in unmated specimens and in mated but not greatly engorged spec imens, this genital apron is bordered on each side by a slight bulge that gives the genital area a trilobed appearance not found in any of the other species with which H. impeltatum may be con fused. This characteristic is maintained = only slightly less distinctness in greatly engorged females. Female body size, in all except runts, is always large. The legs are like those of the male. - 459 - NOTE: Very small, rounded, globose, pale runts appear com. monly with typical specimens. Their diagnostic characters are frequently modified. In field collections, such specimens are virtually impossible to identify. They should not be confused with other species or cast into the subgenus Hyalommina. Among collections of reared adults, gradations from typical males to atypical males closely approximating exceptionally small spec- imens of H. dromedarii are frequently seen. Such atypical spec. imens are obviously poorly nourished in the immature stages and show numerous indications of lack of proper development. If encountered singly, they would be most difficult or impossible to properly identify. =e LOO t= Figures 174 and 175, &, dorsal and ventral views Figure 176, 9, scutum and capitulum, dorsal view Figure 177, 9, genital area. A, partly engorged, B to D, outline and profile; B and C, partly engorged; D, fully engorged. HYALGMMA IMPRESSUM ees an Specimens PLATE LI = 40s HYALOMMA IMPRESSUM Koch, 1844. (Figures 174 to 177) THE WEST AFRICAN HYALOMMA NOTE; In literature on African ticks various subspecific designations of H. impressum apply actually to H. rufipes or to H. truncatum and are treated herein under those species. The "H. umbeum impressum" of Soviet workers is H. rufipes. Koch's wee of H. impressum and of H. mere clearly distinguish these two species. The H. impressum mentioned by Adler and Feldman-Muhsam (1946,1943) ah Palestine resulted from mistaken identity of H. rufipes (Feldman-Muhsam 1954), as did Delpy's H. impressum ae) from Iran (Delpy 1949A,B). The H. impressum of Schulze 1918), from a single specimen collected from a horse in Macedonia, Greece, and presumed to have been carried there as a nymph on a migrating bird, is questionable; this record was repeated by Panda zis (1947). DISTRIBUTION IN THE SUDAN H. impressum is locally common on cattle in central Sudan west of the Nile. Numerous males and fewer greatly engorged fe. males reach the Khartoum quarantine station on Darfur and Kordo. fan cattle and many males are still attached when these cattle reach the Cairo abattoir. The West African hyalomma is not known to be established east of the Nile. Darfur: Zalingei and Sibdo (common on camels and cattle, also occurs on horses; SVS). Kordofan: Delami (cattle; SVS). El Obeid (camels, SVS). Khartoum: See three paragraphs above./ aoe = DISTRIBUTION H. impressum is a West African tick that ranges eastward into the Sudan. It has not been found south of the equator, in the great northern deserts of Africa, or on the Mediterranean littoral. WEST AFRICA: FRENCH WEST AFRICA (As H. impressum sp. nov.: Koch 1822. As fi. gpl impressum typica: SONICS 1919, and iS) e m™m Chodziesner 1922. ressum impressum: Schulze and Schlottke 1930, and Kratz ; ot sete i intermedia: Girard and Rousselot 1945, and Rousselo . As H. Lmpressum: ' Rousselot 1953B). NIGERIA (As H. impressum: Unsworth GER Material in BMNH from Katagum; FH. TW. oe CENTRAL AFRICA: CAMEROONS (As H. impressum; Rageau 1951, 1 neg ee AFRICA (As H. impressum: Rousselot 1953B). EAST AFRICA: SUDAN (As H. impressum: Hoogstraal 1954B. Feldman-Muhsam 1954). ?UGANDA and ERITREA: As H. impressum: Wilson (1949B) and Tonelli-Rondelli (1930A). These may refer to H. rufipes or to H. truncatum. 7 HOSTS Domestic cattle are the chief hosts of H. impressum, and are mentioned by all authors. No other hosts have previously been reliably recorded. In Darfur, camels are frequently infested and horses are sometimes attacked. BIOLOGY Unstudied. DISEASE RELATIONS Unstudied. - 463 - IDENTIFICATION Males: This handsome species is easily recognized and readily distinguished by the following combination of characters: (1) center of subanal shields posterior of central axis of adanal shields, (2) scutum regularly covered by deep, rather large punctations that often tend to obscure the long lateral grooves, and (3) distinctive narrowing of the scutal margin posterior of the lateral grooves, giving it a rectangular outline posteriorly. A moderate amount of circumspiracular pilosity may in some specimens suggest H. rufipes. The festoons are more or less fused, but the central and two santas pairs are always distinct. The scutum is dark reddish brown to black and the legs are brightly ringed. The comparatively narrow, elongate scutal shape is in contrast to the wide outline of H. rufipes; the rectangular pos— terior margin is distinctive of H, impressum. Females: Numerous, regular scutal punctations, similar to those of the male, are distinctive; in some specimens they are more superficial but still numerous. The genital apron is broad ly triangular in outline; it is composed of an anterior narrowly transverse (bulging) ridge and a posterior button (sloping or depressed) not so wide as the anterior ridge. / With respect to the outline and profile of the genital area of this species, no specimens observed during the present study conform to that illustrated by Delpy (1949B, page 106), which is more narrowly triangular (equilateral) and gradually sloping in profile.7 A certain amount of circumspiracular pilosity is present in some females; differences in the genital aprons distinguish these specimens from females of H. rufipes, the scutal punctations of which may be quite similar to those of H. impressum. a LGh = Figures 178 and 179, @, dorsal and ventral views Figures 180 and 181, 9, dorsal and ventral views A, 9 genital area. B to D, 9 genital area outline and profile. B, unengorged. C, partly engorged. D, fully engorged. HYALOMMA MARGINATUM an Specimens PLATE LII - 465 - HYALOMMA MARGINATUM Koch, 1844. (Figures 178 to 181) THE MEDITERRANEAN HYALOMMA NOTE: H. marginatum appears in European literature chiefly as either H. mar meraupetue © or as Le savignyi and in recent Soviet literature as plumbeum e Adler-Feldman_Muhs am (1948) school Smployed Th the name H. “mar inatum, the Delpy (1949) school H. savignyi (Gervais, 1842) ieee the lead of Schulze. The MH. savi a "oe Adler and Feldman-Muhsam (1948) and of some of their ae is H. excavatum. Recently Feldman-Muhsam (1954) concluded, from study of the Koch (1844) type material, that H. mar marginatum is the correct name for this species although reasons for so doing are somewhat obscure. Pomerantzev (1950) states that the name H. plumbeum plumbeum (Panzer, 1795) refers to the present species. ths point merits further investigation. Feldman-Muhsam apparently has not con sidered the possibility of this synonymy. DISTRIBUTION IN THE SUDAN H. marginatum thus far is known from only two localities in central Sudan, one in the west and one in the east. Further search will probably reveal additional populations between these two places. Kordofan: Umm Inderaba (cattle; SVS). Kassala; Kassala (camels; SVS). DISTRIBUTION H. marginatum appears to be most common in southeastern Europe and southern Russia from where it extends eastward into India and Indochina and westward throughout southern Europe, and into the = 466 = Near East including Arabia, and North Africa. Elsewhere in Africa, small populations are scattered through the drier transitional areas just south of the northern deserts from the Red Sea to the Atlantic Ocean. Soviet workers in Transcaucasia consider this to be a "Mediterranean species* in contrast to the other hyalommas of their territory, which they classify as either "widely spread in the southern part of the Palearctic Region" or "peculiar to the zone of western deserts" (Pomerantzev, Matikashvily, and Lototsky 1940). The mapped distribution of H. marginatum (= H. savignyi) as presented by the American Geographic ciety (1954) does not correspond to the range of any recognized species in this genus. NORTH AFRICA; “NORTH AFRICA" (As i. marginatum balcanicum;: Schulze and Schlottke 1930). EGYPT (As Cynorhaestes eee Virey 1822. As H. savignyi; Hoogstra 5 populations exist on the Meena littoral and in desert edge situations of Nile Valley and Delta; somewhat more common in Sinai: Hoogstraal, ms.). LIBYA (Specimens from several localities in HH collection. As H. marginatum balcanicum: Stella 1938). TUNISIA (As H. tunesiacum: Schulze and Schlottke 1930. As H. savignyi: Colas-Belcour and Rageau 1951). MOROCCO (As H. savignyi: Blanc, Martin, and Maurice 1946, 1947A,B. Blanc, Bruneau, Martin, and Maurice 1948. Blanc and Bruneau 1949. Blanc, Martin, and Bruneau 1949. Blanc 1951). ALGERIA / According to Delpy (correspondence), the H. aegyptium of Senevet and subsequent French workers in Algeria, is H. marginatum (= Delpy's H. savi i).- The "H. aegyptiun life cycle studies reported by Wattert (1913) on Aare from Algeria refer to H. marginatum (HH determination of Nuttall lot 1305 in British Museum ataral History). Most of the references = Lot listed for Algeria on page 405 also contain notes on H. meson (as H. Sa) this species is apparently less common there than H. detri um. WEST AFRICA: NIGERIA (As H. marginatum annulipe : Schulze and Schlottke 1930). [Delpy (19498) synonymized H. impressum brunneiparma: Hi at ei tum Schulze and Schlottke, 1930 (from Togo) under H. marginatum (= H. savignyi), but this is actually a synonym of H. albi_ armatum (see page ). The "H. savignyi"® of Tendeiro (1948, POLS TOSIF »1952A,C ,D) from Portugese Guinea is shown by the same author's discussion (1949A) to be H. truncatum. 7 EAST AFRICA: SUDAN (As H. savignyi: Hoogstraal 1954B). fH. marginatum balcanicum, a synonym of H. marginatum, has been reported a Ethiopie and Somalia by Italian workers; these records may be correct but it is more likely that they refer to H, impeltatum.7 KENYA (A single @ in BMNH from a cow, Kisima, Nanyuki, May, 1932, collected after dipping by E. A. Lewis. This specimen is entirely typical of H. marginatum; H.H. det.). Z SOUTHERN AFRICA: Santos Dias (19500) noted that "H. savignyi" had been reported from Angola by A. Morais in 1909, and a this probably refers to H. truncatum (= H. impressum transiens). Fe me NEAR EAST: PALESTINE (As H. marginatum: Bodenheimer 1937. Adler and Feldman-Muhsam 1946,1948). and LEBANON (As H. savignyi: Pigoury 1937). TRANSJORDAN (Hoogstraal, ms.). ~ TURRET TAs H. marginatum baleanicum: Schulze and Schlottke 1930. As H. anatolicum: “Galuzo 1944. As H. savignyi: Kurtu pinar 1954 and Mimioglu 1954. As H. marginatum: eee ms.). CYPRUS / As H. cypriacum: iiniescaee eee 1930. Kratz 1940. The "H. savignyi exsul" of Schulze and Schlottke, 1930, attributed by Delpy CSUSB) to H. savignyi marginatum (= H. savignyi of Delpy) appears rather to be H. See cf. Reete Cot, pp. 551535) 7. TRAN (As H. savignyi: Delpy 19468, 19490). IRAQ (As H. marginatum: Hoogstraal, ms. Hubbard 1955). = 468%= ARABIA: YEMEN (As H. marginatum: Hoogstraal, ms.). EUROPE: SPAIN (As H. hispanum: Koch 1844. As H. savignyi iberum:; Schulze and Schlottke 1930. As H. iberum: Kratz Tarb- H. marginatum hispanum: Schulze and Schlottke 1930. Kratz 1940. As agrees marginatum SETA Gil Collado 1936,1948A. As H. marginatum:; Gil Co o 1948A. Perez Gallardo, Clavero, and Hernandez 1949,1952. As H. savignyi: Gil Collado 1948A. De Prada 1949. Parker, De Prada, Bell, and Lackman 1949. De Prada, Gil Collado, and Mingo Alsina 1951). “f* PORTUGAL: It appears likely that the "H. rufipes glabrata! of Fonesca, Pinto, Colaco, Oliveira, Branco, da Gama, ares Franco, and Lacerda (1951) may refer to H. marginatum; cf. page FRANCE (As H. marginatum: Enigk 1947. As H. savignyi: Lamontellerie 1954. Specimens from Estrel Mts. in EMNA collection; H.H. det.). ITALY (As H. marginatum: Koch 1844. Tonelli-Rondelli 1938. Fnigk 1947. Cavaceppi 1950. As H. marginatum marginatum: Schulze and Schlottke 1930). ALBANIA (As H. marginatum: Enigk 1947). YUGOSLAVIA (As H. marginatum; Enigk 1947. As H. savignyi: Angelovsky 1954. PetrovTtee Tose c : ROMANIA (As H. savignyi: Metianu 1951). GREECE (As H. aegyptium f. brunnipes: Schulze 1919. Kratz 1940. As H. marginatum balcanicum: ulze and Schlottke 1930. As H. marginatum: Inigk 1OL7. Pandazis 1947). BULGARIA (It is possible that the "H. ee of Pavlov 1940,1942, etc., refers to H. marginatum. As H. marginatum: Enigk 1947. Pavlov and Georgiev TS50y. GERMANY (As H. marginatum marginatum: Arrives as nymphs on migrating birds according to Kratz 1920). - 469 . "TNSUL BRIONI® (As H. marginatum brionicum: Schulze and Schlottke 1930. Totze 1933. Gossel 1935). RUSSIA: NOTE: According to Delpy (1949B), Soviet authors have frequently confused H. excavatum (= i. anatolicum) under H. marginatum (= H. savignyi). Most Russian reports of "H. savignyi" appear to apply to H. excavatum (H.H.). a Most Soviet records for H. anatolicum subspp. since 1948 can be considered as applying to H. excavatum and most records for H. marginatum (and in 1950 for H. p. plumbeum) as referring to what nS ae called H. marginatum. Raz. an intensive study of all 2 ATs CA att aac eA oe a eS is concluded that Delpy's remarks in this respect apply chiefly if not entirely to reports by Galuzo (1941 and 1944) and by Galuzo, Bolditzina, and Kaitmazova (1944) on ticks of Kazakstan and control of piroplasmosis vectors in that area. Correspondence with Dr. Delpy concerning this matter has not elicited a reply. On bio- logical grounds, it appears that Galuzo's "H. savignyi - H. anatolicun applies to H. excavatum (= H. anatolicum of Soviet workers). It is possible that the use of the name H. savignyi by Zotova and Bolditzina (1943), who reported on work with 4H. seen and H. savignyi in relation to attempts to infect ticks with brucellosis in the laboratory, also applies to H. excavatum. As H. marginatum: Olenev 1934. Pomerantzev 1934,1946. Pavlovsky and fe 1934. Lototsky and Popov 1934. Galuzo 1935,1941. Galuzo and Bespalov 1935. Arginsky 1937. Bernadskaia 193%. Kurchatov 1939A,B,1940A,B,C,D,E,F,G,1941A,B,C. Pomerant— zev, Matikashvily, and Lototsky 1940. Kurchatov and Sokolov 1940. Grobov 1946. Blagoveshchensky and Serdyukova 1946. Enigk 1947. Chumakov 1948A,B. Markov, Gildenblat, Kurchatov, and Petunin 1948. Piontkovskaia 1949. Gajdusek 1953,1956. Pritulin 1954. As H. marginatum marginatum: Serdyukova 1941. Grobov 1946. Piontkovskaia tor 51049. Paviovsky 1948. Korshunova and Petrova. Piontkovskaia 1949A. Pervomaisky 1949,1950. Gajdusek 1953. Tselishcheva 1953. As both H. marginatum and H. savignyi: Zolotarev 1934. Galuzo 1935 1941 TOLL Zotova and Soltitzina 1943. Galuzo, Bol. ditzina and Kaitmazova 1944. See two paragraphs above. - 470. As H. marginatum balcanicum and H. marginatum olenevi: Schulze and Schice tes 1o30-an . Olenev 1929A,1931A SS atz 1OZ0. As H. pagent bacuense (apparently of Schulze, ms.):; Olenev 1931A,C. s H. marginatum caspium (apparently of Schulze, ms.): Noted by Olenev 90, but described by Kratz 1940. As H. plumbeum plumbeum: Pomerantzev 1950. Piontkovskaia 1951. Meintkova 15 aw Ce: » plumbeum: Shatas 1952. Shatas and Bustrova 1954. Pavlovsky, Pervomaisky, and Chagin 1954. Arakian and Lebedev 1955. Pillipenko and Derevianchenko 1955. Petrishe. hevo 1955. Abramov 1955. Zhmaeva, Pchelkina, Mishchenko, and Karulin 1955. MIDDLE EAST: INDIA (As H. aegyptium f. typica: Sharif 1928). INDOCHINA (As H. dromedarii indosinensis: Toumanoff 1944). FAR EAST: ?CHINA: The "H. supressun rufipes" of Chodziesner (1924) is probably H. marginatum according to ee (1940, p. 554) 7 HOSTS The common hosts of adult H. marginatum are any domestic ani. mals, especially cattle and horses; atso goats, sheep, and camels often serve. In the Crimea, horses have been stressed as hosts by Kurchatov and Sokolov (1940) . A typical female taken from a dog at Amman, Transjordan, by Dr. B. Babudieri, has been seen (Hoog- straal, ms.). Nymphs may also attack domestic animals but are much more frequent on small wild mammals and birds, while larvae feed only on these small animals. Host preferences, especially of immature stages undoubtedly vary somewhat from locality to locality, but the impression of considerable variation between areas appears to be due to incomplete observations by various workers. In Transcaucasia, birds are said to be the chief hosts of immature stages (Pomerantzev, Matikashvily, and Lototsky 1940). In the laboratory, chickens have been used (Zhmaeva, Pchelkina, Mishchenko, and Karulin 1955). = A47L= In Anatolia, adults have been reared from nymphs from hares, hedgehogs, and partridges (Hoogstraal, ms.). Nuttall lot 3278 in BMNH consists of adults reared from nymphs from a hare on the River Tigris, 32°N., November 1917, by Captain P. A. Buxton; H.H. det. A single nymph has been reported from a hare in Iraq (Hubbard 1955). Hosts in Tunisia are cattle, sheep, porcupines, and hares. Adults are also found in gerbil nests (most probably newly molted, before venturing forth to find a larger host: HH). Nymphs have been taken from "Cochevis" (Galerida cristata) (Colas-Belcour and Rageau 1951). In southern Morocco, larvae and nymphs were re- ported from the nests of jirds, Meriones shawi (Blanc, Martin, and Maurice 1946,1947A,B), while others, presumably adults (same authors 1947B), were found on domestic animals and, at certain times of the year, on the grounds of native markets. In Egypt, including Sinai, nymphs, which have been reared to typical adults, have been found on two kinds of hedgehogs, Hemiechinus aegyptius auritus and Paraechinus aethiopicus dorsalis, on fat sandrats, Psammomys o. obesus, and on jirds, Meriones shaw1 and M. crassus. Equally important here are lizards, Acanthodacty— lus boskianus, while lesser gerbils, Gerbillus g&- erbillus are Tess frequently found infested by nymphs (Hoogstraal, ms.). In the Arax valley of Armenia, hosts of immature stages are stated to include reptiles and wild birds (Pomerantzev 1934). Recent Soviet workers on hemorrhagic fever in Crimea report that adults attack cattle, sheep, horses, and men. Larvae and nymphs infest the European hare, Lepus europaeus transsylvanicus, in Crimea but are not found on hedgehogs, bats, rodents, dogs, or wild carnivores. Some immature specimens were taken from gray partridges, Perdix perdix, prairie larks, vetanocors pee calandria, cranes, Grus grus, and sparrows and domestic chickens (cf. Gajdusek 1953 ,1956). An exceptionally interesting study of H. marginatum (= H. p. lumbeum) in the Crimean National Forest Reserve has recently Bae reported by Melnikova (1953), whose chart is reproduced below. eMC No. Hosts No. Hosts No. No. No. No. Max. No. Mean Per Host Examined with Ticks Larvae Nymphs c@ o9 Ticks on Host One Host Red Deer 12 22 fe) O 52 204 90 6.1 Roe Deer 36 i! @) ) A (0) ) 0.0 Hare 32 10 3a 714 (0535) 309 Yb 0) Squirrel 38 al 0 O. Wo 1 0.0 Jay 26 20 1,02 225 O)in W(8) 236 24.0 Cattle 147 64, O O 568 497 fal: 6.2 Pig 28 8 0 0 137 69 50 aS) Chicken 15 15 300 25 (0), (0) 70 21.6 As is easily seen, jays, chickens, and hares are the chief hosts of immature stages in the Crimean forest. Cattle, pigs, and red deer are important adult hosts, and hares may be of some im portance. The absence of ticks on the roe deer is noteworthy. Melnikova (loc. cit.) noted that unfed larvae enter the audit tory canals of jays and chickens and molt there to nymphs and to adults; he found 118 immature ticks in the ears of a single bird. In Eastern Anatolia (Hoogstraal, ms.) partridges with larvae and nymphs of this tick similarly tightly packed in their ears have been observed. The comparative ease with which these birds were shot or even caught by hand suggested that the heavy tick infesta tion impaired the birds! keenness. Infested birds seemed muddled and confused and ran in staggering circles rather than flying or dashing off as did most of the flock. BIOLOGY Life Cycle Life cycle studies of "H. aegyptium" reported by Nuttall (1913B) were undertaken with H. marginatum. Specimens resulting from this work are at present in the Nuttall collection at the British Museum (Natural History). Nuttall found that H. margi- natum may act as a two-host or as a three-host tick; he believed that the "peculiar" two-host life cycle, when nymphs were fed on - 473 - hedgehogs, was due to larvae remaining entangled among the spines. This is, however, probably the typical life cycle in nature. Nuttall summarized his findings as follows: PERIOD DAYS Preoviposition 6 Oviposition to hatching 35 (eggs at 18°C.) Larval prefeeding period Ll Larva feeds 6 Premolting period 16 (larvae at 19°C.) Nymphal prefeeding period a Nymph feeds 6 Premolting period 20 (nymphs at 18%.) Adult prefeeding period 1 Adult (female) on host 6 Total 116 This period Nuttall believed to be the shortest time required for completion of the life cycle. From 4300 to 15500 eggs were laid by single females; the higher number probably approximates the more common figure in nature. Females may remain alive as long as 26 days after oviposition; males live much longer. The longevity of the various stages, presumably unfed, in these ex periments was 345 days for larvae, 89 days for nymphs, and over 421 days for some adults. Hosts were hedgehogs, guinea pigs, and rams. * Subsequently, Nuttall (1915) noted that some adults were still alive 759 days after emerging. Females that had fasted for 817 days were fed on a ram, mated with males that had fasted over 210 days, and were ovipositing when the report was written. Three times as many femles (253) as males (83) were counted in the progeny of a single female. Four years later, Nuttall (1919) observed that males may remain attached to one spot of the host for as long as 122 days. More commonly, however, after preliminary feeding for periods ranging from five to 29 days, they start wandering about in search of females. After mating, when females leave the host, males do - AZ, - little if any wandering. If a female is placed near a male, com siderable excitation is caused. A male may copulate with several females, but females apparently accept only a single male. Copu. lation may be very swift or may apparently extend over several days. Females do not move from their feeding site until they drop from the host. Ecology See also section on HOSTS above. In Algeria, adults are rarely seen in winter but begin to appear in March and continue till October, maximum densities being reached in April, May, and June (Algerian seasons are comparable to those of southern United States). Nymphs are found mostly during the summer. With slight variation, this seasonal picture is typical for H. marginatum wherever it oc- curs. In the Crimean forest (Melnikova 1953) adults are found during the summer, March to September, but rare individuals (?7mostly males) may be seen at any time of the year. Larvae and nymphs infest hares from the end of June to the first half of September. Nymphs are most common on hares in the latter part of August; for example, a hare on 2 August yielded 100 nymphs while another on 17 eee yielded 390 aoa In this forest reserve, H. marginatum (= H. p. plumbeum) occurs in all ecological zones and in all types of forest. in pure stands of conifers, it is, however, comparatively rare. The most favorable habitats appear to be valleys with small open fields between the mountains. In Transcaucasia, this species occurs equally in both high. land and forest zones and in desert and steppe formations and is found in every type of landscape in that area (Pomerantzev, Matikashvily, and Lototsky 1940). In Armenia, it occurs in the Artemesia semidesert of the Arax valley (Pomerantzev 1934) and at altitudes of 6500 feet and over (Lototsky and Popov 1934). On the Crimean steppes, adults are most common in July (May and June according to Kurchatov 1940A). There, the greatest aa AD ee density of this tick is around haystacks and in fallow fields where their immature stage host, the European hare, hides and feeds. mgorged nymphs drop from hares in autumn and overwinter in that stage. They molt in the spring, and adults attack cattle, sheep, horses, and man. The devastation of the Crimea during the war, followed by a great increase in hares and their ticks, was significant epidemiologically in the outbreak of highly virulent, often fatal hemorrhagic fever at that time. REMARKS Gynandromorphs of H. marginatum have been described and il- lustrated by Pervomaisky (ISe0y- The same author (1949) was un. able to secure a complete Fy generation from parthenogenetic fe. males of this species. Schulze (1932C) illustrated certain of the cuticular sense organs of two of his "subspecies" of this species, also "gyno- tropes", males with more dense punctations on the scutum con forming to those of the female scutum in location and distribu. tion. This species has been utilized in a study of sensory phy- siology (Totze 1933). When large numbers of ticks (as H. lumbeum) feed on a res— tricted area of the host, the females and sometimes also males fail to engorge completely and may die; their development is far from normal. When different species are competing for the same feeding area, this additional competitive factor often hinders their normal development (Pavlovsky, Pervomaisky, and Chagin 1954). DISEASE RELATIONS MAN; H. marginatum is considered to be the chief vector of the virus of eceanner eee fever. The extensive geographic range of this tick and its large population in many areas where it occurs ~ factors that suggest a high potential as a medically important species —~ appear to be generally unappreciated outside of Crimea. This species is not involved in the transmission of Omsk hemorrhagic fever, since it does not occur in that area, so far as is presently known. Ore Specimens naturally infected with Q fever (Coxiella burnetii) have been found; this species is a vector of the organism transmits it through all stages including the egz. Brucellosis or undulent fever organisms, Brucella melitensis, survive some time in this tick, which is said by some Soviet workers to be a carrier and transmitter of this pathogen. Some Russian studies of ticks as animals sustaining natural foci of tularemia have negated the importance of H. marginatum (as H. plumbeum) in this regard, although other workers have reported the finding of naturally infected specimens. CATTLE: Theileriasis (Theileria spp.). HORSES: Theileriasis (Theileria equi ) and piroplasmosis (Piroplasma caballi). Transovarial transmission of the latter organism to the seventh generation has been demonstrated. GUINEAPIGS: Rickettsiae pathogenic to these animals and transovarially transmitted to the F3 generation of H. marginatun have been reported. IDENTIFICATION Males: The combination of characters for recognizing typical males is: (1) the center of the subanal shields is directly pos terior of the central axis of the adanal shields (which are large and broad); (2) the lateral grooves are long, reaching approximate. ly to the eyes, but they are frequently somewhat obscured by dense punctations or by lack of discreteness, especially anteriorly; (3) the scutal punctations are dense and large in the distal and scapular fields, but variable elsewhere, being usually smaller and more shallow and less dense centrally; (4) the posteromedian groove reaches the scutal midlength, it is narrow anteriorly and wider posteriorly; the paramedian grooves are about half as long as the posteromedian groove and taper from a pointed apex to wide in the festoon area; a narrow heavily punctate ridge lies between the paramedian grooves and the lateral grooves. - 477 - The scutum is usually comparatively narrowly elongate, meas— uring approximately 4.0 m. long and 2.5 mm. wide. Its color is typically dark brown to black, but reddish specimens also occur; the legs may be entirely reddish or reddish centrally on each segment with paler anterior and posterior bands (see next para. graph). The scutal punctations may be dense enough to suggest H. rufipes, but those in the center are shallower and smaller than elsewhere, while in H. rufipes they are deep and quite uni_ form in size and depth. The parma may appear to be merely a median festoon and is the same color as the rest of the scutum. The subanal and adanal shields have rounded contours; the adanal shields are quite large. Heavily punctate males, as seen among series from Libya, Algeria, Egypt, Romania, and elsewhere may suggest H. turanicun (page 531). The legs of H. eS however, lack the con spicuous enamelling characteristic of those of H. turanicun. Females: The genital apron is a large, robust widely trans verse oval or triangle with a strongly bulging profile; it is most characteristic. The scutal punctation consists of numerous small, shallow, distant punctations over the entire surface (they may be almost medium size and slightly deeper and closer), and a few larger and deeper punctations scattered among them chiefly on the anterior half of the scutum. The basic punctation in some specimens is so shallow as to give scutum a rather smooth appear- ance, especially posteriorly. The central field between the cer— vical grooves is usually lighter (more reddish) than the dark lateral fields. The scutum appears to be exceptionally wide, its length.width ratio being about equal or shorter than wide. Females of H. marginatum and H. turanicum are quite similar but the bright enamelling of the leg segments of H. turanicum distinguishes them. In most specimens of H. turanicum the genital apron is not so widely triangular ‘and the scutal punc- tations are more numerous and discrete than in H. turanicunm. The larva and nymph of H. marginatum have been described and illustrated by Bernadskaia (193%) and Peldman-Muhsam (1948). SA Figures 182 and 183, co, dorsal and ventral views Figures 184 and 185, 9, dorsal and ventral views A, 9 genital area. B to D, © genital area outline and profile. B, unengorged. C, partly engorged. D, fully engorged. HY ALOMMA RUFIPES ee an Specimens PLATE LIII - 479 - HYALOMMA RUFIPES Koch, 1844. (Figures 182 to 185) THE HAIRY HY ALOMMA NOTE: H. rufipes is a most distinctive tick although Schulze, Delpy, and Feldman-Muhsam in their earlier reports confused H. rufipes under the name H. impressum. Nuttall during the first = of the 1900s identified African specimens as H. aegyptium impressum and many British and some Italian workers of this period Totlousd this precedent. The only other African species of wide range, H. truncatum, was referred to by this school as H. aegyptiun. A few Russian synonyms are listed in the distribution section ca but contemporary Soviet usage concerning the nomenclature of this tick appears to be incorrect. Ee N, <0. GS BQUATORIA PROVINCE RECORDS 7 i Lomas Syncerus caffer aequinoctialis Dec 3. Khor Waat Syncerus caffer aequinoctialis - (MCZ) (Allab) 2 7, Belorne domestic cattle Jan (2) 1 3 Torit domestic cattle Feb 3} 9» Dorit domestic cattle Nov ib 3B, Storer domestic cattle Dec 1 ouba: domestic cattle Jan 1 2 Terakeka domestic cattle Mar (SVS) 2 4 Tali Post domestic cattle Mar (SVS) 1 5 Meridi domestic cattle Jan (SVS) 1 1 Yambio domestic cattle Jan DISTRIBUTION IN THE SUDAN H. rufipes is widely spread in the Sudan but is numerous only in the semiarid central area. The following are localities from which specimens (all from cattle unless otherwise noted) have been seen; = Oe Bahr El Ghazal: Galual-Nyang Forest (Very common on buffalos and giraffes; SVS, HH. The absence of this tick on the numerous tiang examined in this area is notable). Aliab (buffalos; SVS). Lau (SGC). Lake Nyubor, Boro, Khor Shammam, and near Raga (SVS). Yirol (horses; Svs) Upper Nile: Bor, Ler, and Fangak (SVS). Malakal (HH). Blue Nile: Common at Wad Medani (HH). Abu Zor, Hosh, near Ethiopian border, and Lake Ras Amer (camels and cattle; SCC). Kordofan; “Western Jebels® and Umm Inderaba (SVS). Darfur: Nyala, Zalingei, Radom, Sibdo, Muhagariya, and Sahafa (common on camels, cattle, sheep, and horses; SVS). Kassala: Kassala (camels and cattle; SVS). Port Sudan (SVS). Khartoum; Khartoum; Numerous specimens have been collected from sheep, goats, and horses, but few from cattle (HH). At the Khartoum quarantine station many adults are found on Darfur and Kordofan cattle. The Sudan Government collection contains a series of adults reared from nymphs from a kite, Milvus grans, by H.H. King, 20 September 1922. Northern: Shendi (camels and donkey; SVS). Wadi Halfa (camels; SGC, HH). DISTRIBUTION H. rufipes is widely distributed in many drier parts of Afri- ca but it is quite localized and seldom very common in any locality. The hairy hyalomma is not known from many of the more westerly areas of Africa. It is present in the Yemen (Southwestern Arabia); and in North Africa occurs in Egypt and Libya but is not known fur- ther west on the Mediterranean littoral. Elsewhere, H. rufipes occurs in Palestine, Iraq, Eastern Ana. tolia, and Russia (Transcaucasia, Astrakhan, Kazakhstan). Soviet workers find H. rufipes in such small numbers and in such scattered = 4m localities that Pomerantzev (1950) believes its presence in Russia is due to small local populations established from nymphs from migrating birds. Schulze (1918) reported a Macedonian specimen (as H. impressum) that may have been imported on a bird. This distributional picture is indeed unique, and it may be surmised that H. rufipes is a species of the Ethiopian Faunal Re gion that has extended its range beyond these confines as a result of transportation by migrating birds, which are important hosts of immature staces (page 486). Note: All records below are for H. rufipes or H. r. rufipes; other combinations are so stated. NORTH AFRICA: BGYPI (Common on domestic animals in the Nile Valley only; also arrives at the Cairo abattoir on cattle from the Sudan and East Africa: Hoogstraal, ms. See immature HOSTS below). LIBYA (Scattered populations on Mediterranean littoral: Hoogstraal, ms.). WEST AFRICA: NIGERIA (As H. impressum rufipes: Unsworth 1949,1952. As H. impressum subsp.: Gambles 1951. As H. rufipes: Theiler 1956. Material from Katagum and Oban in BMNH collections; H.d. det.). FRENCH WEST AFRICA (As H. savi i impressa: Girard and Rousselot 1945. Rousselot 1946. ?As H. rufipes glabrum: Rousselot 1951 and Villiers 1955; it is assumed onm ec do not aie refer to H. turanicum. Rousselot 1953B. Theiler 1956). errr, CENTRAL AFRICA: FRENCH EQUATORIAL AFRICA (Rousselot 1953B). BELGIAN CONGO (Rare: Theiler and Robinson 1954. Theiler 1956). EAST AFRICA: SUDAN (Hoogstraal 1954B. Feldman-Muhsam 1954. Theiler 1956). ETHIOPIA (As H. aegyptium impressum f. typica: Stella 1939A, B,1940. As H. rufipes: Rsticn 56). ERITREA (As H. impressum impressum; Tonelli-Rondelli 1930A. As H. impressum rufipes: Niro 1935. Numerous specimens in BMNH and HA ieee FRENCH SOMALILAND (Hoogstraal 1953D). BRITISH SOMALILAND (As = oe H. grossum; Pocock 1900. As H. aegyptium impressum: Stella TSs8h 1359 . Numerous specimens from camels < EMNA collections; H.H. det.). ITALIAN SOMALILAND (As H. aegyptium impressum: Paoli 1916. Tonelli-Rondelli 1926A.— ne ooo Niro 1935. Stella 1938A,1940. As H. impressum rufipes: Tonelli-Rondelli 1935. Stella 1940. Specimens in HH collections. See NOTE under EAST AFRICA for H. impressum, p. 463). KENYA / Materials identified and variously reported by E. A. Lewis as H. aegyptium impressum, H. impressum albiparmatum, or H. rufipes are st invariably a mixture of H. truncatum and H. rufipes; sometimes H. impeltatum and H. albi tum are included under these names in Lowest collections now in the british Museum (Natural History) (H.H. det.). As H. rufipes: Binns (1951,1952). Theiler (1956). Note: In Sates @ probability of mixed species in reports by Lewis, the synonymy of the GARE names is uncertain. As H. wh Brassey—Edwards (1932). As H. Aaa Daubney H. impressum rufipes: Daubney eate (Wilson 1953. Theiler 1956. See HOSTS below). TANGA. NYIKA (As H. aegyptium impressum: Cornell 1936. As H. rufipes: Theiler 1956. ee HOSTS Seton). = SOUTHERN AFRICA: "SOUTH AFRICA" (Koch 1844. As H, impressum rufipes: Schulze and Schlottke 1930). ANGOLA: Absent: Sousa Dias (1950). Theiler (1956) .7 NORTHERN Oe S (Matthysse SON, Theiler and Robinson 1954. Theiler 1956). SOUTHERN RHODESIA (As H. ae ium impressum: Jack 1921,1928,1937,1942. As H. rufipes: wel Ter OF NYASALAND (No available Secords) — KOEERBIQUE (As H. impressum rufipes: Theiler 1943B. Santos Dias 1952D ,1953a, 1950 TondeTeo 1955. As H. rufipes: Theiler 1956). BHCHUANALAND (Specimens from Chena Te EMH collections; H.H. det. Theiler 1956). SWAZILAND (Uncommon: Theiler 1956). SOUTHWEST AFRICA (As H. impressum rufipes: Schulze 1936A. As H. impressum: Schulze SLO. As H. eee Theiler 1956. See WOSTS below). UNION OF SOUTH AFRICA (The H. oop of Donitz 1910B and of Cooley 1934 apparently includes H. truncatum and = 403 = H. rufipes. As H. aegyptium impressum: Howard 1908. Bedford 19328, »1936. exander, Mason, and Jeitz 1939. McIntyre 1939. du Toit 1942,1947. du Toit and Monnig 1942. As H. aegyptium: Clark 1933. As H. impressum rufipes: Theiler 1943B. As [ rufipes: Thorburn 1952. Werte T1952. Fisiler 1956). ISLAND GROUPS: MADAGASCAR (Hoogstraal 1953E. Theiler 1956). COMORES (Kratz 1920; cf. immature HOSTS below). NEAR EAST: TURKEY (Rare in eastern Anatolia: Hoogstraal, ms.). PALESTINE (As H. impressum, rare: Bodenheimer 1937. Adler and caer gee iste DOISy. YEMEN and IRAQ (Common: Hoogstraal, Msc Je RUSSIA: (As H. aequipunctatum: Olenev 1931A,C. Galuzo and Bespalov 1035. As ES impressum; Pomerantzev, Matikashvily, and Lototsky 1940. As H. marginatum impressum: Pomerantzev 1946. Tselishcheva 1953. “As H. plumbeum impressum: Pomerantzev 1950). NOTE; The record of H. impressum rufipes from China (Chod_ ziesner 1924) probably refers to a heavily punctate H. marginatum (Kratz 1940). H. rufipes has been stated to occur in Portugal by Kaplan and Hulse in their review of prevalence of Q fever in Europe; this apparent error derives from the report by Fonesca, aos Colacao, Oliveira, Branco, da Gama, Franco, and Lacerda 1951) that "H. rufipes glabrum™ is associated with Q fever there. This is assumed To nae to H. marginatum. HOSTS Domestic cattle appear to be the most common hosts of this tick. They are mentioned by practically every author and are the most frequent hosts of specimens in museum collections. Other common domestic hosts are horses, sheep, and goats. Bedford (1932B) states that dogs and cats are also infested; specimens from these hosts are present in British Museum (Natural History) and HH collections. In semidesert areas, camels are frequently parasitized (Sudan records; HH mss.; numerous Somaliland spec- imens in BMNH collections; Hoogstraal 1953D). Among wild ani. mals, the buffalo and giraffe are common hosts, as is the rhinoceros. Antelopes and certain birds are less common hosts = hb of adults, and a variety of small mammals are occasionally ine fested. Immature stages feed on a large variety of birds and also on hares. Adults Domestic animals: See two paragraphs above. Man: Howard (1908). Bedford (1932B). J. B. Walker (cor_ respondence; 9 tick from Tanganyika). Wild animals: Rhinoceros (Two collections in BMNH from Kenya). Buffalo (Santos Dias 1952D,1953B. Onderstepoort col. lection from Northern Rhodesia. EMNH collection from Kenya, Sudan records above). Eland (Schulze 1936A. Two collections in BMNH from Southwest Africa. HH collection from Tanganyika. Onderstepoort collection from South Africa and Tanganyika). Bushbuck (MCZ collection from Tanganyika). Duiker (Sylvicapra immi) (Bedford 1932B). Sable antelope (Santos Dias z Gemsbok (Onderstepoort collection from Southwest Africa). Giraffe (Santos Dias 1952D,1953B. Onderstepoort collection from northern Kenya and Southwest Africa. Sudan records above). Jackal (Canis mesomelas schmidti) (Stella 1939B). Zebra (Santos Dias 1952D. EMNH collection from Tanganyika). Hare (Howard 1908. Onderstepoort collection from South Africa). “Fowls™ (Howard 1908). Ostrich (Howard 1908. Bedford 1932B. ¢@ in HH collection, from “west of Afmadu", Somalia, 1952, Col. D. Davis legit). Guinea fowl (Santos Dias 1953B). Adults from the following birds are present in the Onderste- poort collection (Theiler, Pie cian ie AEE ostrich (Southwest Africa), swallow (Southern Rhodesia), Cape dikkop (Burhinops c. capensis from South Africa), and mocking chat (Thammolaea c. Cinnamomeiventris from South Africa). Immature Stages Hosts of the immature stages noted by Bedford (1932B,1936) are not listed here since it is questionable that larvae and nymphs - 485 - of H. rufipes and of H. truncatum (= H. transiens) could be dif- EGGS at that time. Nymphs have been reported from a hare (Alexander, Mason, and Neitz 1939), and from a kite (Sudan records above); the former workers induced five of the twelve nymphs to reattach to a guinea pig. Kratz (1940) records the finding of a nymph, which molted into a male mo pes: on a female comorant caught on the high seas between the northern trip of Madagascar and the Comores Archipelago. The Onderstepoort collection (Theiler, correspondence) has larvae (L) and/or nymphs (N) from the following South African birds: Namaqua thrush, Afrocichla smithi Cape thrush, Afrocichla 0. Olivacea (2 collections) White-throated seed_eater, Crithagra a. albogularis t collections) Mocking chat, Thamnolaea c. cinnamomelventris Red-winged starling, drus m. morio Starling (Southwest mS Boubou shrike, Lanjarius f. ferrugineus Gray tit, Parus afer ~ Fiscal flycatcher, Eien silens Cape barn owl, Tyto a affinis The same collection contains nymphs from a hare and a rock hare (Pronolagus randensis) in South Africa and from a hare in Uganda. i tt ad hai 5 el | Sec In Egypt, nymphs (reared to adults in the laboratory) have been found only on birds (Hoogstraal, ms.) although adults are locally common on domestic animals. The hosts have been: Wheatear (European form), Oenanthe o. oenanthe Blackeared wheatear (Eastern form), Oenanthe hi spanica melanoleuca The former bird breeds throughout most of Europe east to Central and northern Asia and to northern Alaska; it winters in Arabia and tropical Africa, also in Asia to India. The latter “breeds in the Crimea, Bulgaria, and almost throughout the Balkan peninsula, Asia Minor, Palestine, and western Persia, etc.; winters in Egypt and Sinai to the Sudan, Ethiopia, the Red Sea coast, and has straggled = 4BOve to the southern Sahara, British Islands, Malta, and northwest Africa® (Meinertzagen 1930). Possibilities for the wide dis_ persal of this tick are easily recognized. BIOLOGY Life Cycle Under laboratory conditions, H. rufipes is a two host tick although it possibly may also ia Re Sra three host type of life cycle. Theiler (1943B and 1955 correspondence) has summarized the developmental stages as follows: PERIOD DAYS (1943B) (1955) Preoviposition 460, V2 Zato, 9 Oviposition period 37 to 59 Oviposition to hatching 34, to 66 28 to 66 Larval prefeeding period ? Larva feeds Servo / Premolting period Zz to 15 Nymphal prefeeding period Y Nymph feeds 7 to 10 Premolting period 14 to 95 (Larvae and nymphs on host) 13 to 45 average 14 Adult prefeeding period ? Adult (female) feeds 5 to 6 5 to 12 It appears that the minimum time for completing the life cycle is between four and five months but double this period may be re— quired under local conditions. "The life cycle of H. aegyptium (= ?H. rufipes, possibly mixed with H. truncatum: HH) is of particular value in that it illus— trates the influence of vermin in the distribution of the species. On sheep, cattle, and domestic fowl it behaves as a three host tick, requiring a separate host for the larval, nymphal, and adult stages. On the hare, H. sem peae will feed as a larva, become engorged, molt as a nymph without leaving the host, feed as a = AO) ae nymph on the same individual host, and then drop off the host for molting. Thus on the hare the life cycle requires only two hosts" (Brassey-Edwards 1932). This interesting phenomenon should be re— investigated. The long oviposition period is especially noteworthy. Unfed larvae may survive a year, unfed nymphs three months, and unfed adults for longer than a year (Theiler 1943B). Enigk (1953) ob served unfed adults surviving up to two years. Howard (1908) considered H. rufipes as a two-host tick with one generation a year in South Africa. He described, illustrated, and discussed the immature stages but did not differentiate them from those of H. truncatum which he apparently did not rear. Jack (1928) noted a twochost and a three-host type of life cycle for this tick. Ecology Thorburn (1952) states that on cattle the chief site of in- festation of this tick is in the tail region. Specimens in the present collection are from the flanks, genitalia, udders, and perianal regions. The anal area is mentioned by Matthysse (1954). Nymphs are always, in our experience, on the crown of the head of their avian hosts. du Toit and Monnig (1942) record the finding of a male at— tached to the hard palate of the mouth of a cow, and indicated that on the farm where this occurred this phenomenon had been ob. served on several occasions. H. rufipes ranges through the more arid areas of tropical and southern Africa but only localized populations maintain themselves in the severely arid conditions of northern Africa. It exists where annual rainfall is from ten to thirty inches a year. It may also thrive in irrigated areas with diminished rainfall or where a long, severe dry season occurs between an annual rainy season of approximately forty inches. In the Sudan, it is more common in the drier savannah and semiarid central areas than in the southern forest and savannah areas; it occurs in the Nile Valley, but is not known in extreme desert conditions. In Egypt, = 4855 H. rufipes is found only in the Nile Valley, never in extreme desert areas. The hairy hyalomma is included in Wilson's (1953) Amblyomma emma — R. pravus (= R. neavi) association (see page J; which is found in areas where rainfall rarely exceeds 25 inches annually. The only ecological survey of this tick is that of Theiler (1956) who lists the areas of its occurrence and absence in south. ern Africa. It occurs in all desert and semidesert areas with rainfall up to thirty inches annually, but at higher altitudes or in semitropical conditions, where the relative humidity is higher, it is absent even though annual rainfall is little or no greater than in some of the hotter, drier areas. It does not occur in winter rainfall areas, where rain falls throughout the year, or in coastal areas with high relative humidity as a result of proximity to the sea. Temperature appears to be a limiting factor of lesser importance since H. rufipes ranges from hot deserts into areas with up to 120 days of frost annual. ly. Increase in temperature associated with increase in relative humidity restricts the tick's range. Other factors being equal, the hairy hyalomma occurs in most vegetational types except for ested areas of central Africa. In many regions it is active the year around, but in others more so in summer than in winter. In Russia, H. rufipes has been reported (as H. impressum) from the western deserts of Transcaucasia (Pomerantzev, Matikash. vily, and Lototsky 1940) and, in western Tadzhikistan, from mountain pastures but not in the valleys (as H. aequi punctatum) (Galuzo and Bespalov 1935). - DISEASE RELATIONS MAN: Nymphs infected with boutonneuse fever (Rickettsia conorii) have been taken from a hare in South Africa. CATTLE: H. rufipes causes abscesses and sloughing of the host skin. These areas often serve as points of penetration of the screwworm miso bezziana Villem. This tick may also be associated with footrot of sheep, a secondary infection by = 409= bacteria, and lesions from its bites may also lead to lameness in sheep. The possibility that the hairy tick may be a vector of sweating sickness (virus) of cattle has been mentioned. IDENTIFICATION Male: H. rufipes, almost without exception, is a large, robust, shiny black tick with a comparatively broad body outline. Its scutum is densely and rather uniformly punctate, so much so that the lateral grooves are obscured. The punctations are close together and while they give the impression of being rather uni_ form, they usually grade imperceptibly from large posteriorly to smaller anteriorly. There is little differentiation of the caudal area. Ventrally, the subanal shields are directly pos— terior of the adanals and the circumspiracular area is more or less hirsute, but a fair to dense accumlation of hairs always occursin this area. The ventral integument is usually somewhat more hirsute than in other species. The legs are reddish brown with bright, paler rings. The posterior body margin is typically broadly rounded but not infrequently may be somewhat constricted and thus tend to approach the form of that in H. impressum. New ly molted specimens are reddish brown and rare preserved individ. uals show this color. The body shape of some specimens is more narrowly elongate than is typical for this species, but such individuals always appear to be rather weak and poorly nourished. Females: This sex often reaches considerable size. Cir cumspiracular pilosity and color are like those of the male; scutal punctations are like those of the anterior part of the male scutum. The genital apron is a wide shield that bulges strongly from a deep anterior and posterior indentation. The outline of the apron is much like that of H. marginatum and H. impressum; however the division of the apron of latter spe cies into an anterior ridge and a posterior button readily separates this from H. rufipes. The larva has been sketched by Bedford (1934) and Theiler (19433). = eh JO Figures 186 and 187, @, dorsal and ventral views Figures 188 and 189, 9, dorsal and ventral views A, 9, genital area. BtoD,Q, genital area outline and profile. B, unengorged. C, partly engorged. D, fully engorged. HYALOMMA TRUNC ATUM an Specimens PLATE LIV - 491 - HYALOMMA TRUNCATUM Koch, 1844. (Figures 186 to 189) THE AFRICAN HYALOMMA NOTE: The Nuttall school referred to H. truncatum as H. oeepl During the past five years, most authors nave called Ss species H. transiens, the authority for which has been at— tributed to Schulze (1019) or to Delpy (1946A). Feldman_-Muhsam's (1954) studies of Koch's type specimens leave no doubt that this species is Koch's (1844) H. truncatum. Feldman-Muhsam (op. cit.) has also compared the type specimens of several of Schulze's African "species™ and found them to be identical with H. truncatunm. These are noted below in the distribution section. Lea Boa EQUATORIA PROVINCE RHCORDS aL Lugurren Phacochoerus aethiopicus bufo i) soratt Sus scrofa sennaarensis LS eKajorKaji cerus caffer aequinoctialis 1 1 Kapoeta omestic cattle ys fort domestic cattle Se eee LOTASG domestic cattle iL Lalanga domestic cattle A —-daleigbl domestic cattle 12 4 # Tombe domestic cattle 4 8 Kapoeta domestic cattle 2 lLoronyo domestic goats 2 lori domestic goats Tortoise 1 LS Juba Kinixys belliana DISTRIBUTION IN THE SUDAN The following material has been studied: = 492% Dec (SVS) (svs) (Sis) (Svs) Bahr El Ghazal: Galual-Nyang Forest (Large numbers of adults from several giraffes in March, May, and June; SVS. Moderate num. bers of adults from forest buffalos from February through April; SVS, HH. Small numbers of adults from tiang, roan antelope, domestic horse, and on ground from February to April; SVS, HH). Fanjak (Small numbers of adults from cattle, February and March; SVS, HH). Wau (roan antelope; SGC). Yirol (cattle; SVS). Note: The following records consist of one to ten adult specimens per collection unless otherwise mentioned. Upper Nile: Makier (cattle; SVS). Blue Nile: Lake Ras Amer (camel; SGC). Hassa Heissa (camel; G. Kohls det., G. B. Thompson, correspondence). Wad Medani (cattle and camels; HH). Darfur: Nyala (camel; SVS). Muhagariya (horses, donkeys, cattle, and camels; SVS). Zalingei (camels, cattle, horses, donkeys, and goats; SVS). Kulme (no host record; BMNH). Radom (cattle; SVS). Kordofan: Talodi and Heiban (cattle; SVS). J Khartoum: Apparently not established in this Province but arrives 1n fairly large numbers on Kordofan and Darfur cattle for export to foreign markets; HH.7 Kassala: Kassala (goats; SVS). See EGYPT below. Northern; Rare in this Province but arrives at the Wadi Halfa Quarantine on cattle en route to Egypt. Known only from report by Chodziesner (1924) from Delgo. DISTRIBUTION H. truncatum is the sole endemic representative of this genus that is widely spread throughout the Ethiopian Faunal Region (Figure 1) and nowhere else. It commonly occurs in the drier parts of this Region but appears to increase in numbers - 493 - towards and north of the equator. It is rare or absent in forests of western Africa. ATLANTIC OCEAN: CANARY ISLANDS (Specimens from dogs, Tene- rife, 1900, 1n BMNH collections; HH det.). NORTH AFRICA: BGYPT (Occurs only in Gebel Elba area of ex. treme Southeastern Egypt adjacent to Sudan frontier (Kassala Prov. ince); numerous males arrive at the Cairo abattoir on cattle from the Sudan and East Africa but this species has not established itself as a result of these introductions: Hoogstraal, ms.). WEST AFRICA: FRENCH WEST AFRICA (As H. truncatum sp. nov.; Koch T82Z. As ft. tium impressum transiens: ochulze 1919. As H. transiens: Rousselot ; ). GAMBIA (Numerous spec. imens in single lot from cattle in BMNH collections; HH det.). PORTUGESE GUINEA (As H. savignyi: Tendeiro 1948,1949A, 1951F,1952A,C,D. From Tendeiro'ts TOLOA discussion it is evident that he is referring to H. truncatum (= H. transiens) but prefers to call it H. savignyi. “It is likely that one to three other Hyalomma species occur in Portugese Guinea). NIGERIA (As H. aegyptium; Simpson 1912A,B. As H. impressum transiens: Unsworth Asp As H. impressum subsp.: Zanibios TST. Is W. transiens: Unsworth 1952. See HOSTS below). GOLD COAST (As H. aegyptium: Simpson 1914). TOGO (Feldman-Muhsam 1954 states aT H. impressum brunneiparmatum Schulze and Schlottke, 1930, is a synonym of H. truncatum; however, from examination of Miss J. B. Walker's Kenya-reared material of H. albiparmatun, in which the parma varies in size and color, it Is evident that H. brunneiparmatum is a synonym of H. albiparmatum). CENTRAL AFRICA: CAMEROONS (As H. aegyptium impressum transiens; Chodziesner 1024. As H. transiens: Rageau TSBL, TORS. Rousselot 1951,1953B. Unsworth 1952). FRENCH EQUATORIAL AFRICA (As H. nitidum from "New Cameroons®; Schulze 1919. Chodziesner 1924. As H. impressum nitidum: Schulze and Schlottke 1930. Kratz See also 1940. Feldman-Muhsam 1954. As H. eon impressum transiens: Chodziesner 1924. As H. transiens: ousselot 1O51, 195 He BELGIAN CONGO (As H. transiens: Theiler and Robinson 1954 ° - 494 - EAST AFRICA: “EAST AFRICA" (As H. planum and H. zambesianun: Schulze and Schlottke 1930). SUDAN (As H. aegyptium eee ressum eae Chodziesner 1924. Kratz 1940. As i. im Ssain Tetetooe eLpes: ochulze and Schlottke 1930. As H. transiens: Hoogstraal IRIE ETHIOPIA (As H. impressum transiens and H. impressum nitida: Stella 1939B,1940. As f tium impressum transiens: Chodzies. ner 1924). ERITREA (As H. impressum transiens: Tonelli-Rondelli 1930A,1932C. Niro 1935.— Sete TOSSES . Common in many parts of Eritrea: HH). FRENCH SOMALILAND (As H. transiens: Hoogstraal 1953D). BRITISH SOMALILAND (Numerous specimens from camels in PMNH collections; HH det.). ITALIAN SOMALILAND (As H. aegyptiun impressum: Paoli 1916. As H. aegyptium impressum form transiens: Tonelli-nondelli 1926A,1935. ella O. See HOSTS below). KENYA / As H. impressum transiens: Daubney (1937). As H. transiens: Binns eae 1952). As H. impressum near planum: ~ Fotheringham and Lewis (1937). As H. truncatum: Feldman-Muhsam (1954). Hoogstraal (1954C). See also H. albiparmatum, p. See HOSTS below. ve H. lewisi Schulze (1936E) is a synonym of H. truncatum and not of H. excavatum as stated by Delpy (1949E); it 1s also not a "“Hyalommina® as stated by Schulze (19365). Schul ze_identified material of "H. lewisi” consists of small, stunted, misshapen i. truncatum (seen by HH); in this Feldman-Muhsam (1954) is in agree. ment. Kratz (1940) also referred to H. lewisi from Kenya. See page Lewis (see bibliography) mentions Hyalomma ticks under a variety of names. Most specimens in his Tas Sree aes now in British Museum (Natural History) are H. truncatum among which H. rufipes is frequently mixed and other species aré sometimes eae The H. ee specimens had been identified by Lewis as H. im ressum, H. dromedarii, and iH. eer this confusion is unders andable due to the unsatisfactory informa. tion available in literature at that time./ UGANDA (As H. impressum transiens: Wilson 1950C. As H. transiens: Wilson oOmmon in many Uganda collections ‘studied by Theiler and ey HH). - 4% - TANGANYIKA / As H. planum and H. aegyptium albiparmatum: Schulze (1919). i! H. aegyptium impressum transiens: Chodzies. ner (1924). As H. impressum transiens and as H. Lewisi; Schulze (1936E). Kratz UiSIOy- See KENYA above. As H. aegyptiun: Cornell 1936. ?As H. impressum planum f. rhinscemsee Schulze and Schlottke (1930) ee CsLoy; the synonymy of this name is uncertain but it is suspected to apply to H. truncatum. See HOSTS below.7 T uA? eRe SOUTHERN AFRICA: ANGOLA (As H. impressum transiens: Sousa Dias 1950. Santos Dias 19500. As H. transiens: Theiler and Robinson 1954. Santos Dias 19500 noted that 7H. savignyi™ had been reported from Angola by A. Morais in 1909, ai this may refer to H. truncatum (= H. impressum transiens), MOZAMBIQUE (As H. impressum transiens: Theiler 19Z3B. Santos Dias 1947B 1953B,H, . Tendeiro 1955. As H. truncatum: Theiler 1956). NORTHERN RHODESIA (As H. transiens: Theiler and Robinson 1954. Matthysse 1954. See HOSTS below. SOUTHERN RHODESIA (As H. aegyptium impressum transiens: Chodziesner 1924. As H. ae ae Lawrence 1959. As i. a. aegyptium: Jack 1942. So . truncatum: Theiler 1956. See Hoste tol elow). NYASALAND (As H. impressum transiens: Wilson 1943 ,1946. As iH. impressum; TOEOEy Wilson SOUTHWEST AFRICA (As H. ae ium impressum transiens: Chodziesner 1924. As H. impressum transiens: Kratz LO40. As H. aeryptium aegyptiun: BodTond LOS2E- As fH. transiens: Fiedler ose ee HO elow). SWAZILAND (As H. a. aegyptium: Bedford 1932B. As H. truncatum: Theiler 1956).~ BECHURTILEND (As H. aeryptium impressum transiens: Chodziesner 1924. As H. trun SS um: iatier To er 1956). /BASUTOLAND: Absent; Theiler (1956)._7 UNION OF SOUTH AFRICA (The "H. aegyptium™ of Donitz 1910B and of Cooley 1934 apparently include ee H. rufipes and H. truncatum. As H. a. aegyptium: Lounsbury 190/C. = Theiler T505B,1906. Howard 10 edford 1920,1926,1927,1932B,1936. Nieschulz and du Toit 1937. P. J. du Toit 1931. Finlayson, Grobler, and Smithers 1940. R. du Toit 1942A,B,1947A. As H. impressum impressum; Theiler 1943B. As H. transiens: Erasmus 5 orburn 1952. Neitz 1954. As H. truncatum: Feldman. - 496 - Muhsam 1954. Theiler 1956. As H. aegypticum (sic): Gear 1954. See HOSTS below). "SHORES OF THE ZAMBESI" (As H. zambesiacum: Schulze and Schlottke 1930. Kratz 1940). dal ARABIA: YEMEN (Hoogstraal, ms.). OUTLYING ISLANDS: MADAGASCAR (Recently introduced: Hoog- straal 1953E. Theiler 1956). SEYCHELLES (Desai 1941; not stated whether introduced or established). ZANZIBAR (As H. aegyptium: Aders 1917). HOSTS Domestic cattle and goats are the most common hosts of H. truncatum but other large wild or domestic mammals may be in- fested. Wild carnivores are seldom recorded as hosts. Rarely, small mammals, birds, or tortoises are also attacked. Immature stages are definitely known from birds and hares but most published remarks concerning these stages should be accepted with reservation because of questionable identity. Adults Domestic animals: Cattle (Bedford 1932B, Schulze 1936, Fotheringham and Lewis 1937, Sousa Dias 1950, Wilson 1943,1946, 1950B, Rousselot 1951, Rageau 1951,1953, Santos Dias 1953B. Sudan records above. Numerous specimens in various collections examined for the present study). Goats (Bedford 1932B, Rousse lot 1951, Hoogstraal 1953D,E. Numerous BMNH specimens. Sudan records above). Sheep (Bedford 1932B, Wilson 1950B, Sousa Dias 1950, Rousselot 1951, Hoogstraal 1953D. EMNH specimens. Sudan records above). Camels (Aders 1917, Rousselot 1951. Numerous BMNH specimens from British Somaliland. Hoogstraal, Yemen ms. Sudan records above). Horses (Bedford 1932B, Sousa Dias 1950, Rageau 1953. Sudan records above). Donkeys (Bedford 1932B, Rousselot 1951). Mules, dogs, and rarely cats (Bedford 1932B). Dog (HMNH specimens from Canary Islands, Eritrea, and Transvaal). - 497 - Wild antelopes: Tiang (Sudan records above). Roan antelo (Bedford 19528, Fesley 1934, Tonelli-Rondelli 1930A. BMNH a imens from Nigeria. Sudan records above). “Ozanna grandicornis' (Santos Dias 1952D). Wildebeest (Matthysse 195Z. Se Se specimens from South Africa). Nyasa wildebeest (J. B. Walker specimens from Tanganyika). Lichtenstein's hartebeest (Santos Dias 1952D,1953B). Brindled gnu or blue hartebeest (Bedford 1932B, Santos Dias 1953B). Sassaby or bastard hartebeest (BMNH specimens from South Africa). Eland (Chodziesner 1924, Bedford 1932B, Wilson 1950B, Santos Dias 1953B. BMNH specimens from Southern Rhodesia and Southwest Africa. Onderstepoort specimens from South Africa). Greater kudu (Santos Dias 1953B). Bushbuck (MCZ specimens from South Africa). Western defassa waterbuck egee 1953). Gemsbok (Onderstepoort specimens from South Afri ca). Other wild animals: Hedgehog (Bedford 1936). Hares (BMNH specimens from Kenya and Nigeria). Bushpig (Santos Dias 1953B. HH specimens from Eritrea. Sudan records above). Warthog (Santos Dias 1953B, Bedford 1932B, Rageau 1953. Numerous BMNH specimens from Kenya and Nigeria. Sudan records above). White or square. lipped rhinoceros, southern race (MCZ specimens). Black or narrow lipped rhinoceros (Schulze 1919, Schulze and Schlottke 1930. BMNH specimens from Kenya). Buffalo (Schulze 1919, Wilson 1950B, Santos Dias 1952D,H,1953B. MCZ and BMNH specimens from Kenya. J. B. Walker specimens from Tanganyika. Sudan records above). Dwarf buffalo (Rageau 1953). Giraffe (Chodziesner 1924. MCZ specimens from Kenya. BMNH specimens from Tanganyika. Onder- stepoort specimens from Transvaal and Southwest Africa. Numerous Sudan specimens recorded above). Burchell's zebra (Santos Dias 1952D ,1953B). Zebra (Matthysse 1954. BMNH specimens from Kenya. Onderstepoort specimens from Northern Rhodesia). Lion and antbear (Wilson 1950B). Leopard (BMNH specimens from Kenya. Onderstepoort specimens from Southern Rhodesia). Jackal and African porcupine (Matthysse 1954. Onderstepoort specimens from Northern Rhodesia). Reptiles: Tortoise (Sudan records above). Birds: Cape thick. kmee, Burhinops capensis (Bedford 1932B). Ostrich (Ic and in HH collection, ven west of Afmadu", Somalia, 1952, Col. D. Davis legit. Ostriches in Southwest Africa (Theiler, ee = 498 = Man: Several specimens from Kataguna and Katagum, Nigeria, and from Kenya in BMNH collections (HH det.). Immature Stages Nymphs on dogs and hedgehogs (Rousselot 1951). Larvae and nymphs sometimes on cattle, sheep, and goats (Fotheringham and Lewis 1937). Nymphs on hares (Wilson 1946,1950B, Sousa Dias 1950. Fiedler 1953). Larvae from a hornbill, Tokus flavirostris leucomelas (Santos Dias 1952D). "“Immatures"™ from a pied crow, orvus albus albus in Transvaal (Theiler, correspondence). BIOLOGY Life Cycle Unstudied. Wilson (1946) was unable to rear this species in Nyasaland. Ecology The African hyalomma, another xerophilic member of this genus, obviously differs somewhat from H. rufipes in ecological requirements but the limiting factors are not yet recognized clearly enough for proper elucidation. As stated above, the distribution of H. truncatum is strictly limited to the Ethio- pian Faunal Region and its range is widespread and fairly continuous within these confines except in heavily forested and high rainfall areas. Wilson (1953) includes H. truncatum in the A. gemma — R. pravus (= R. neavi) association (see page ) that occurs where annual rainfall seldom exceeds 25 inches. In southern Africa (Theiler 1956), the range of H. trunca tum differs from that of H. rufipes in that the former is absent ‘at higher elevations with high rainfall but present in cooler lowland winter rainfall areas. In regions with 25 inches of annual rainfall populations are rare and isolated. In Equator jal Africa, however, the African hyalomma does tolerate this and a slightly higher range of rainfall (HH). Here a combina. tion of factors including higher temperatures, long dry seasons, - 499 - and lower average relative humidity probably modify this tick's ecological thresholds (HH). In southern Africa, low temperature and high altitude do not in themselves limit the range of H. truncatum. It occurs in all types of South African vegetation except in short grass of the highveld, a mountainous zone associated with high rain. fall, and is rare or absent where snow falls. From the size and variety of collections examined, it ap. pears that in comparison with H. rufipes, H. truncatum may be somewhat less numerous and more widely ranging in southern Africa but that the reverse is true towards and beyond the equator. This matter, however, requires more careful study. H. truncatum is rare or never present in the forests of western rica. This species was not collected in high rainfall areas of the Cameroons (Unsworth 1952), and is unusual if not entirely absent on the humid west bank of Equatoria Province in the Sudan. In Northern Province of Nyasaland, where H. truncatum is the only species of this genus that is found, females engorge on cattle chiefly during the dry season (March, April, May) but also in small numbers during other months of the dry sea son. Nymphs were found on hares early in the rainy season (October) and also in December (Wilson 1946). Adults attach in the brush of hair at the tip of the tail, between the hooves, in the inguinal and perianal areas, and on the scrotum and udders. A hymenopteran parasite, Hunterellus theilerae, has re. cently been described from nymphs of H. truncatum of Southwest Africa and from nymphs of Rhipice halus oculatus from Transvaal (Fiedler 1953). Cooley (33D Teared Hunterellus hookeri from nymphs of "H. aegyptium” in South Africa, but it appears that he included both H. truncatum and H. rufipes under this name. = 500 = DISEASE RELATIONS as Tick paralysis (toxin or venom). Q fever (Coxiella burnetii). CATTLE: Sweating sickness (virus). Lameness and paralysis in calves (toxin or venom). SHEEP; This tick may be associated with footrot of sheep, a secondary infection by bacteria, and lesions from its bites may lead to lameness. HORSES: Not a vector of horsesickness (virus). NOTE: This species should be considered as strongly suspect in the transmission of rickettsial organisms among domestic and wild animals. REMARKS A somewhat deformed specimen of H. truncatum has been des cribed and sketched by Santos Dias (1947B) and another (as H. savignyi) by Tendeiro (1951F). In various papers by E. A. Lewis on work in Kenya, based largely on H. truncatum (cf. Hoogstraal 1954C), the author refers to rearing experiments by Nuttall (1913) and Patton and Cragg (1913) as being accomplished with the same species. Since Nuttall's material originated in Algeria and Patton and Cragg'ts in India, these workers obviously utilized different species. The material used by Nuttall, now in British Museum (Natural History) collections, reported as H. aegyptium, is H. margina. tum. The 4500 specimens of H. truncatum from throughout Africa that have been examined for the present study are highly distinctive and show considerably less variation than en countered among most other species in this genus. This ob servation is diametrically opposed to Feldman-Muhsamts (1954) remarks; “Examination of laboratory-bred material ......... = Ole showed an enormous range of variation between the offspring of one female“. As much caution must be employed in evaluating laboratory. reared specimens as in evaluating field-collected material. Under abnormal, artificial conditions, some individuals that would not survive in the field may be protected enough to maintain the life they would otherwise lose under inclement conditions. Artificial conditions in themselves obviously induce morphological modifica tions. At any rate, extremely few atypical specimens are found in field collections. IDENTIFICATION Males: The scutum is black or reddish-black and measures approximately 3.3 mm. long and 2.3 m. wide. It is characterized by long, deep, distinct, cleanly cut lateral grooves; smooth, glossy, impunctate surface except caudally, where there is a dense patch of large, contiguous punctations. The scutum is narrowed posterior of the spiracular plate, but the posterior margin is usually not so squarely truncate as in H. impressum. The festoons number seven and the central one is not morpholog- ically differentiated, as compared with H. albiparmatum in which the central festoon forms a parma resembling a ne nse: cellu loid watch cover, variable in size, shape, and color. Ventrally, the small rectangular subanal shields lie posterior of the axis of the larger, rectangular adanal shields. The legs are red dish brown with bright paler rings. Variable field collected males may be small and stunted and lack the subanal shields. Such specimens are the basis of Schulze's so called “H. lewisi” in the “subgenus Hyalommina” (page 521). Collections froma few areas show somewhat more than ordinary scutal punctation. The long, clear, cleanly cut lateral grooves indicate that such specimens are H. truncatum and not lightly punctate H. impressum, as does also The com. paratively more rounded posterior margin of the body. Although H. marginatum occurs with H. truncatum only in rare localities at the northern periphery of the range of the latter, it may be well to add that the scutum of H. truncatum is characterized by fewer scapular and central punctations; = 90 = longer and cleaner lateral grooves; glossier surface; often some what smaller size; obsolescence of posteromedian and paramedian grooves in the caudal field of dense punctations; and narrowed posterior margin. Field collected specimens of these two spe cies that might be confused have not been observed during the present study; Feldman-Muhsam (1954), however, states that laboratory-bred series might be confused. Females are easily recognized if only by the genital apron, the character of which is accentuated rather than decreased by enormous engorgement. The apron is a transversely elongate oval of somewhat variable shape; in profile it is surmounted anterior— ly by a narrow, bulging lip; centrally it is deeply depressed (or concave); posteriorly it is bounded by a mre or less dis. tinct lip that never protrudes as mich as the anterior lip. Feldman-Muhsam's (1954) figure 2F of this apron is a surprising- ly unsatisfactory representation of its actual appearance and supports the assumption that this species did not prosper during the laboratory study devoted to it. The scutum of practically every field collected female is blackish and with few punctations among which a few fine ones may be scattered. The scutum of a few specimens bears larger, superficial punctations scattered about its surface; that of greatly engorged specimens, as usual in this genus, is rugose. In the few specimens with a more punctate scutum, the genital apron is nevertheless highly distinctive and the glossy scutal appearance is retained. Note that no known characters distinguish the females of H. truncatum from those of the less common and more restricted B. albiparmatun. HEE NON-SUDANESE SPECIES OF HYALOMMA (Figures 190 to 211) In order to better understand this group, illustrations and selected data for all species presently—recognized in the genus Hyalomma have been assembled in this section. KEY TO ALL PRESENTLY RHCOGNIZED SPECIES OF THE GENUS HYALOMMA MALES 1. Without subanal shields. Festoons not fused. Adanal shields un. usually large and wide. Scutum with few large, scattered puncta. tions and long,clear lateral grooves. (India). (Subgenus Hyalommina)*....cscccccecccccsscccscccccscscccsccells HUSSAINI Figures 19S and 199 With subanal shields (exceptional, deformed runts may lack subanal SHIGIAS) ce.co-c cso oie sais dons Seo oe as wie bie eis oe wo iosemtwieletetelecehe “sretewetetns *The status of H. kumari Sharif, 1928, which falls into this classification, is uncertain. = 504 — Zs Coxa I simple, with two wide, short spurs from posterior margin. Scutum smooth, shiny with few, scattered, large punctations; festoons unfused; lacking lateral grooves and caudal depression. Adanal shields large, much like those of H. hussaini; sub. anal shields minute. (Tortoise para. site; Mediterranean and Black Sea areas, southern Russia to western Middle East). (Subgenus Hyalommasta)............H. ABGYPTIUM Figures 190 and 191 Coxa I deeply divided into narrow external branch and wider internal branch. Festoons partly fused. With short or long lateral grooves. Adanal shields not so large and wide as above. Subanal shields typically larger than above (frequently though abnormally as small as above). (Palearctic, Oriental, and Ethiopian Faunal Regions). (Sub genus SUSI) Mi ROME OUASOOUS 0c 6 65005 SERGE BONNOBUOORDOBODE:, Center of subanal shields characteris_ tically exterior of the axis of the adanal SHweildisis cyoisveveroiere c.c. eave ous colevererarelercicieleeievevetevciers ous c ests oe sec Center of subanal shields characteris. tically in line with the axis of the adanal shields (frequently laterally displaceds when ereatly, ENneOrsed) mane cece < cic ciclo esviceeeciscce 00 Spiracular plates of female type, with short, blunt "tail", (Camel parasite, Iran to Egypt; large size, UP ORS MMPI OTE )\aisralsraisse's cla eictetsleletete Male sta seinle. HeodatlnIOCHULAE Figures 202 and 203 Spiracular plates of normal male type, with long Cheese aso eeeeoee SSS SR OR ne — 505 e 5s Lateral grooves reaching scutal mid— length; scutum largely covered by small and medium-size punctations. (Medium-size; Near East; North, West, and coastal East Africa).......cceecceeeeeeeeceeeeHs IMPELTATUM Figures 170 and 171 Lateral grooves confined to posterior third of scutum; scutum smooth with few, scattered, usually large puncta. tions; posterior grooves characteris. tically marked. (Large ticks of camels wherever these animals now or recently OCCUFTOD ).. 0. ecccecccccccccsccscccccccccccccscoseeH. DROMEDARIL Figures 1 Lateral grooves confined to posterior third of scutum; scutum almost im. punctate except in caudal depressed area. (Usually small ticks; Mediter ranean Basin, Near and Middle East; East African coastal areas and northern Sudan) os cccccccscccccccccccccccccccecccsceseseses els EXCAVATUM Figures 1 Lateral grooves reaching at least scutal midlength (if specimen is heavily punc tate, Crooves. May De ODSCULE) «<.c sinc ce cc cess sicielsie cee v.cleicisie eeleiell Scutum smooth, shiny with rare, scattered large punctations; lateral grooves long and usually very distinct; posterior grooves well marked; legs usually not LT OC sjereralc c/o cle'sle/ciele/eiele eievelsislelesieielelaisieie.s ctais. ea aisjeimtarie ll aD Er uUl Figures 160 and [61 *A closely related "form" of uncertain taxonomic status, “H. scupense”, with some biological and morphological characters eae from those of typical H. marginatum, occurs in parts of the Soviet Union and southeastern hee: =9500.= 10. Scutum either densely punctate, ir regularly punctate, or lightly punc- tate except in caudal area where punc— Latlonapercedonsesicnep ence oes eee ear cce sets t ese sec emseO Scutum densely, entirely, and almost always uniformly covered by puncta tions often obscuring the lateral PEOOMGGs Meee cece cc as ccs ose ce ciicccise ssc oss ciccisecieicicsiecisiciie eee Scutum lightly or irregularly punc. tate, or densely punctate only pos. Wepenlorel ho GOG0 00 6 DOU OO DODO DOONNOSO00G00G eeeeoe ARITA EO CROCE Scutum strongly narrowed postero. laterally to form an almost recti. linear posterior margin. (West ENC) Meee nares sees esas tema ss case eiteeie elie ME NEOSUM Figures 17] and 175 Seutum bluntly rounded posteriorly... ....cccccccsccccccccces ell Circumspiracular area glabrous. Scutum brownish, punctations not so dense as in H. rufipes. Legs with dorsal enamelling on middle segments of two hind pairs. (Russia, Middle East, and South African Karr00)....ccccccccccsccccccccccsccecscsceoHs IURANICUM Figures 20 and 205 Circumspiracular area hirsute. Scutum black, punctations usually very dense. Leg segments usually distinctly annulated but lacking dorsal enamelling. (Near East, Arabia, Northeast Africa, tropical andeSouth Afriica)sccecis cc's cs «sos o'so sos cs's cesieeelese Hot RUFIPES Figures 182 and 183 - 507 = ll. 12. Lateral grooves long but often not well marked. Scutum irregularly punctate, posterior area not stri- kingly differentiated, posterior margin bluntly rounded. (Far, Middle, and Near East, Arabia, Northwest Africa, rare in North Gast ALTICA) sc1s.c sreisiiecsia sxeicieteeisie aierel osisic eiels oiaisicraaiclaiollio MAN GEN ALUM Figures 1 Lateral grooves long, distinct, deeply and cleanly delineated. Scutum smooth and shiny except posteriorly where it is densely punctate, posterior margin usual. ly rectangular. (Strictly African, notytneluding, North Africa)cccccssscccccceacdeccescctuceciaemle Central festoon undifferentiated. (Throughout Ethiopian Faunal een tens ree nee, ey ere TRUNC ATUM Figures 1 Central festoon forming a whitish or brownish parma. (East and West BPTICE). oc ccccccceccesccweses ces sasecesessiccieceH. ALBIPARMATUM Figures 194 and 195 = 50S— les FEMALES Genital apron divided into two lateral fields by a vertical median depression extending from posterior margin to mid. nae of apron. = Nea smooth. Subgenus Hyalommina)....seecseccecececeees cece esells) HUSOALNE Figures 200 and 201 Genital apron not divided by a Vert le auaimcGs OnweTOOVO ss sicies + sicleieiisieisisic #1 0\m0 6.010 «alse cls male o/eis.c)o% Coxa I simple, with two wide, short spurs from posterior margin. Scutum shiny, smooth, with few scattered, large, deep punctations. Genital apron a flat, widely triangular shield that becomes greatly de pressed posteriorly upon engorge— ment. (Subgenus Hyalommasta).........-..-- daslrestee H. AEGYPTIUM Figures 192 and 193 Coxa I deeply divided into a narrow external branch and a wide internal branch. (Subgenus Hyalomma)....sccssecessccescccccceececeeed Scutum regularly and completely covered by rather uniform, discrete punctations of moderate or large SHIPS GS SSO USO OO OO IOICIS CUU'O.0:0 SHACCEEISTIISIOtL O10 DOCG C Scutum with few to moderate number CLs iT Erepudar | PUNCLALLONS <1 crcisieisioeicis vice ccs ces esiecic clee ease *Exceptional specimens of H. turanicum may lack this aspect. Others of H. marginatum and H. impeitatum may approach this aspect. Sis) = Ae Genital apron a wide triangle composed of a protruding anterior ridge and a deeply, abruptly depressed, narrower POSLELLONM [DUG LOM ee clelslelsiofetsicle’ «cle siele elelelsloleleiolsie esiheietl om MER moO Figures 176 and 17”? Genital apron a wide shield with a broad posterior margin and not Galvided: aS) QDOVE cc:c:crcisicvelelce's'e 16 0'cle clove clersiecreve eleleletotele cieteversie crete Circumspiracular area hirsute. Middle segments of two pairs of hind legs lacking dorsal enamel. EVID Helavalalovese a /clevaisterevavelelelere'e aia aiaia/a%e\ o/s/aeelelalelsleleceteinvesejaieis lieu eter Figures 184 and 185 Circumspiracular area glabrous, or with few hairs. Middle segments of two pairs of hind legs enamelled Glopes WU hive RBG on So 6 OD CODOoOOO OS OOO On cao H. TURANICUM Figures 2 Genital apron an elongate triangle or GUIS CULT Sian ise lal reeed oa loro ve aucefoucave osanekeioreie eieieie eusiersveveis: syeveieretevedle Genital apron distinctly wider than long, a transverse oval, triangle, or Taal hc aoa eis va heater aan Gio wane evel ere naleks (aie teVornaietsiolnsisieie ere LO Genital apron a narrowly pointed triangle with lateral margins definitely longer than dorsal margin; gradually depressed in profile. Scutum usually as wide as or wider than long; with few, scattered large punctations; uncommonly with scat— tering of smaller punctations; surface frequently rugose. Usually Mar GO LCI crc.os:n/ois,cis/oerese siaiasicistersietete Seis eleieia exe aiajoisll ¢ DRCMMD ARE Figures 102 and 165 = OLOre Genital apron circular or a less narrow- ly pointed triangle with lateral margins approximating dorsal margin in length. Scutum usually somewhat longer than wide; punctations variable; usually less rugose. Size moderate (except H. schulzei) to SMA cr overetobetoverey cet cvorercrakerete cla crieieteatokerclereiaveisic (cial cievele mraveverecckeverctaroareie Genital apron a triangle bounded on each side by a lobe; greatly bulging in pro file. Scutum usually with sinuous pos. terior margin, numerous small puncta tions, and two irregular, submedian rows of large, deep punctations. Usual. BRP aes lescehrey Eris GaGa OO ODOOROO NGOS COC Cus eeeeeeeH. IMPELTATUM Figures 172 an Genital apron circular or triangular but lacking lateral lobes; greatly bulging or gradually depressed in profile. Scutum usually with very PEW MPUNCT ALT ONSisc sc cle's/e'< « sis slolelelelololohstcloloioictelotslerelerclelerelerelelsielslcle 7 Genital apron triangular; gradually depressed except when greatly em gorged but never convex in profile. Scutum smooth and shiny with very few, scattered punctations of large size; uncommonly with superficial small punctations. Usually moderate aA raialelelotelolelelcteletclelslelereialel\clele)eteleleielalsistaiets eleieisieleleiesie rien DE LIUCL UM Figures 160 and LOL Genital apron small, triangular or circular, convex or bulging in pro file. Scutum usually lacking smooth, shiny appearance of H. detritum; few, scattered punctations, those present small or moderate size; almost never large. Usually small size and narrow SIAC ise alaleietelalelalsiale|oic/e/ers) ateleleiela/e\aiele slotelelele/els eles eeieieicl nA CAVALUM Figures 168 an =, Sib — 10. TRS Ws Genital apron a transverse oval, in profile deeply concave poste. riorly and with a narrow, bulging lip anteriorly. Scutum usually smooth and shiny with few scattered punctations, with or without in conspicuous interstitial punctations. (Ethiopian FaunalRestontonly)s.ceccccenccessceceke coe name etl Genital apron not deeply concave in profile. (Absent or exceptional and only peripherally in Ethiopian Faunal ReHiGH) al; Jota G6oRa hen, Tinne, auOTarmin ahtyrh telat Common, widely ranging species throughout Ethiopian Faunal Region..............eH. TRUNCATUM Figures 188 an Uncommon species, known by small populations only from Kenya, Tanga. nyika, and Togo. Differentiated only by association with male.................H. ALBIPARMATUM Figures an Genital apron a large, transverse oval; flat, somewhat protruding, or convex in profile. Circumspiracular area usually markedly pilose. Usually large size. (Palearctic Faunal Region ON yin as cies sieve. cleie siviels sieisinieltioleieicie cs gninsin ins ocisisteaere oll om CMU LsGral Figures” ~an Genital apron a large robust shield; transversely oval or widely triangular in outline; strongly bulging in pro- file; scutum with moderate to numerous punctations of variable size.....seeeeeeeeeeeee oH. MARGINATUM Figures 150 an aoe ee Figures 190 and 191, co, dorsal and ventral views Figures 192 and 193, 9, dorsal and ventral views A, Q, genital area. B to D, genital apron, outline and profile. B, unengorged. C, partly engorged. D, fully engorged. HYALOMMA ABGYPTIUM Specimens from Land Tortoise, Bastern Anatolia Hoogstraal Collection PLATE LV - 513 - HYALOMMA AEGYPTIUM (Linne, 1758). (Figures 190 to 193) THE TORTOISE HYALOMMA The much mooted name H. aegyptium has been frequently used as a “catchall" by workers in many countries for a number of spe- cies. Consequently much literature referring to Hyalomma ticks has been hopelessly confused, even to the present = King (1926) lumped all Sudan species under the name H. aegyptium*, as did most other workers on African and Near Eastern oe of his time. H. aegyptium (Linne, 1758) is now recognized as a distinct parasite of tortoises in the Mediterranean area and Near East. In Russia it is confined to Crimea, Georgia, Armenia, the Caucasus, Azerbaijan, Turkmen, Uzbek, and Tadzhik (Pomerantzev 1950). It is common in many parts of Asia Minor (Hoogstraal, ms.) and occurs in Afghanistan (Anastos 1954). The tortoise hyalomma does not occur in the Sudan, elsewhere in tropical or southern Africa, or in Europe away from the Medi. terranean and Black Sea areas. Although originally described from Egypt, where tortoises are said to have been numerous on the Medi_ terranean littoral, H. aegyptium apparently is now extinct here. In present-day Egypt, tortoises occur only a few miles east of the Libyan border — extremely rarely as far as seventy miles east of Libya — and in Sinai a few miles west of Palestine. No ticks have been found on recent Egyptian tortoises, except on specimens from Palestine in Cairo pet-shops. Tortoises are the hosts of predilection of adult H. aegyptium. Exceptionally, lizards, hares, and hedgehogs are attacked. Witte other mammals may be infested rarely, long lists of various hosts for this tick are all erroneous, based as they are on old records in which all species were lumped under the name H. aegyptium. In *With reference to reports of "H. aegyptium" from the Sudan, it should be noted that the actual tick species with which O'Farrell (19134,B), did his interesting work on an entomogenus trypanosome, Crithidia (?Herpetomonas ) hyalommae, cannot now be determined. = SA — Anatolia, however, when rearing larvae and nymphs from tick-infested animals, it was found that these stages commonly attack tortoises, lizards (Agama), partridges, man, hares, hedgehogs, and a wide variety of rodents (Hoogstraal, ms.). A number of Russian host records were presented by Olenev (19288). The life cycle of "H. aegyptium" described by Nuttall (1915) applies to H. marginatum. eerie lot 1305a in British Museum (Natural History) was used for this study. No material for his lot X (from Rome) can now be located. It appears that the study of the external morphology of each stage and of the bionomics of "H. aegyptium in India (Sharif 1924) applies actually to H. excavatum, but this is not certain. A popular article concerning the actual H. aegyptium has recently appeared in the Illustrated London News Te 1950). Based on living ticks arriving in the British Isles on pet-shop tortoises from southern Europe, this account should interest persons who frequently encounter this name promiscuously used in the literature. Distribution data in the Browning paper are from literature references under H. aegyptium, and, therefore, far more extensive than the actual geographic range of this species in nature. Contempor iblished reports on disease transmission by ticks ortunately continue to perpetuate the early confusion ee aA eS een COnrusron in identification of species in Ss genus. The tortoise para Eaceed (ee SSS eS eoues site, H. ae tium, has never been incriminated as a vector or aos Sone — ———. oe i = reservoir of pathogenic organisms of man, other mammals, or irds. H. aegyptium does, however, transmit two sporozoa to land tortoises in northwestern Africa and in the Near East. These are Haemo arina mauritanica and H. stepanovi (cf. Sergent and Sergen > Laveran and Negre 1905, Nicolle and Comte 1905, Laveran and Pettit 1910, Brumpt 1938; and Laveran 1901, Marzinowsky 1927, Popovici-Baznosano 1901,1906,1907, and Reichenow 1910). Another parasite from this tick Coelomoplasma eee has been briefly described by Brumpt (1938) er er classification as to group (see also Brumpt 19385). Further studies on these parasites have not been encountered. = Silene Certain unusual morphological characters of this easily recog nized species cause the tortoise hyalomma to be considered in a separate subgenus, Hyalommasta Schulze, 1930. H. aegyptium ctata Schulze, 1919 (see also Kratz 1940), a name proposed for a single specimen from Malta and subsequently overlooked by even Schulze himself, is undoubtedly a synonym of H. aegyptium. Pomerantzev (1950) considers Dermacentor rosmari Ass (1935), described from nymphs from walrus in the Kara (White) Sea and said to be the northernmost ticks on record, as a synonym of H. aecyptium. On the basis of the original description and illus. trations of D. rosmari, its systematic position is uncertain but this synonymy is hardly convincing. The synonymous name H. syriacum Koch, 1844, has frequently been used for the actual H. aegyptium. So far as has been deter— mined, H. syriacum has been confused with no other species. Both sexes can be readily distinguished from key characters provided herein. = 5167= Figures 194 and 195, co’, dorsal and ventral views Figures 196 and 197, 9, dorsal and ventral views A, 9, genital area. B and C, 9, genital area, outline and profile, unengorged. HYALOMMA ALBIPARMATUM Specimens from cattle, Kenya; from Nuttall lot 3773, exchange, British Museum (Natural History). Hoogstraal collection. PLATE LVI - 517 - HYALOMMA ALBIPARMATUM Schulze and Schlottke, 1930 (= H. BRUNNEIPARMATUM S. and S., 1930). (Figures 194 to 197) THE PARMATED AFRICAN HYALOMMA H. albiparmatum, presently known only from Kenya, Tanganyika, and Togo, 1s SimLlar to H. truncatum except that the central fes. toon is differentiated as a parma in the form of a celluloidlike cover of variable size, shape, and color. No constant characters have been discovered for differentiating females of these two species. Delpy (1949B) considered H. impressum albiparmatum Schulze and Schlottke, 1930, from East Africa, as a synonym of H. mar_ ginatum (= H. savignyi) or questionably of H. truncatum (= H. transiens); he considered H. brunneiparmatum o ze and Schlottke (1930) to be a synonym of H. marginatum (= H. savignyi). Feldman-Muhsam (1954) stated, after examining Kenya laboratory- reared material of H. albiparmatum submitted by J. B. Walker that H. albiparmatum is a synonym of H. truncatum. Dr. Theiler and the writer, after examination of the same material, are convinced that Walker's H. albiparmatum is a distinct genetic entity worthy of full species ar Tt is also apparent from Walker's material and from other series in British Museum (Natural History) collec- tions that H. impressum brunneiparmatum Schulze and Schlottke, 1930, from Togo is a synonym of H. albiparmatum and not of H. truncatum as stated by Feldman-Muhsam SEL. Miss Walker is preparing a report on this species for publication in Parasitology. Hosts of the parmated African hyalomma appear to be the same as those of H. truncatun. British Museum (Natural History) collections contain material from cattle and from a rhinoceros from Zatta Plains, Kajiado, Namanga, and the Masai Reserve, Kenya (HH det.). J. B. Walker (correspondence) has seen specimens from a rhinoceros from Iringa District, Tanganyika, and from a warthog in Chunya District, Tanganyika. Skips Figures 198 and 199, co’, dorsal and ventral views Figures 200 and 201, 9, dorsal and ventral views A, 9, genital area. B to E, o, genital apron, outline and profile; all unengorged; E dried, contracted specimen. HYALOMMA HUSSAINTI Specimens from Bihar, India (det. Sharif) (Rocky Mountain Laboratory collection). PLATE LVII = Bulls) HYALOMMA HUSSAINI Sharif, 1928. (Figures 198 to 201) HUSSAIN'S INDIAN HYALOMMA AND REMARKS ON THE SUBGENUS HYALOMMINA Special note should be made of the subgenus Hyalommina, es. tablished by Schulze (1919) for the new species H. rhipicephaloides from the Red Sea area. Subsequently, Schulze (1936) SSS H. lewisi from Tanganyika (and Kenya) in this subgenus and Sharif (1928 and 1936) included H. kumari and H. hussaini from India. These are discussed below. The criterion proposed by Schulze for this subgenus is the absence of subanal shields. With regard to the so-called “H. rhipicephaloides", it has been our experience during field collecting and study of Hyalomma material from the Near East, Asia Minor, Arabia, North Africa, and tropical Africa, that weak, poorly developed, apparently undernourished, runts of any Hyalomma species frequently lack subanal shields. Such individuals may be part of a series in which some are typical of a common species (Such as H. excavatun) cies in morphological characters, except that they lack subanal shields. “tt has ote erence and from series in British Museum (Natural History) collections, especially those of the late Professor Buxton from Iraq and Palestine, that when nymphs are removed from a bird, lizard, or small mammal and placed in a vial to molt, the resultant adults, obviously affected by abnormal, artificial conditions, are frequently frail and lack subanal shields. This feature is the rule rather than the exception among adults reared from nymphs that have become overgrown by the host skin (see below and page 447). Schulze (1932C) referred to "Hyalomma (Hyalommina) rhipi- cephaloides" as a “half endoparasite™ (and compared it with a 520r— Amblyomma nymphs to support a morphological theory). The concep. tion of a separate species with unique "half-endoparasitic" habits is not supported by field and laboratory observations. In several Hyalomma species observed in Egypt, long-feeding immature stages become overgrown by the host skin. Poorly developed adult "Hyalom minas" result from these nymphs. If removed early enough, such nymphs may molt into typical though frail adults of recognized species with or without subanal shields. Other larvae and nymphs that attach to the ears, which do not react to the engorging ticks by producing a large amount of tissue, usually develop normally. On the basis of many such variants among specimens of H. excavatum examined for the present study it is apparent that Delpy's (19498) synonymy of H. rhipicephaloides under H. exca vatum is correct. H. lewisi Schulze, 1936, from tropical Africa is the result of similar misinterpretation of H. truncatum by Schulze and his students. In the present collection, a few specimens of H. truncatum are poorly developed and lack subanal shields. These are similar to specimens (in the Rocky Mountain Laboratory) determined by Schulze as H. lewisi from Kenya and Tanganyika. There is no question but that H. lewisi is a synonym of H. truncatun. Delpy (19498), probably inadvertently, listed Hv lewisi as a synonym of H. excavatum. Kratz (19403 retained H. [ewisi in the "subgenus Hyalommina" even though he noted that "some spec- imens retained subanal shields while others lacked them". With regard to H. hussaini Sharif, 1928, the Rocky Mountain Laboratory collections contain enough constant specimens of this species (described below) from India to indicate beyond a doubt that H. hussaini is a valid species. It is coincidental that this Species conforms to the criteria proposed for the subgenus Hyalommina; H. hussaini rather than H. rhipicephaloides might, eee be considered as the type species of this subgenus. The absence of subanal shields apparently has become a genetically- established character in Indian populations. As stated below, other constant male and female characters also validate this spe- Sey = cies. Thus, recognition of the subgenus Hyalommina would be justi- fied. It is, however, likely that the absence of subanal shields is not a genetic character in Hyalomma populations of Africa and the Near East. Conclusions on the subgenus Hyalommina may be summarized as follows; Such an entity apparently does exist, but criteria proposed for it apply to a species (H. hussaini and possibly H. kumari) different from that originally proposed as the type for this subgenus (H. rhipicephaloides), this latter species being merely a morphological variant of H. excavatum. These conclusions are based on study of series of preserved specimens, on field rearing of specimens, and on laboratory ob. servations of wild-caught subdermal specimens from rodents. More formal laboratory studies on the phenomenon of loss of subanal shields among other species are indicated. Sharif (1928) also described H. hussaini brevipunctata and H. kumari from Indian populations on the basis of sti ht dif. ferences in color, lateral grooves and tarsi. No specimens of these forms have been available for the present study. Sharif (1928) lists specimens of H. hussaini from the fol. lowing India areas: Bihar, Orissa, Central Provinces and Madras and Bombay Presidencies. The subspecies brevipunctata is listed from the same areas as well as from ee eee is also known from the first localities and Assam and Punjab. Hosts are cattle, buffalos, horses, goats, sheep, dogs, tiger, and various kinds of deer. Sharif (1930) illustrated a specimen of H. hussaini with unequal adanal shields. Material from Portugese India has been reported (Santos Dias 19543). Both sexes of H. hussaini have such unique morphological characters that it is difficult to comprehend why Delpy (19498) placed this species in synonymy under H. excavatum. Sharif's (1928) original description is excellent as are the illustrations of the male. This sex is characterized by large, broad adanal shields, absence of subanal shields; bright, shiny scutum with long, pronounced lateral grooves; long, narrow posteromedian ipo = grooves, shorter and wider paramedian grooves; rarity of punctations that are widely scattered over scutal surface but usually arranged in lines bordering the posterior grooves, and small size (less than 3.00 mm. long and 2.00 m. wide). The female also has a smooth, shiny scutum with few punctations, those present are similar to those of the male. Porose areas of the. females at hand are notably large and distinct. The genital apron . is unique in that it is divided by a medioposterior depression that is either a narrow, median, posterior groove or expanded posterior- ly to include the posterior periphery. In outline the genital apron is subrectangular to subtriangular, in profile it is more or less gradually depressed posteriorly. The female size unengorged is only slightly greater than that of the male. = 52350— ITIAT aly Id OIIAX FIVId ees ‘suotyerysnTIT 6 s0g “TT@8H “H °D “Ad Wory UOTYOSTTOD TeeryssooH *beay Teryjueg ‘Teueo worjy uoutoeds IAZTNHOS VWWO'TY AH H *Traepeuoip *H ‘eqeTd TeTnoeitds ‘y¥ *TezTnyos *y feqyetTd reqtmoeatds ‘g SMOTA TeljUeA pue Tessop {9 *¢oz pue zoz sem3ty - 524 - HYALOMMA SCHULZEI Olenev, 1931(B). (Figures 202 and 203,337 and 338) THE NEAR EASTERN CAMEL HYALOMMA H. schulzei a hyalomma of restricted geosraphical range, appears from male characters to be a giant relative of H. dromedarii. The fe male genital apron, however, differs so greatly from that of H. drome darii that the relationship of these two species does not appear to be actually so close as previously considered. It is also likely that females associated by Olenev with males of H. schulzei in the original description of the species included some specimens of H. dromedarii. - The geographic range of H. schulzei extends from Iran (Olenev 1931A,B, Delpy 1937) and Afghanistan (Anastos 1954), through Iraq (Hoogstraal, ms.), and Palestine (Schulze 1936C; Kratz 1940, Adler and Feldman_Muhsam 1946,1948) into Sinai and the Eastern and Western Deserts of Egypt some 200 miles west of Alexandria (Hoogstraal, ms.). It is absent in Russia (Pomerantzev 1950) and is not represented in the present collections from Libya, Yemen, coastal East Africa, or Turkey. Hosts of adults mentioned by all authors and represented in our collections are camels, and Pomerantzev (1950) adds cattle. Hosts of immature stages have not previously been determined. In Sinai and Egypt (Hoogstraal, ms.), nymphs (reared to adults) have been found on hares, Lepus capensis subsp., and fat sandrats, Psammomys o. obesus. Unfed, newly molted adults have been taken from bur— rows of jirds, Meriones c. crassus. The male is large and when engorged measures up to 8,00 mn. in length and 7.00 mm. in width. The extreme width is due to con siderable lateral stretching of the integument during feeding; un. engorged specimens are approximately 4.00 mm. wide. The scutum of specimens at hand and previously described has parallel lateral margins while those of H. dromedarii are usually convexly arched. The palpi of available specimens are notably short and robust; the parma is subrectangular; in H. dromedarii it is typically tri- engular. The most notable distinction between the two species is =VS2 = the short, almost tailless, female-type spiracular plate of the male H. schulzei that is usually surrounded by numerous hairs. This type of spiracular plate is found in the male of no other species in the genus. An approach to this spiracular form occurs in "H. scu ense"; see page 418.7 In other characters, these two species are similar. Females of these two species are also superficially similar but the genital apron of H. schulzei is considerably different and the scutum (of available specimens) is slightly more elongate than that of H. dromedarii. The genital apron is subrectangular with slightly convex anterior and posterior margins and rounded junctures; this convexity is increased with extreme engorgement. In profile, the apron is flat when unengorged but after feeding it protrudes and is markedly depressed posteriorly. The female is illustrated herein in the APPENDIX (PLATE XIXC) inasmich as distinguishing features were recognized only near the termination of the present study after other illustrations had been numbered. Delpy and Adler and Feldman-Muhsam did not describe the female genital apron of the meagre amount of material available to them. The short, wide scutal outline with a bluntly rounded posterior margin, as illustrated, is similar in all available specimens and corresponds to that delineated by Pomerantzev (1950). Other workers however, have shown it as more elongate and with the posterior margin gradually converging to a fairly narrow point. = 5260= Figures 204 and 205, o@, dorsal and ventral views Figures 206 and 207, 9, dorsal and ventral views A, 9, genital area, unengorged. B to H, 9, genital apron, outline. B and C, wnengorged. D and E, slightly engorged. F to H, moderately engorged. J, c', Leg III, dorsolateral view to show enamelling. HYALOMMA TURANICUM South African Karroo Specimens Reared by Dr. G. Theiler PLATE LIX = BUS HYALOMMA TURANICUM Pomerantzev, 1946 (= H. RUFIPES GLABRUM Delpy, 1949) NEW NAME COMBINATION (Figures 204 to 207) THE ENAMEL-LEGGED HYALOMMA H. turanicum, is considered by South African and French work. ers as a subspecies of H. rufipes and by Russian workers as a sub. species of H. mrginatum (= H. plumbeum of Pomerantzev). The specific entity of H. glabrum has been demonstrated in laboratory rearings by Theiler, who has kindly provided material for the present study and has provided a manuscript (Theiler 1956) on distribution and ecology in South Africa for use herein in ad vance of her own publication, H. turanicum (= H. abrum) appears to have been introduced into the arid South African Karroo on Persian sheep. It is um known elsewhere in Africa. Persian sheep were originally intro- duced into South Africa in 1872, having been purchased from a ship from the Mediterranean then anchored in Table Bay (Lounsbury 1904E). Subsequently others were imported from Aden. The exact locality from whence any of these importations originated does not appear definitely to be known. A number of flocks were scat. tered about South Africa at the time (1904) Lounsbury reported their high degree of immunity to heartwater. H. rufipes glabrum was briefly described by Delpy (1949A) from material rear rom females from Karroo sheep sent to him by Theiler. The source of this material has been identified in correspondence with Dr. Delpy and Dr. Theiler; it is not found in the literature. Subsequently, Delpy (1949C,1952) indicated that H. rufipes glabrum is a poorly known, two host tick, that it also occurs in Iran, and that it is not of considerable im portance in the transmission of bovine theileriasis, Theileria annulata. This species occurs in southern Russia (Pomerantzev = 5265 1950; as H. plumbeun turanicum) and a single specimen has been col. lected in Afghanistan (Anastos 1954; as H. rufipes glabrum). H. marginatum turanicum Pomerantzev, 1946 /Subsequently (1950) considered as a subspecies of H. plumbeum by eae author 7 is obviously the same tick as H. rufipes flabrum Delpy, 1949. Tt is necessary, therefore, to cive the Soviet name priority. There is no biological reason or taxonomic utility for considering H. turanicum as a subspecies of any species. Theiler (1956) continues to apply the name H. glabrum to South African populations of this species. Rousselot (1951) reported H. rufipes glabrum from French West Africa but omitted it from his T9SsD aE » apparently after having reconsidered the identification of pertinent material. Villiers (1955) also used this name for material from the same area. It is assumed that these data refer actually to H. rufipes. Fonesca, Pinto, Colaco, Oliveira, Branco, da Gama, Soares Franco, and Lacerda (1951) reported "H. rufipes glabrata" fron Portugal, but Theiler, who has seen their material, states (cor_ respondence) that it is an entirely different species, probably H. marginatun. According to Pomerantzev (1950) the biology of H. turanicun was reported in 1945 by Lototsky, pp. 69-130, but it has been impossible to secure further details for the present study. The distribution of this tick is stated to be southern Khazakstan, Middle Asia, and Iran. To this range, the South African Karroo and Afghanistan should be added. Where it occurs in the Soviet area, H. turanicum ranges through a number of altitudinal zones. It lives in tugai meadows, in semidesert, and in low areas of fields at the base of hills and on mountain slopes to the wooded belts. Adult hosts are "large and small horn-bearing animals", especially cattle, and also horses. A single Afghanistan male was taken from a camel (Anastos 1954). Immature stages attack birds; these hosts are listed below and those most commonly infested are preceded by an asterisk. = G23) = Alectoris kakelik kakelik (?= A. x eca falki), Chukar partridge Phasianus chrysomelas bianchii (= P. colchicus bianchii), Ring- neck pheasan Columba livia neglecta, Rock dove Columba eversmanni, Bastern stock pigeon Streptopelia turtur arenicola, Turtle dove SS Neophron percnopterus, Egyptian vulture Coracias garrulus, Roller Merops persicus, Iranian beeater Upupa epops, Hoopoe Sturnus garis dresseri, Starling Pastor roseus, Rosy starling Carduelis carduelis subcaniceps, Goldfinch Passer domesticus bactrianus, House sparrow *Passer montanus zaissanensis, Tree sparrow Miliaria calandra buturlini (= Emberiza c. buturlini), Corn bunting Mmberiza stewarti, White-capped bunting — *Fmberiza buchanani huttoni, Grey-necked bunting Emberiza icterica, Redheaded bunting *Melanocorypha calandra, Calandra lark SEL Ee acutirostris, Oriental short—toed lark *[Tauda guleula inconspicua, Oriental sky lark Anthus eee griseus, Tawny pipit tephronota, Sitta Rock nuthatch Lanius == Lesser grey shrike Lanius collurio isabellinus, Red-backed shrike Phylloscopus nitidus viridanus, Greenish willow warbler S iia curruca, Lesser whitethroat a althaea, Hume's lesser whitethroat Saxicola torgquata, Stone chat Oenanthe ca but scutum Longer®. The Kaka specimen is a poorly preserved nymph that appears to be I. rasus but cannot definitely be assigned to this species. i DISTRIBUTION The actual range (and identity) of subspecies of Ixodes rasus is unknown (see REMARKS below). The species is especially numerous in many parts of West and Central Africa and locally common in East Africa. Material on which the few records of South African specimens are based should be restudied. =) 550) = WEST AFRICA: NIGERIA (Pearse 1929). FRENCH WEST AFRICA (Neumann 1899,1901,1911. Rousselot 1951,1953B. Villiers 1955). Saaee (Nuttall and Warburton 1911. Nuttall 1916. Stewart 1936). CENTRAL AFRICA: FERNANDO PO (Schulze 1943A). CAMEROONS (Neumann 1899,1901,1911. Ziemann 1912A. Nuttall 1916. Jojot 1921. Rageau 1953. Schulze 1943A. Rageau 1951,1953. Dezest 1953. See HOSTS below). FRENCH EQUATORIAL AFRICA (Bequaert 1931 refers the type locality to French Equatorial Africa, not to Belgian Congo. Fiasson 1943B. Rousselot 1951,1953B). RIO MUNI (Nuttall 1916. Schulze 1943A). BELGIAN CONGO (See French Equa. torial Africa above. Neumann 1899,1901,1911. Nuttall 1916. Nuttall and Warburton 1916. Schwetz 1927C,1932. Schouteden 1927. Bequaert 1930A,B,1931. Schoenaers 1951A. Rousselot 1951,1953B. Theiler and Robinson 1954). EAST AFRICA: SUDAN (King 1926. Hoogstraal 195/B). ETHIOPIA (Nuttall 1916). KENYA (Neave 1912. Nuttall 1916. Anderson 1924A. Lewis 1931A,B,C,1939A. See HOSTS below). UGANDA (Nuttall and Warbur_ ton 1911. Mettam 1932,1933. See HOSTS below). TANGANYIKA (Neumann 1899,1901,1907C ,1910B,1911. Morstatt 1913. Bequaert 1930A. Reichenow 1941B. Schulze 1943A). SOUTHERN AFRICA: SOUTHERN RHODESIA (Nuttall 1916). /7UNION OF SOUTH AFRICA (Bedford 1929A,1932B. Cooley 1934. Some or all of this material refers actually to Ixodes pilosus: Arthur, cor- respondence./ HOSTS The range of hosts of adult Ixodes rasus includes a large variety of animals from mice and elephant shrews (small insecti- vores) to leopards, large antelopes, and domestic dogs. It is difficult to determine whether this tick displays predilection for any group of mammals. The picture for immature-stage host preferences is equally uncertain. Whether individual "subspecies" have characteristic preferences differing from each other remains to be demonstrated. - 551 - Published reports of adults and immature stages found together on a single host are categorized below by associated stage and sex for what they may be worth in eventually answering questions of host preferences and biology of this species. Stage or sex not specified: Domestic cattle from Uganda, leopard and domestic dog from Gold Coast (Nuttall and Warburton 1911). Domestic sheep from Kenya (Lewis 1931B). Cricetomys emini (giant forest rat), Crocidura sp. (shrew), Crocidura manni, Hybomys univittatus (back-striped mouse), Leggada NS a al Tmisculoides (pygmy mouse), Lemniscomys striatus (striped grass- mouse), Rattus rattus (house rat), TteriTins acilus angelus (gerbil), Thyronomys swinderianus (cane rat) factes 1S From Nigeria). Buffalo, blue duiker, forest or red duiker, and domestic cattle (Mettam 1932, Ae eee). Black duiker (Villiers 1955, from Ivory Coast). Warthog (Ziemann 1912A, from Cameroons). Cane—rat (Schwetz 1927, from Belgian Congo). Okapi (Schouteden 1927, from Belgian Congo). Impala (Bedford 1932B and Cooley 1934, from South Africa). Tree hyrax, Dendrohyrax (Schoenaers 1951A, from Belgian Congo). Larvae alone: Common on small and large birds in Kenya (state ment Should be checked for accuracy) (Lewis 1939A). Nymphs alone: Bdeogale mongoose (Schulze 1943A, from Tanga- nyika). Man (Nuttall 1916, from Ethiopia). "Antilope brune" (Bequaert 1931, from Belgian Congo). Females with larvae: Okapi (Schwetz 1927, from Belgian Congo). Females with larvae and s: Akeley's suni or dwarf nymph antelope, Nesotragus moschatus aie i, (from Kenya; this female tick corresponds more closely to the description of Ixodes rasus cumulatimpunctatus than any other specimens that I have seen). Mole rat, Tashyoryctes sp. (HH legit at Njoro, Kenya). ae pra Females with nymphs; Water chevrotain, giant elephant shrew, giant forest rat, = a pig, mongoose, and domestic dog (Schulze 1943A, from Tanganyika and Cameroons); Hyrax (Procavia sp.) (Bedford 1929A,1932B, from South Africa). "White mongoose" (specimens seen by HH, from Cameroons). Females alone: Domestic dog (Schulze 1943A, from Tanganyika; Rageau 1951, from Cameroons; Nuttall 1916, from Gold Coast). Do mestic cattle (Nuttall and Warburton 1911, from Uganda*). Domestic goat (Nuttall 1916, from Belgian Congo). Tree hyrax, Dendrohyrax arboreus (Bequaert 1931, from Belgian Congo). Hyrax (leccavio) (Schwetz 19270, from Belgian Congo). Giant forest rat (Cricetomys) (Schulze 1943A, from Rio Muni; others seen by HH, from Uganda). Grass rat (Arvicanthis abyssinicus nubilans) and groove-toothed rat (Otomys tropicalis elgonis) (spec- imens from the latter host closely correspond to See: ption of Ixodes rasus cumlatimpunctatus (HH legit at Njoro, Kenya). Duiker (Bequaert 1930, from aa Serval from Kenya, pangolin and bush pigs from Cameroons (Nuttall 1916). Leopard (Nuttall et al 1911, from Cameroons*). Large erey mongoose eee caffer) ? and ered mongoose (Myonax cauui) (Bedford 1 Tom south Africa). Females and males: "Man and domestic dog" (Nuttall et al, from Gold Coast®). Hyrax (Neumann 1899, from French Equatorial Africa — not Belgian Congo according to Bequaert 1931) 3; bush pig (Schulze 1943A, from Tanganyika); leopard (Nuttall 1916, from Southern Rhodesia; others seen by HH, from Kenya). Pangolin (Specimens seen by HH, from Uganda). Males alone: Hare (Sudan record above). Caged chimpanzee (Speci mens seen by HH, from Cameroons). *Stages or sex not reported in literature cited, but determined from Nuttall collection in British Museum (Natural History). - 553 - BIOLOGY After having collected numerous specimens of Ixodes rasus from many small mammals in Kenya, from sea level to 8000 feet elevation, it was surprising to find this species so rare in the Sudan. From Pearse's (1929) report on Nigerian ectoparasites, I. rasus appears to be common on small mammals there. The reasons for these considerable differences in local populations remain to be explained. Nuttall (1911) categorized I. rasus in the biological group within the genus Ixodes in which males and females are usually found together on a host that either wanders or does not travel far and in the subgroup in which the sexes are often found in cone on the host. Present evidence partially supports the inclusion of Ixodes rasus in the last subgroup and careful col. lecting may subsequently prove that this species should be so considered. Systematic search probably will show Ixodes rasus to be a common tick in many parts of tropical Africa. Examination of rodents in certain areas should prove fruitful. Seeking for every tick on a variety of animals will undoubtedly reveal a few, small inconspicuous, well-hidden Ixodes ticks along with more apparent, more numerous, larger, and more colorful ticks of other genera. Questions concerning the biology of Ixodes rasus in its various forms (i.e. subspecies) offer a challenge in one of the most fascinating zoological areas of the world. DISEASE RELATIONS Unstudied. REMARKS Schulze (1941) noted certain features of the haller's organ of I. rasus (cf. also K. W. Neumann 1943). A comparison of this - 554 - organ with that of other Ixodes species is presented in table form by Arthur (1956B). a Remarks on morphology of this species are contained in papers by K. W. Neumann. These have not yet been seen and complete ref-— erences are not available except for: (1942) Z. Morph. bkol. Tiere, 38(2):358, 362. (Morphology of subcoxa of subspecies cumlatim. punctatus); and (1925) Zool. Jb. (Anat.), 69(2):286, fig. 8. Jakob B) included this species in his review of the relationship of tick genera from the standpoint of comparative morphology. Both sexes are readily distinguished by their completely closed, usually circular anal grooves, an easily observed character in all specimens except some greatly engorged females in which the anal area is depressed and the associated grooves are difficult to dis. cern. Only one other African species, Ixodes ugandanus Neumann, 1906 (with which Ixodes SS Warburton, ios from Uganda is quite possibly synonymous), is known to have closed anal grooves; these are not circular but characteristically oval and unite in a slight posterior elongation. I. ugandanus has not yet been found in the Sudan. ee ene Schulze (1943A) indicated that circular anal grooves are char- acteristic of this species but that some specimens in which these grooves are expanded to a broad oval outline are merely atypical individuals of the same species. According to Schulze, the anal grooves of I. rasus may even be narrowed posteriorly. Schulze (1943A) differentiated three subspecies of I. rasus and one "related species" as follows: iI. rasus rasus; Short, broad palpi and hypostome; retro- grade auriculae (lateral spurs of ventral basis capituli); pro nounced single punctations on scutum. (From Cameroons). I. rasus SS ae Long, narrow palpi and hypo- stome; perpendicular auric ae; and small scutal punctations, some of which appear to be formed of a small group of smaller, contiguous punctations. (Tanganyika to Fernando Po). i. rasus eidmanni: The same characters as given above for I. rasus rasus, but "more strongly chitinized and darker", and - 555 - denticles of hypostome with a small apical "hook", proximity of sensory organs in the integument, and absence of a definite "peripheral zone" of the integument. (Rio Muni, or Spanish Guinea). iI. vanidicus: Similar to I. rasus but with anal grooves horseshoe shaped or circular but not closed posteriorly. (Tanga. nyika to Cameroons). IDENTIFICATION Males are characterized by circular anal grooves (see REMARKS above), narrow marginal fold beside the scutum, fairly many to numerous fine scutal punctations, and short palpi with segments 2 and 3 of about equal length. This reddish brown species is about 2.8 mm. long and 1.8 mm. wide and has a broadly rounded posterior margin. The tarsi are usually humped but in some specimens which otherwise conform to this description they are tapering. Females also have closed anal grooves. Their scutal puncta- tions are similar to those of males. The hypostome and palpi are long and narrow and the basis capituli ventrally has a large spur (auricula) arising from each lateral margin. The tarsi are either tapering or somewhat humped. Note; As presently considered, any African Ixodes male or female with circular, closed anal grooves is I. rasus. Within the large amount of material seen in various Collections there is considerable variation in most other characters, the significance of which await to be determined by Dr. Arthur. =) 550l= 227 229 Figures 226 and 227, co, dorsal and ventral views Figures 228 and 229, 9, dorsal and ventral views IXODES SCHILLINGSI Q Sudan specimen oO Kenya specimen PLATE LXV = 557 = IXODES SCHILLINGSI Neumann, 1901. (Figures 226 to 229) THE COLOBUS RUSSET TICK Lb, ON Fore: BQUATORIA PROVINCE RECORD BE Lotti Forest Colobus polykomos dodingae Apr Lotti Forest is at 4500 feet elevation. This is the only record of this species from the Sudan. About a dozen monkeys of the same species examined in Lotti Forest and at other places at different altitudes in the same forest were free of ticks. An equal number of monkeys, Cercopithecus mitis stuhlmanni, from Lotti Forest were tickless, as were numerous specimens of other monkeys (Setzer 1956) examined elsewhere in the Sudan. DISTRIBUTION I. schillingsi, a parasite of East African Colobus monkeys, does not appear to range outside of wooded savannah and forested areas. It is the sole described African representative of a complex of species occurring in Asia, Madagascar, Australia, and Europe. EAST AFRICA: SUDAN (Hoogstraal 1953E,1954B. Arthur, ms.). KENYA (Nuttall 1916. Lumsden 1955. Arthur, ms. See HOSTS below). TANGANYIKA (Neumann 1901,1907C,1910B,1911. Nuttall and Warburton 1911. Morstatt 1913. Arthur, ms.). ZANZIBAR (Arthur, ms.). SOUTHERN AFRICA; MOZAMBIQUE (Neumann 1919A,1911. Nuttall and Warburton LOlL. Santos Dias 1953B). HOSTS Man is parasitized by I. s. schillingsi according to a single record for a female tick of this species —n Njoro, Kenya (Lumsden = SEE = 1955). Aside from this, Colobus monkeys, also known as guereza or leaf-eating monkeys, are the only known hosts. As stated below, all other records from different mammals, including man and monkeys other than Colobus refer to undescribed Ixodes spe— cies or subspecies. Colobus polykomos caudatus (Neumann 1901,1907C,1910B. Nuttall and Warburton Totty C. polykomos dodingae (Sudan record above). Colobus sp. (Nuttall seo Anderson (1924B) recorded Rattus rattus kijabius as a host in Kenya. Specimens of this tick species collecte Anderson in Kenya, now in British Museum (Natural History) collections, are labelled as from Colobus monkeys and others bear no host data (Arthur, ms.). The rodent host is believed to be an error. Lewis (1931C) listed this tick from a duiker, bushbuck, and domestic cattle in Kenya. These four collections, two from bush. bucks and one each from the other hosts have been examined at British Museum (Natural History) and found to refer to an entirely different species. They were then referred to Dr. Arthur, who considers them to be an undescribed subspecies of Ixodes pilosus (Arthur, ms.). Rageau (1953B) reported Tene a) man and from another kind of monkey, Cercopithecus (= Lasiopyga) cephus cephus. These represent an SS species re to Ixodes SchiTTin si (Arthur, ms.). BIOLOGY This species is a parasite of Colobus monkeys and uncommonly of man. It is said to prefer young hosts and attaches exclusively to the eye according to collecting notes furnished Neumann (1901). Females have been taken from around the eyes and ears and in the axillae of adult Colobus monkeys in Kenya and on the eyelid of a Colobus monkey in the Sudan. The male has been found only in copes on the host (and once alone on a tree trunk) and its feeding its are not known. Lar. vae, nymphs and both sexes of adults may occur on the same host (Arthur, ms.). - 559 = Extremely few ticks of any genus attack primates as hosts of predilection (Hoogstraal 1953E). Ixodes schillingsi appears to be one of these few. Monkeys, lemurs, and other primates usually groom themselves and each other so meticulously that external parasites have little chance of surviving on them. Certain exceptions should, however, be noted. When wishing to examine baboons for ectoparasites, one should choose large, lone males who wander separate from the group. Individual baboon hobos have yielded as many as 350 ticks (Rhipice- halus simus simus) while others living in groups in the same areas ave been free of ticks or have been infested by only one or two specimens. The striking rarity with which one observes easily— visible groups of Colobus monkeys grooming each other immediately suggests that because this practice is so infrequently indulged, I. schillingsi has managed to survive on this genus of monkeys But not among others. REMARKS The close relation of Ixodes schillingsi to Ixodes lunatus Neumann, 1907, a Madagascan parasite or er that also infests insectivores (Hoogstraal 1953E), has been described by Colas Belcour and Grenier (1942) who consider that these two ticks might be regarded as related subspecies. The features of the haller's organ of I. schillingsi are listed in a table by Arthur (1956B). IDENTIFICATION Males are easily recognized among the African Ixodes fauna by the following characters: Anal grooves racket-shaped (ante- riorly slightly wider than "horseshoe shaped"), not closed. Legs normal length, but fourth pair comparatively long, coxae with first three pairs bearing a small internal spur posterior- ly, coxa IV with a small external spur. Scutum with one lateral fold, convex, with many large, subequal punctations. Basis capituli with a sharp, pointed posteromedian spur ventrally. Measures about 2.3 mm. long and 1.4 m. wide. = 560 = Females have elliptical, open anal grooves. Coxa TI has a concave posterior border prolonged internally to a long spine and externally to a shorter spine. The scutum is longer than broad, broadest at anterior third; it lacks lateral grooves, has numerous deep, wniform punctations, and its cervical grooves diverge from the anterior third and are deeper posteriorly than they are anteriorly. The basis capituli ventrally has pointed, retrograde spurs (auriculae). Total length of gorged specimens may reach 7.0 mm. The nymph has been described and figured by Nuttall and Warburton 1). The larva is described by Arthur (ms.), who also redescribes the nymph and both sexes of adults. = 5oly= Figures 230 and 231, 9, dorsal and ventral views IXODES SIMPLEX SIMPLEX Kenya specimen PLATE LXVI - 562 — IXODES SIMPLEX SIMPLEX Neumann, 1906. (Figures 230 and 231) THE SHORTLEGGED BAT-RUSSET-TICK Tis Ne Orec! EQUATORIA PROVINCE RECORD n Katire Rhinolophus clivosus zambesiensis May (CNHM) Katire is at 3500 feet elevation in the Imatong Mountains. The tick noted above was sent for identification after having been re— moved at Chicago Natural History Museum from a host collected by Mr. John Owen. This is the only record of this species from the Sudan. DISTRIBUTION The subspecies simplex has a wide distribution throughout the warmer parts of the Old World. In Africa, it is thus far known only from the Sudan, Kenya Ys where the only other subspecies, africanus Arthur, 1956(A), also occurs 7, and the Union of South Africa. Elsewhere, it is found in the Near East, southern Europe, and Asia from Shanghai to Japan. A larger, closely related South African form has been noted by Arthur (1956A) as Ixodes sp. incer- tae. Africa L' CENTRAL AFRICA: BELGIAN CONGO. The subspecies mentioned by Bequaert (19305,1031) is not now certain. Records from French Equatorial Africa (Neumann 1906,1911, and Nuttall and Warburton 1911), refer actually to mterial from Kashmir or a neighboring country in southern Asia; cf. Arthur (1956A).7 EAST AFRICA: SUDAN (As I. simplex: Hoogstraal 1954B. As I. simplex simplex: Arthur 1956K). KENYA /Arthur 1956A. Note: I. simplex was listed by Love— ridge (19364) from Kenya; his specimens, from Mt. Elgon, are not = 502\— available for subspecies determination. Material from the crater of Mt. Menengai (HH legit) recently has been described by Arthur (1956A) as I. simplex Seay SOUTHERN AFRICA: UNION OF SOUTH AFRICA / Arthur (1956A). Note: According to Bedford (1932B), the record of I. pilosus howardi from a bat, reported by Howard (1908), may be = simplex, but the material is not now available and the subspecies or spe— cies cannot be determined. The specimens referred to as I. simplex by Zumpt (1950B) have not been reexamined. A closely related species or subspecies, of uncertain taxonomic status, is noted by Arthur (1956A) from the Irene caves near Pretoria /. Z2NORTH AFRICA: Specimen with this designation noted by Arthur (19564) 7 Near Kast PALESTINE (Arthur 1956A). Europe GREECE (Schulze 1937B. Pandazis 1947. Arthur 19564). FRANCE (Arthur 1956A). Asia "KASHMIR OR A NEIGHBORING COUNTRY" /Type locality according to Arthur (19564) 7. CHINA (Neumann 1906,1911. Arthur 1956A). JAPAN (Kishida 1930. Arthur 19564). HOSTS Bats known to be hosts of I. simplex simplex or of "I. simplex" are those of the 2 Seione hus , eas Rhinolophidae, the horseshoe bats; Miniopterus, eed bats (all pre viously unreported); and Myotis, mouse-eared bats. Both latter genera are in the family Vespertilionidae and both families are in the suborder Microchiroptera (insectivorous bats). See ( Rhinolo ae uinum (Neumann elige Rhinolophus eloquens Loveridge TOIGR ~ Rhinolophus clivosus auger (= geoffroyi auger apes 1950B). R. sp. of Howard aunts R. clivosus auger = geoffroyi auger*) according to Bedford (19328).” Rhinolophus clivosus zambesiensis (Sudan record above). Myotis (= Vespertilio) sp. (Neumann 1906). Myotis tricolor (neafont 19 stunder SOOM 328), rene caves material mentione er AFRICA above. Myotis macrodactylus (Arthur 1956A). Miniopterus natalensis arenarius and Miniopterus schreibersi (subsp. probably japoniae*) from Japan, M. schreibersi and M. Se schreibersi (Arthur NE BIOLOGY Aside from the fact that larvae, nymphs, and females are taken on bats, nothing is known concerning the biology of I. simplex. Males either do not take blood or feed very rapidly and ae ckly secrete themselves thereafter; they should be searched for in re- treats frequented by bats. Ixodes simplex is widely spread through the tropics and temperate climates of o: world and must be an un commonly adaptable tick. Its hosts! ability to fly undoubtedly accounts in part for the great range of this species. DISEASE RELATIONS Unknown. REMARKS The haller*s organ of both subspecies of I. simplex is like that of I. vespertilionis (Arthur 1956B). *I am indebted to C. C. Sanborn, Curator of Mammals at Chicago Natural History Museum, and an outstanding authority on bats, for checking the bat host names in this section. = 505 T= IDENTIFICATION Females are readily separated from those of the only other known bat-infesting species of this genus, I. vespertilionis by the fact that I. simplex has normal_length legs, ug. e last pair is longer than usual (all pairs of legs of I. vespertilionis are exceedingly long). Anal grooves are short and divergent. The scutum is slightly longer than broad, has gently curved postero. lateral margins, and converging anterolateral margins; widely scattered, subequal punctations, shallow cervical grooves, no lateral grooves; its color is brownish, reddish, or yellowish. The basis capituli is triangular, without cornua or auriculae. Coxae are flat and without spurs. For a fuller description, see Arthur (1956A). Males are unknown. Immature stages are described by Arthur (19564). =~ 566 — Figures 232 and 233, co, dorsal and ventral views Figures 234 and 235, 9, dorsal and ventral views IXODES VESPERTILIONIS English specimens. © loaned by british Museum (Natural History). 9 loaned by Dr. D.R. Arthur PLATE LXVII = 567 - IXODES VESPERTILIONIS Koch, 1844. (Figures 232 to 235) THE LONG-LEGGED BAT-RUSSET TICK EP ON = 40)" Cc. EQUATORIA PROVINCE RECORD al Torit Bat, unidentified Feb This is the only specimen of I. vespertilionis known from the Sudan. DISTRIBUTION I. vespertilionis is widely distributed in the Old World and is known from scattered areas in Africa where search will probably reveal numerous new locality records. The distribution of I. vespertilionis was first summarized by Nuttall and Warburton (1911) and Later, more extensively, by Neumann (1916). The present distributional summary is based on the latter paper, with only subsequent reports added. More recent— ly, Arthur (1956A) has brought these records up to date. Africa NORTH AFRICA: ALGERIA (Neumann 1916. Hirst 1916. Nuttall 1916.” Senevet 1937). MOROCCO (Arthur 1956A). EAST AFRICA: SUDAN (Hoogstraal 1954B. Arthur 1956A). UGANDA (Arthur 1956A). KENYA (HH collecting in crater of Mt. Menengai). SOUTHERN AFRICA: UNION OF SOUTH AFRICA (Arthur 1956A. See NOTE five paragraphs below. = 568 = Europe GIBRALTAR (Neumann 1916). PORTUGAL (Hirst 1916). SPAIN et 1916. Schulze 1927. Gil Collado 1936,1938,1948). FRANCE Neumann 1916. Hirst 1916. Jeannel 1926. Schulze 1927. Senevet™ 1937. Cooreman 1954A,B. Lamontellerie 1954. Arthur 19564). GER. MANY (Neumann 1916. Schulze 1923B,1944B. Schulze and Schlottke 1929). SWITZERLAND (Arthur 1956A). AUSTRIA (Neumann 1916. Nuttall 1916). BRITISH ISLES (Neumann 1916. Nuttall 1916. Hirst 1916. MacLeod 1939. Arthur 1948,1953A,1956A). BELGIUM (Bequaert 1913. Schmitz and Bequaert 1914. Leruth 1939B. Cooreman 1951). LUXEM. BURG (Leruth 19398). NETHERLANDS (van Eyndhoven 1939,1953). ITALY (Neumann 1916. Tonelli-Rondelli 1930A). SARDINIA (Kohls, correspondence). GREECE (Schulze 1936. Pandazis 1947. Arthur 1956A). HUNGARY (Neumann 1916. Hirst 1916. Kotlan 19214,B). CZECHOSLOVAKIA (Neumann 1910C. Rosicky 1953). BULGARIA (Schulze 1927). YUGOSLAVIA (Neumann 1916. Oswald 1939). ROMANIA (Leruth 1939A. Cooreman 1951). CRETE (Hirst 1916). NOTE: Schulze (1927) listed a nymph from Rhinolophus hipposiderus (sic) (?ferrum-equinum) at "Zelebor™ (7Burope). Near East TURKEY (Arthur 1956A). PALESTINE (Arthur 19564). IRAN (Olenev 1927,1931. Pomerantzev 1937,1950). RUSSIA (Olenev 1927, 1929,1931. Pomerantzev 1937,1950. Karpov and Popov 1944). Far East JAPAN (From Sawada, Myiagi, Honshu, A. J. Nicholson legit: Kohls, correspondence). NOTE: I. vespertilionis has been reported from Australia by Nuttall and Warburton (oT on ) and quoted by Neumann (1916), Ferguson (1925), and Leruth (19398), but not subsequently veri fied. The host was listed as Vesperugo tricolor. The only ves. pertilionid bat known to have the specific name tricolor is rious tricolor* of East and South Africa. The collecting locality for adie EEE EE EEE eee #I am indebted to C. C. Sanborn, Curator of Mammals at Chicago Natural History Museum, and an outstanding authority on bats, for checking the bat host names in this section. = 569 — the tick is Kingwilliamstown. There is a city of this name in Cape Province, South Africa but, so far as known, none in Aus. tralia. The specimen came from the Rothschild collection, a frequent source of South African material for Nuttall in the early 1900s. With little hesitation, therefore, this may be considered to be a South African record. Fielding (1926) re corded I. vespertilionis from North Queensland bats, but this appears to be merely a repetition of the earlier literature statement. Taylor and Murray (1946, p. 41) state that this spe cies is “doubtfully Australian" and the "original specimen is unfortunately lost". HOSTS All authors list bats except for the probably adventitious Hungarian record from a domestic dog (Kotlan 1921A,B). Immature stages and femles are usually found on bats; males only in caves inhabited by bats. The host of only one of the few specimens originating in Africa has been reported. Nuttallts (1916) Algerian specimen came from Pipistrellus (= Vespertilio) kuhlii. The host number of the specimen collected in Torit was inadvertently not included in the vial. The nymphs taken in Kenya (HH) were found, together with the holotype female of Ixodes simplex africanus, on Miniop- terus natalensis arenarius. So far as known, there are no other records of both I. vespertilionis and I. simplex subspp. froma single collection. “Myotis tricolor of South Africa has been dis— cussed in the NOTE SS Hipposideros caffer is also a host in South Africa (Arthur 1956A). re. Genera of European bats reported by Neumann (1916) are Rhino— lophus, Plecotus, Pipistrellus, and Myotis. The most commonly listed hosts are R. ferrum-equinum - hipposideros; the former species is also the host of the Japanese specimen reported by Kohls (correspondence). A few other Rhinolophus species are also mentioned by various authors. = 9/0%— BIOLOGY Judging from its considerable geographical range, this spe— cies is able to adjust to marked climatic variations only par- tially modified by protected cave environment. I. vespertilionis is rare on bats in Equatoria Province east of the Nile. Over a thousand bats, representing almost every spe- cies in Eastern Equatoria, have been carefully searched without finding more than the single specimen listed above. There has been little opportunity to examine carefully many caves. Males have been collected only from caves and other retreats in which bats assemble. No males have been found on bats. Nuttall and Warburton (1911) postulated that males may either feed very rapidly and then leave the host or that they may not feed at all Neumann (1916) believed that the various degrees of engorgement in which male specimens are found might not necessarily prove that males do feed but rather may be an indication of degree of nymphal feeding. This conclusion is based on the atrophy of the male hypostome in comparison with its robust development in fe males and in immature stages. Neumann (loc. cit.) mentioned the preponderance of numbers of males in relation to females and immature stages in collections and surmised that this may be due to the conspicuousness of the male's vagabond search for females. Females secrete themselves between stones of the caves to digest their blood meals. They probably oviposit in these niches, though this is not certain. Engorged nymphs are sometimes found in similar situations. When females are found on the host, immature stages are frequently found with them. Feeding is probably comparatively rapid, otherwise it is logical to assume that females and nymphs would have been more frequently reported from bats. Arthur's (1956A) comparison of data from Switzerland and from Macedonia leads him to believe that, because there is a reasonably high catch of partially and fully engorged ticks be- tween October and January and a number of unfed nymphs and fe males during the summer, feeding is accomplished mainly during - 571 - the winter months. This picture, possibly modified by the host's seasonal breeding cycle and activity, requires further observation. REMARKS The exceptionally long legs of this species is a character shared by many chiropteran parasites, notably the Streblidae and Nycteribidae (Diptera) and Argas boueti (cf. Figures 33 and 34). This feature is, however, not shared by all bat parasites, es. pecially those which are strongly appressed laterally, as fleas, or appressed dorsoventrally as bugs of the families Cimicidae and Polyctenidae. Except for Argas boueti, all the known chiropteran. infesting Argas species have normal-length legs, and indeed some, as for instance Argas transgariepinus White, 1846 (cf. Hoogstraal 19524), have comparatively short legs. Certain morphological peculiarities of adults and immature stages have been briefly mentioned by Arthur (1953A). The haller's organ is described by Arthur (1956B); it is like that of I. simplex subspp. Schulze (1938A, figure 28) has utilized this species to il. lustrate the thesis of morphological indicators due to pressure within the developing nympb. The subgeneric position of this species has been discussed by Neumann (1916), but this is moot; Arthur (1956A), the out— standing contemporary specialist on this genus, disregards it until further study can be undertaken. IDENTIFICATION Both sexes and the immature stages of I. vespertilionis are unique in the extreme elongation of the legs. The long anal grooves of both sexes are open; those of the male slightly con. verge posteriorly, but female anal grooves are parallel. The male scutum has a few large punctations in three rows and nu. merous fine, scattered punctations; the female scutum has nu. merous small, shallow punctations. = OIA = The larva and ae were partially illustrated and briefly described by Nutt Warburton (1911) but Arthur (1956A) provides complete descriptions of both sexes and of the immature stages. = Jia MARGAROPUS INTRODUCTION The genus Margaropus, closely related to Boophilus and com fined to Re seis consists of only ae Species, M. winthemi Karsch, 1879, of southern Africa and Madagascar, and Mi. reidi sp. nov. of the Sudan. Earlier assertions that M. wintheni is a South American tick apparently are erroneous. Usual remarks in the introductory sections for each genus treated in the present work are, in the case of Margaropus, in. corporated into the text below and do not require Sess: here. Illustrations of nymphal M. reidi sp. nov. and of M. winthemi, together with a review of the latter species, are given in the APPENDIX, pages 896 to 905. The unexpected circumstance of the very recent acquisition of the new species necessitates this treat. ment. KEY TO THE GENUS MARGAROPUS* MALES Six pairs of hair tufts and ventral hook on posterior body margin; a caudal pro. jection present when engorged. Adanal shields sharply pointed distally, acces— sory shields absent. Scutal outline convex laterally and bluntly rounded posteriorly. Free segments of leg IV as Wide as long. (South African winter Hor sertick) avs fdenecen toes co web dt ae ae ee wialvie’e sid oe eecleroclice WENT HEME Figures 359 and 300, 363 to 367 *The characters provided in the key, together with those in the generic key, are sufficient to comprise an adequate diagnosis for each species in this genus. - 574 - Dense row of scattered hairs on posterior body margin; hook lacking; caudal pro- jection not known to be present. Adanal shields bluntly pointed distally; acces. sory shields present. Scutal outline parallel laterally and gradually rounded posteriorly. Intermediate segments of leg IV slightly longer than wide. (Sudanesenei raffle tick) o's. das clos tecc.cccialeiccicice« dense. 0 ccMeeREIDI Figures 236 and 237 FEMALES Palpal segments 2 and 3 not separated by a slight constriction; basis capituli dorsally three times as wide as long; porose areas transversely oval. Scutum widest at level of eyes, rounded poste— TLOLIY cicicle » aatela/c¥eic o.oo. 0 o\a'sisiala|c\e o,01c olele's slele(icieisiaio.s o0je'o «sieMe: WIN HEMI Figures 361 and 362 368 to 372 Palpal segments 2 and 3 separated by a slight constriction; basis capituli dorsally twice as wide as long, porose areas vertically subtrianguler. Scutum widest midway between scapulae and eye level, bluntly pointed posteriorly........cccccccsccceeeeeeM. REIDI Figures 238 and 239 - 575 - Figures 236 and 237, @, dorsal and ventral views Figures 238 and 239, 9, dorsal and ventral views MARGAROPUS REIDI SP. NOV. Holotype and Allotype PLATE LXVIII = 5 7e.= MARGAROPUS REIDI SP. NOV. (Figures 236 to 239, and 342 to 358) THE SUDANESE BEADY-LEGGED TICK DISTRIBUTION IN THE SUDAN Bahr El Ghazal: All from Giraffa camelopardalis subsp., E. T. M. Reid legit: Liednhom (south bank o te it iver); lc, 1999, from muzzle, March 1955. One nymph, same locality, 8 March 1955. Twenty-four nymphs, Guar, Galual_Nyang Forest, 15 May 1953. DISTRIBUTION M. reidi sp. nov. is known only by the above specimens from the Sudan. It is related to the South African winter horse tick, M. winthemi, the range of which is charted by Theiler and Salisbury (1956). The small males may easily have been overlooked by ear lier collectors and the larger females may previously have been quickly identified as Boophilus by nonspecialists. More careful search should reveal the wider distribution of this interesting tick in the Sudan and possibly elsewhere in Africa. See M. winthemi (page 900). HOSTS The only known hosts are three specimens of the western Sw danese form of giraffe, Giraffa camelopardalis subsp., the actual subspecific identity of which is Sree by Setzer (1956) to be a moot subject that cannot presently be decided. See M. winthemi (page 900). BIOLOGY Unstudied. See M. winthemi (page 900). - 577 - DISEASE RELATIONS Unstudied. REMARKS M. winthemi and M. reidi sp. nov. are readily distinguished by key characters (pages 574 and 575). The HOLOTYPE co is deposited in the United States National Museum, number 2225. The ALLOTYPE 9 is deposited in the same institution, together with one of the PARATYPE nymphs collected on 15 May 1953. The data for these specimens are presented a. bove. A single &, 9, and nymph (15 May 1953) PARATYPE are depos- ited in the Onderstepoort collection (Dr. G. Theiler), East African Veterinary Research Organization (Miss J. B. Walker), British Mu seum (Natural History), Rocky Mountain Laboratory, and Chicago Natural History Museum. The remainder of the PARATYPE material is in the writer's collection. See M. winthemi (pages 899 to 905). DESCRIPTION MALE (Figures 236, 237, 342 to 346): Length overall approx. imately 3.0 mm., width approximately 1.4 mm. Color (dry) dark reddish brown, legs yellowish; integument white, yellowish, or dark brown. Outline oval with integument bulging beyond scutum laterally and posteriorly, but lacking caudal protrusion (avail. able specimens are moderately engorged). Capitulum; Basis capituli twice as wide as long; lateral margins markedly diverging anteriorly, basal margin slightly concave; dorsal surface bearing a horizontal row of twenty hairs at level of midlength; ventrally more elongate, length and width equal, lateral margins slightly concave, basal margin convex. Palpi comparatively short; overall length and width of each palpus approximately equal; segment 1 forming a short, wide pedicle for segment 2. Segment 2 subquadrate, approximately twice as wide as long; outer margin slightly expanded basally, inner margin = 51S = straight or slightly convex; anterior and posterior margins straight, parallel. Segment 3 triangular, outer margin approximately twice as long as inner margin. Palpi ventrally with segments 2 and 3 of al. most equal length, segmen earing a small triangular retrograde spur extending just beyond the apex of segment 2; segment 4 small, inserted in pit of segment 3; segment 1 forming a slight pedicle for palpi and bearing at its inner basal margin a slight knob with at least one bristle. Hypostome twice as long as wide, apical margin broadly rounded notched medially; a dense corona present; dentition 5/5, with nine to eleven denticles in each file, files of equal length. Scutum; Outline narrowly elongate with lateral margins par— allel, posterior margin bluntly rounded; lateral margins widening slightly over scapulae; anterior emargination deep. Posteromedian and paramedian grooves shallow, narrow, short, and indistinct, situated at level of spiracular plates. Cervical grooves shallow but distinctly divergent to lateral margin posterior of eyes. Eyes on lateral margin at level of coxa 2; small but distinct, rounded and slightly raised. Surface of scutum smooth and shiny, with a narrow median field of Tew scattered punctations extending the full length of the scutum; a single row of twelve hairs along posterior margin and a few hairs among posterior grooves; area between cervical grooves and lateral margins punctate and bearing scattered fine, long, white hairs; a row of hairs also situated on anterior margin between cervical grooves. iIntegument bulging around scutum from level of eyes posteriorly; Tenn shos with reg- ularly scattered, dense, long, white, conspicuous hairs bordering scutum to level of spiracular plates; thence these hairs more confined to narrow lateral surface of integument around posterior margin of scutum, few or no hairs on dorsal surface in this area. Spiracular plate subcircular, with one and a half rows of large goblets surrounding aperture. Genital aperture situated at midlevel of coxa II, anterior and posterior margins parallel, lateral margins slightly converging posteriorly. Genital grooves extending from genital aperture to level of anus, slightly teal gent. Adanal shields commencing at anterior level of coxa IV and extending beyond level of anus almost to posterior margin of body (depending on degree of integumental stretching due to feeding), narrowly elongate, slightly expanded just anterior of anus, - 579 - tapering and bluntly rounded distally; divided from each other by narrow area of integument; unattached to integument posterior of anus; surface bearing a few scattered hairs distally. Accessory shields extending from level of anus to approximately same Leve as apex of adanal shields, tapering, bluntly rounded apically; narrower than adanal shields. Hairs on ventral surface fairly dense and evenly distributed but shorter than those on lateral surfaces. Legs with free segments appearing beaded due to width and constriction between several segments; length-width ratio of seg- ments variable but width of none equalling length of same segment; free segments with numerous long, fine hairs dorsally and ventral. ly. Tarsi apically extended into a downward projecting, spurlike point; IV also with a short ventral subapical spur; outline nar- rowly elongate; claws and pads arising from dorsal surface at apex; claws long, narrow, recurved at some distance beyond pads. Coxa I narrowly, elongately subtriangular and extending anterior Ty almost to anterior margin of basis capituli, visible from dorsal view; posteriorly deeply cleft to form a widely triangular outer spur and a narrower, tapered inner spur; other coxae sub. rectangular with rounded margins; posterior margins slightly cleft; all coxae with numerous hairs. FEMALE (Figures 238, 239, 347 to 351): Unengorged specimens are very slightly larger than males; engorged specimens measure up to 6.2 mm. long but retain the linear, podshape appearance of boophilid females due to the severely parallel lateral margins of the body. Female characters recall those of the male but the leg segments are less expanded and the palpi are more elongate with a slight constriction between the second and third segments. Capitulum; Basis capituli from one and a half to twice as wide as long; basal margin moderately concave and joined to lateral margins by slight expansions suggestive of cornua; lateral margins concave to widest point at anterior third, thence recurved to anterior margin; porose areas small, shallow, and indistinct, subtriangular or pear shaped; bearing four or five hairs lateral. ly on dorsal surface. Palpi more elongate than those of mle, segments 2 and 3 separated by a pediclelike constriction and of 26560. approximately equal length; segment 2 arising from pedicle, with straight outer margin, convex inner margin, and subparallel ante_ rior and posterior margins; segment 3 with lateral margins slightly converging to bluntly rounded anterior margin, posterior margin straight though forming a slight, more or less downcurved projec_ tion at juncture with inner margin; segment 3 ventrally with short, broad spur reaching basal margin; segment 1 narrow, pediclelike, not observed to bear a ventral knob as in male; palpi with hairs as illustrated. Hypostome larger but otherwise similar to that of male. Scutum three-fourths as wide as long; outline slightly convex anterior of eyes, slightly converging from eyes to juncture of cervical grooves, abruptly converging posteriorly; anterior emar— gination deep. Cervical grooves reaching lateral margins as in male and delimiting an anterolateral area that is hirsute and somewhat punctate as in male. Surface otherwise smooth except for a transverse row of hairs bordering anterior margin between cervical grooves and a few scattered hairs arising from punctations in cen tral field. Eyes on lateral margins at scutal midlength, slightly convex. Spiracular plate of similar construction to that of male but differing slightly in outline. Genital apron a wide oval at level of coxae II. Genital grooves Seen to level of spiracular plates, thence divergent to level of posterior margin of anus. Inte nt with numerous fairly regular rows of short hairs on both eeeat and ventral surfaces; hairs more dense on posterior margin between spiracular plates. Legs similar to those of male except that the free segments are narrower and tarsi more elongate; tarsus IV lacking ventral subapical spur; anterior projection of coxa I variable, as dis. tinctly produced as that of male or more truncate. NYMPH (Figures 352 to 358): Capitulum. Basis capituli three times as wide as long, with straight basal margin, rounded junctures, and divergent lateral margins. Palpi four times as long as wide; segment 1 forming a slight ei: segments 2 and 3 of approximately equal length and subrectangular; apex OL yes more or less bluntly rounded; segment 3 ventrally with a short, wide spur not reaching basal margin of segment. Hypostome similar to that of adults but with smaller corona and 3/3 dentition in files of eight denticles. Scutum with lengthwidth ratio approximately equal; outline gradually diverging from scapulae to midlength, thence abruptly converging to narrow, bluntly rounded posterior margin. Surface lightly shagreened and with a few scattered hairs; cervical grooves as in adults (not visible in all specimens); eyes small, indistinct or invisible. Spiracular plate subcircular, with six large goblets in a circle. Tntegument with long hairs. Legs with length-width ratio of free segments intermediate between those of male and females sexes; coxa I not so deeply cleft and trumcate anteriorly; tarsi short, robust, not tapered downwards as in adults; dorsal margin gradually tapering; claws and pads as in adults. LARVA: Unknown. —poCe = RH IP ICEPHALUS INTRODUCTION It appears well established that continental Africa is the place of origin and center of distribution of Rhipicephalus ticks. Of the 46 rhipicephalid species and subspecies that aumpt (1950A) recognized in his preliminary generic revision, 39 (selatids the now cosmopolitan R. S. san ineus ) are endemic in the Ethiopian Faunal Region; two are Orien and five range from southern Europe and northern Africa into "Russia. The genus Oh cohen comprises almost a third of the known Sudan tick fauna contains more than twice as many endemic African species as any other ixodid genus in the Ethiopian Faunal Region. King (1926) listed five forms of Rhipicephalus from the Su. dan. In addition, R. capensis has been apparently erroneously referred to the Sudan 942B). During the present study, seventeen species and two B aidttionall subspecies, or nineteen different forms, have been discovered in the Sudan. Zumpt*s (1950A) major contribution in bringing together the basic taxonomic data for this difficult genus will facilitate greatly the final revision of rhipicephalid species. Earlier, Theiler (1947,1949B,1950A,B) had commenced study of individual species, an effort that is still underway (Theiler and Robinson 1953B, Theiler, Walker, and Wiley 1956). Very careful and thor. ough studies are also in progress by Walker (1956), who has several further reports completed or in an advanced stage of preparation. Since Zumpt's classic preliminary work, Wilson (1954) described a new species, R. hurti, from Kenya and Santos Dias has provided a number of new names, the validity of which are uncertain. Obviously, new criteria must be sought to sep. arate many rhipicephalid species and subspecies. In an attempt to apply characters surrounding the female genital aperture, Feldman-Muhsam (1952A) has distinguished another species, R. secundus, among populations appearing to be R. s. serie. However, problems of morphological and biological criteria for this genus are not likely to be solved until more exhaustive field research and patient laboratory investigation have been devoted to them. = Oot = Study of comparative morphology in the genus Rhipicephalus and of chaetotaxy in the family Ixodidae led Pomerantzev to reconstruct radially generic concepts of this family. This approach merits further investigation; however, the pitfalls of hasty conclusions based on worn or poorly—preserved field col. lected specimens, in which the chaetotaxic picture is imperfect, may result in additional confusion of species concepts (Hoog- straal 1955C). Rhipicephalids tend to considerable variation in appearance and morphological details owing to crowding on the host, welfare ‘of immature stages, and availability of suitable hosts, factors that play a part in the determination of size, robustness, and even certain physical characteristics. Distinguishing characters in many specimens tend to become so generalized that diagnosis is difficult. This is especially true for females. The question of biological races remains to be explored; many data suggest this phenomenon to be operable in certain groups of rhipicephalids. The genus is divided into clearly defined species and species variable enough to cause confusion. It contains extremely common as well as rare species. Host predilections within this genus are fairly wide among several groups of available animals, although the lack of interest in other animals easily available in the same area is conspicuous by rarity of records of their infestation. A few species, such as R. pravus, have an exceedingly wide host range, being common. ly taken on man and all domestic and many feral animals, such as carnivores, antelopes, hares, birds, elephant shrews, elephants, buffalos, and others. Other species, such as R. distinctus from hyraxes, are know only from a single kind of host. It is sig- nificant that immatures and adults of most rhipicephalid species do not attack birds and reptiles. The life cycle is either the two host or the three host type and hosts of immature stages may be either the same as those parasitized by adults or smaller and different animals. In some species, records of larvae from both cattle and rodents are so common as to confuse the picture of the preferred hosts of this stage. R. appendiculatus is an interesting example in point. In R. s. simus there is so much data indicating immature stage predilection for burrowing rodents that it is disconcerting to find that in certain areas where this tick is common larvae are =) 5Ole = frequently found on other animals. Reasons for these differences are beyond our present ability to explain. Immature stages of many rhipicephalid species remain un. described and distinguishing criteria for a number of those that are known are insufficient for identification of field collected material. Ecological stratification is quite restricted, various spe— cies being confined to forests, highlands, semidesert areas, or certain rainfall conditions. The degree and distribution of relative humidity appear to be the most critical of limiting factors. Vegetation types associated with this factor and im fluenced by the length of the rainy season or proximity to moisture laden air beside the seas can often be associated with rhipicephalid distribution. Economically, many species are of considerable importance as reservoirs and vectors of a variety of animal and some human pathogens. The kennel tick, R. s. sanguineus, has been shown to have a particularly wide spectrum of setaar or potential relation. ships as a vector of diseases. wnGS5 1. Re KEY TO SUDAN SPECIES OF RHIPICEPHALUS MALES Eyes convex or hemispherical, distinctly furrowed laterally or protruding from a depression ("orbited"). Coxa I with distinct; dorsal: projects One. cccicslccicciciale sis/siolcielneiee caisiaicalelcice Eyes flat or slightly rounded, not convex, hemispherical, furrowed, or orbited. Coxa I with or without dorsal PIFOGECTLON sercreielcieictcicic.c'clercvle cicle aie slaleisighe clots elavetalotecinie siaiote senile Eyes hemispherical, in a depression (orbited). Adanal shields large, enormously widened posterolaterally. Scutum dark with dense medium and large size punctations; color con trasting with reddish body integument and saffron legs. Frequently large (about 5.0 mm. long). (Common through MRIGN OF SCUAEM ) cease sieisie < c(eicinis «selene cictcioeattaoeien ape a ienon Figures 265 and 266 Eyes convex, with an encircling fur- row. Adanal shields mildly rounded laterally, not exceptionally large. Color overall brownish. Scutum with moderately numerous fine and medium size punctations. Fairly small (about 3.5 mm. long). (Southeastern Sudan)...............-R. PRAVUS Figures 285 and 250 Coxa I with distinctly pointed dorsal projection*, (Localized areas in southern LET ee cre ects cetera eave Sie acer Sn Cie ane CORSE Ee Coxa I without a distinctly pointed projection though a smaller, rounded hump may be visible in its place. (ioredormless widely distributed) «>see << siete aia cinsio me cite oe Su ppsont R. simus group),a small parasite of canerats only, typically bears this projection but this projection is so small and frequently so reduced that it is not considered distinct enough to include in this section. = 500 = 4. Lateral grooves faint, absent, or indicated only by a row of puncta tions. we arepspectes eae «Aster cisee cise eas closes cecane ane SD Lateral; grooves: GisStincG, aS, SUCH cccincccccccscecccccescsececd Scutal punctations in more or less definite rows of R. simus type, interstitials variable but always insignificant in comparison with PFLMANY. DUNCLALLONG s.c,01 sisieieee cuieicleeiice sce eee ceteccte ees suis ss p60 Scutal punctations scattered, not AN) OWS) LaLT ily, NUMOTOUS 0\eiteteleteleveletn eisialete ayetoroielerclete clelelercterere tote of Posteromedian and paramedian grooves absent. Adanal shields with inner margin in a straight line centrally and with a peculiar protrusion at juncture of inner and posterior MAL ZINS cesses e cece ccccceeccccseceecsceeceseeh. LONGICOXATUS Figures Posteromedian groove long, narrow, paramedian grooves shorter, wider, less well defined. Adanal shields with inner margin concave centrally and its juncture with posterior Margin TOUNCEd....csseecceececccseccccccccccceveeehs BEQUAERTI Figures 249 Basis capituli sharply angled lat_ erally and with long cornua. Scutal punctations large, unequal, unevenly distributed, small laterally, larger BN OTL OTN -Vis tai oicie ereioia’a sys eiajalsioieieleleisierejelerole's ste sveveteTe eleiclalslert of AttNOLL) L Figure Basis capituli slightly convex lat— erally and with brief cornua. Scutal punctations medium size, superficial, 2 dense, fine laterally.......cccccccccceccccecceceeshs MUHLENSI Figures 281 and 202 *R. distinctus atypically may have apparently shallow lateral grooves due to the size and depth of the row of large punctations in its bed. = 957 = 8. 10. Scutal punctations few, in four irregular rows (sims type). Posteromedian and paramedian grooves present but sometimes much MOGUC COaiafaieroielclaloreraravateralavelnvesciclelelolele elelcisloveioielelerelclercreteleleleieielcicieteleler? Scutal punctations scattered, not in TOWS oaictolcis cleroveile’cioree/erevere/elee eisieiele era \ojolole s/efs/etajaleieielows eisveicrotorerenlG Lateral grooves containing prom. inent, large punctations. Adanal shields with rounded or angular posterior marginal junctures but these not produced into spurlike WOO LTTE Soro iaielsiciele\s sie) « a'e o's ee e/nieielal sieleicisicicioqiemsiciciweie'siclt LO ENGL OD Figures 2 Lateral grooves without prominent punctations. Adanal shields with both posterior marginal junctures extended, the outer juncture spur— like, the inner rounded or spurlike; accessory shields distinct and DOLMGEA< cc'scc e's cies sissiesisieie s ojsie ce owls eelsiae cee scneltenl uote Figures 31 Scutal punctations moderate size, rather few and well spaced central— ly, even fewer or none laterally. Cervical fields moderately or norm reticulate. (Common only in Yei DiStriict) cv ciee cic sieves aoe atictareictn,cisto Nolsinrgorejose te JAE PEND ICURERUS Figures 240 to DL Scutal punctations larse, dense, partly confluent. Cervical fields markedly reticulate. (Always rare)............R. SUPERTRITUS Figures Ora aT. 12. Scutal punctations either excessively rare or in a pattern of four more or less linear rows (sometimes with one or two partial additional rows); among these interstitial or secondary puncta. tions are present or absent, if present they are smaller and more superficial*......ccccccccceccesele Scutal punctations not in these patterns, rather dense, not separated into primary and interstitial punctations but uniform or mixed. (Uncommon southern species)....cccessecsececeeeelO Scutum impunctate or with excessively few, shallow punctations. Adanal shields typically tending towards sickleshape but quite variable in series from single hosts. Postero~ median and paramedian grooves pres. ent or absent. Lateral grooves distinct or indicated only by a row of punctations. Coxa I with a small dorsal projection which may be mich reduced. (Small, rare, pearshaped, variable species, confined to cane BES) aie vine's cicie'e sisiels oie /ainiiaieisiatsie aamciancioineisisleisiciais laa eal ECON Figures 297 and 298 Scutum with more, larger, and deeper punctations. Other combinations of characters Git ROTIN ly a telslelelelolereicleiolcisielelelelele cleleiciclelelsicielolvcleletsicrall *[The linear punctation pattern of heavily punctate specimens of Resets ineus in southern Sudan may be somewhat obscured by dense, on large interstitial punctations, but can be discerned by turning the specimen obliquely to the source of the light. - 589 — 13. Posteromedian and paramedian grooves pronounced, deep, and wide*. Inter. stitial punctations varying from faint to large and numerous enough to some. what obscure basic pattern of four rows of larger punctations. (Common, widely distributed in Sudan)......cscceeccesceeseeeeRe S. SANGUINEUS Figures 259 and 200, 293 and 294. Posteromedian and paramedian grooves indistinct, shallow, or BUSOU cinnen es Ceo reise me ce sacle sian tay 14. Adanal shields elongately triangular with rounded marginal junctures. Only middle festoon protrudes. Posterior grooves absent, rarely very faintly indicated (but never distinct). Scutum arched; interstitial punctations usual. ly absent or insignificant, rarely more definite but never confusing basic pattern. (Common, widely distributed ireSuden) Sobcees.eaoteewee es carsee seencesceceeecls Se OlmlS Figures 301 and 302 Adanal shields either distinctly sickle. shaped or with peculiar inner or outer posterior protrusion. (More localized in southern Sudan and less common)..cscccccccccccccccscseseld A SS Da a ————————— *This character is constant in this variable species in all but a very few individuals obviously misformed in several characters. = 590.— 15. 16. 17. Adanal shields sickleshaped. Three middle festoons may protrude. Postero. median groove shallow but usually dis. cernible. Punctations slightly more dense bordering posteromedian groove. Interstitial punctations varying from absent to fairly numerous but not obscuring basic pattern. (Specimens may integrade with R. s. sims)...........-B. S. SENEGALENSIS Figures Adanal shields unusually wide, with rounded outer and posterior margins but with an elongated point at the juncture of the posterior and the con cave inner margin, this point reaches the festoons. Scutum exceptionally flat and broad....ccececccccccsccccccccsceseeeeehs CUSPIDATUS Figures 2 Lateral grooves replaced by a line of almost contiguous punctations. Pos— terior grooves merely faint, shagreened lines. Punctations numerous, close or contiguous, mixed large and small. (A mountain-inhabiting species).....ccccseccccescceesR. KOCHI Figures 269 and 270 Lateral grooves present. Other charac. ters 'V ELT OULS avavoiavere sve eloteioie eloteleye/elere clei elele aiprele ici s/elelw sicieislewveloteleee/ Posteromedian and paramedian grooves wide and deep (like those of R. s. a Adanal shields Typical. s y broadly tri ar (like those of R. 8S. ineus) but may be reduced To 2 Shee Punctations deep, dense, medium to large, some confluent; fewer laterally.....ccccccccccesseccceelhts SULCATUS Figures 309 and 310 Posteromedian and paramedian grooves long and very narrow or vague or GHSOMO LO ae era eon cero bic aeiclove dieieve aie sine Semis cleeniswecw esse lo - 591 - 18. Scutal punctations uniformly shallow, small or medium size, dense and close everywhere except in scapular areas, on festoons, outside lateral margins, and in narrow area just inside lateral margins. Posteromedian and paramedian grooves vague or obsolete. Adanal shields typically sickleshape (reduced in smell Specimens )s2scc.cede sewiaeees secewics cccceceneltc LONGUS Figures 277 and 278 Scutal punctations uniformly large, deep, dense but not contiguous in central area, variable in size in cervical area, fewer laterally. Posteromedian and paramedian grooves long and narrow. Adanal shields broadly triangular but with rounded margins, inner margin slightly con cave and more or less pointed at juncture with posterior margin, other junctures rounded..ccccccccccccccsccsecse eR. COMPOSITUS Figures = SES als FEMALES Eyes convex or hemispherical, distinctly OW GiaT Eau GYEOSSLON siscisisisic« «\a\¢ alnle «s\n o/0/c'eaials's'vic\es\ainle oF Byes flat or slightly rounded o. , Anaplasmosis: Gallsickness, Anaplasma marginale, of cattle is carried by the kennel tick. Redwater: R. s. sanguineus may be a vector of Babesia bigemina of cattle. Trypanosomiasis: The causative organism of surra, Trypanosoma evansl c ‘ls annamense), dies quickly in this tick, and aa of an African trypanosomiasis, T. congolense, is not transmitted by it. Another flagellate, Crithidia christophersi, has been reported to occur in the kennel tick. Rabies; Tompkins (1953) compares textbook statements that rabies 1s not transmissible by arthropods to saying that the aard- vark is immortal because nobody has seen a Gead one. He failed to become more rabid than this after serving as a host for a kennel tick that almost certainly had previously fed on a rabid fox. More than this nobody knows concerning rabies and ticks. Rodent Gestodes: The mouse tapeworm, Hymenolepis microstoma, is sald to be transmitted by this tick. Canine Filariasis: The canine filaria, Dipetalonema grassii, is sald to be transmitted by the kennel tick, which may also trans- mit Dirofilaria immitis and, questionably, Dipetalonema reconditun. So far as known, the role of R. s. sanguineus in the transmission of these parasites in nature has not been elucidated. Salmonella: In dogs and laboratory animals; see Dogs above. Experimental (Miscellaneous) Mistaken Identity: Cryptoplasma rhipicephali Chatton and Blane (1010A), grouped with the aemogregarines, Was indicated by the same authors (1916B) to be really the tick's spermatozoa. REMARKS Taxonomy In his generic revision of Rhipicephalus , Zumpt (1950A) considers R. sanguineus as a group 0 Subspecies, which, besides = Al the typical form, includes the subspecies sulcatus Neumann, 1908, of Africa, and both rossicus Yakimoff and Yakimoft, 1911, and schulzei Olenev, 1929, of Russia. R. sulcatus, now considered as a distinct species, is treated separately in the present study. The Soviets (Pomerantzev 1950) consider the Asiatic forms also as distinct species and add the following related species: R. turanicus Pomerantzev, 1940; R. pumilio Schulze, 1922; R. leporis Pomerantzev, 1946; and R. schulzei Olenev, 1929. ak Owing to the obvious difficulty of an independent evaluation of Soviet species, Zumpt's (loc. cit.) terminology is utilized in the present report. However, a4 survey of our own material from the Near East and ecological observations leave us, at present, uncertain over which of these two schools of thought is the cor rect one. More recently, Feldman_Muhsam (1952A) has designated as a very closely related species, R. secundus, distinguishable from R. s. Sanguineus only by the form of the female genital aperture and of the capitulum of larvae and nymphs. Although Feldman Muhsam!s reared material shows these differences, Mr. Kaiser and I have been unable to distinguish R. secundus after weeks of study of a very considerable number of kennel ticks from tropical Africa, North Africa, Arabia, and the Near East, Feldman-Muhsam (1953) did not recognize R. secundus in American specimens she studied, but claims to have found it among materials from Palestine, Turkey, Yugoslavia, France, Algeria, and French West Africa (and — un. published — Yemen and Egypt). Specimens identified as R. secundus by Feldman_Muhsam have been reported from Iraq along with R. s. sanguineus (Hubbard 1955). nai It appears obvious that full understanding and agreement of the status of subspecies and species related to R. sanguineus awaits more refined laboratory and field techniques than have yet been accorded this problem; possibly a more advanced con. Sideration of species criteria and of taxonomic tools than have yet been applied to ticks; and freer exchange of ideas and inter_ course in presently antagonistic regions of the world. It will be noted in the section on IDENTIFICATION, below, that an important diagnostic criterion for this tick is the met ace presence of a scutal pattern of punctations arranged in four more or less regular longitudinal rows. This characteristic defines the R. simus group and it is suggested (page 751) that R. s. sanguineus leus might logically be considered as a member of the same group. Structure and Physiology Integumentary sense organs, which are fixed in number and location, and which are essentially similar in all stages of the tick, though more primitive in larvae, have been described and silustrated by Binnik and Zumpt (1949). See also Lombardini 1950). The inte ent and sections of the scutum have been illus— trated by SORTS (1943B), who also mentioned the color of the gut contents. Water balance studies of various ticks, as discussed for 0. moubata (page 153), reported by Lees (1946A), include the obser- vation that R. s. sanguineus falls in about the middle of the range among the species studied with respect to its power of limiting evaporation. Variations in this capacity may reflect specific differences in the nature of the epicuticular lipoid among these species. See also Lees (1947). Teratolocical (malformed) specimens have been occasionally reported. Warburton ana Nuttall (19095) illustrated a Gold Coast specimen with duplication of the posterior parts of the body. Nuttall (1914A) described asymmetrical specimens and others lacking one leg. Sharif (1930) also noted absence of legs. Others have been described by Sharif (1930) and Pavlovsky (1940). These reports have been incorporated in an overall review of the subject by Schulze (1950B). Posteriorly joined adanal shields were illustrated by Santos Dias (1955A). A remarkable larva in which one of the palps appears to be partially converted into a typical leg has been described and illustrated by Pav- lovsky (1940) and reviewed by Campana (1947). A gynancromorph has been described by Pereira and de Castro (1945). It is somewhat interestins, in view of the considerable attention devoted to this subject and to teratological specimens in some schools, that no other such observations concerning this common and widely ranging tick have been reported. Tick feeding from tick: A male with its mouthparts inserted in the integument of an engorged female has been described by Sharif (1930). In Egypt we observe that males of various tick species insert their mouthparts into the female body cavity when the latter is engorged and both are confined in tubes for several days after having been collected. This is especially common among specimens of B. annulatus (= B. calcaratus).7 Comparative measurements ("allometrie") of sexual variations among the Ixodidae have been investigated by Chabaud and Choquet (1953). For the kennel tick, the length of the adanal shields and the spiracular plates in relation to the tick's length is logarithmically illustrated. Because of sexual and nutritional dimorphism, these authors consider ticks as important biometric tools. They also believe that certain cases of intersexual ticks may result from nutritional allometry. If a male structure is similar to that of a female it has an isometric growth, but if different an allometric growth. Variation in body size and morphology parallels that already discussed under R. appenerculatus (page a) and the same comments apply. This subject has been studied by Cunliffe (1914A) ana Pervomaisky (1954). See also remarks under IDENTIFICATION, below. Internal anatomy was briefly described and diagrammatically illustrated by Regendanz and Reichenow (1941). Haller's organ of this tick (= R. macropis) has been illus. trated by Schulze (1941). Oviposition and the larva have been described by Samson (1908). Tacluded in this report is a sketch of the female laying eggs, a generalized discussion of the subject, and a short des— cription of the morphology, including the internal organs (illus. - 719 - trated) of the larva. Spermatogenesis has been briefly described by Sharma (1943). Cytology in the kennel tick was studied by Stella (19383), who “examined the course of oogenesis and spermatogenesis ..... from the larva to the adult (stages) ...... The gonads assume their definitive aspect and begin to function only after the last nymphal casting. The maturation with expulsion of both polar bodies occurs inside the female in the ovary. The fertile sperna tozoa are formed only in the adult male and are of a unique type with flagellum and undulating membrane. The chromosomes are probably 46 in number, grouped in twelve tetrads". This paper is nicely illustrated and the reproductive organs of both sexes are described and illustrated. Cytoplasmic inclusions in the oogenesis of the kennel tick have been extensively reported by Das (1939), who correlated his own observations with a considerable amount of controversial conclusions by other workers. Chromosome studies have been briefly reported by Dutt (1952). Adult development within the nymph has been studied by Yalvac (1939). Excretory habits and mechanisms were investigated by Enigk and Grittner (1Se2- An "anatomical.physiological study of R. s. sanguineus" by Stella (1922 ears conclusions which I have iGarzlated freely as follows: ™(1) In the female, anatomical and histologic differentiation of most organs commences with digestion of blood; the gut develops completely and begins to function; the glands begin to activate secretion; the gonads enlarge and mature. The period of greatest functional activity corresponds nearly to the duration of the meal. Abandoning the host, the female begins oviposition, and during this period completely digests the engorged blood, thus its body, at first turgid and hard, becomes flaccid and soft. The malpighian tubules by this time are elaborating the substance SS 20-— of excretion, which fills the rectal ampule, and continue elimina. tion during deposition of eggs. The glands concerned with ovi- position become active the moment it commences; thus the accessory glands of the uterus secrete a substance to facilitate the passage of the eggs to the genital aperture; the gene's organ secretes a substance which cooperates in the formation of a gluey substance. The salivary and coxal glands, the function of which is associated strictly with the digestion of blood, work afterwards at the meal, reducing and degenerating it so that by the time the female has finished oviposition hardly any traces (of ingested blood) can be found." "(2) The young, unfed male, whose genital organs are small, shows indifference, but, differently from females, these organs mature and form sperms although the individual has not yet taken a blood meal. The gut utilizes the gross substance accumulated during the immature stage, and this is sufficient for mating, after which the individual dies. The salivary glands develop only slightly and do not secrete anticoagulin or toxin; the coxal glands do not become differentiated, and the malpighian tubules are small and in a state of repose.®™ "In those males that feed on blood at a certain time in their life, always after having first mated, an immediate arousing of all the physiological activities commences in those organs that, except for the gonads, have been in a torpor. The salivary glands begin functioning, as well as the coxal glands, and produce substances of agglutination and anticoagulin for the duration of the blood meal. The gut, which has completely absorbed the reserve substance, proceeds to digest the blood and the malpighian tubules send the excretory products which they have elaborated to the rectal ampule. Successively, sperm formation increases the secretory activity of the seminal vescile and of the appendage of the white gland so that a flood of viscous secretion for the dilution and nutriment of the sperm appears." "From the examination of the anatomical and physiological results it is evident that in the male and female of R. sanguineus there exist noticeable differences in the development and period of function of the various organs," etc. eels Miscellaneous Symbiotes of the kennel tick have been noted by Cowdry (1923) and Jaschke (1933). See also Muécrow (1932). Artificial feeding of this and other ticks by a capillary tube eee eer SeeLle é : : arrangement in order to accomplish physiological and pathogen transmission studies was described by Chabaud (1950). Whether host immunity is provoked by the feeding of this and other ticks has been investigated by Brumpt and Chabaud (1947) and Chabaud (1950B), who concluded that this phenomenon does not occur when the kennel tick feeds on dogs. Cuineapigs, however, react with an almost complete immunity and rabbits with a partial immunity (1950B). Infestation of guineapigs by R. s. sanguineus does not protect the host against Dermacentor pictus. "The Kind of host and kind of tick, as well as See methods, in. fluence experiments of this nature, and the earlier, classical results of Trager (1939A,B,1940) should not be generalized. IDENT IFICATION Males, though extremely variable in many morphological char. acters, retain a set of certain features that are constant and differentiate them easily. (1) Posteromedian and paramedian grooves are always present and distinct; and (2) no matter how variable the general scutal punctations may be, four more or less regular rows of widely spaced punctations, always larger and most commonly somewhat deeper than all others on the scutun, can be distinguished (when the specimen is turned obliquely to the source of the light) extending from the level of the para. median grooves to the level of the eyes. Males vary considerably in overall size, measuring from 1.7 mm. to 4.4 mm. long, or even more. Variation among most characters, except interstitial punctation, usually can be asso. ciated with size and robustness, though, uncommonly, a large or even enormous specimen is seen with weakly chitinized features normally associated with otherwise poorly developed, runty spec- imens. a A typical male, measuring some 3.0 mm. long, is pale yellow, brown, or reddish brown; has a pearshaped body; slightly convex scutum; deep lateral grooves and distinct festoons; a narrowly elongate or oval but medially expanded, deep posteromedian groove; shorter and broader but equally deep paramedian grooves; very slightly convex eyes; sharply angular basis capituli; elon gate, angular, adanal shields that in the posterior half of their length are usually distinctly widened and anteriorly are elongately subtriangular; and fairly distinct accessory shields. Scutal punctations always consist of four regular or irregular rows of fairly large more or less discrete punctations, among which a variable number of small to large interstitial punctations are indiscriminately scattered. If the specimen is engorged, the pearshape of the body is frequently exaggerated by integumental bulging laterally and posteriorly, from eye to eye, and the legs are successively larger from the anterior to the posterior pair; the posterior pairs are often massive and armed with a formidable tarsal hook ventrally. The single character that most frequently confuses identity of this species is the density and size of interstitial puncta tions. The general conception of this species is one of a tick with four more or less definite rows of larger punctations among which rather few and smaller insignificant punctations are pro- miscuously scattered. This form (Figure 289) is found throughout northern Sudan and Egypt, but in southern Sudan occurs, to the best of our knowledge, only on larger groundfeeding birds, such as bustards and storks. In typical southern Sudan specimens, interstitial punctations are larger, deeper, and more numerous (Figure 293). If it were not for the four rows of largest punctations, one would be tempted to refer to much of this material as R. sulcatus (as, indeed, some students of African and South American ticks are doing). In central Sudan, most specimens are fairly heavily punctate. (Although we have large files of notes on variation among specimens from throughout the world, a further discussion of this matter should await a thor- oughly exhaustive study). Spiracular plates in this species are surprisingly variable. These differences appear to be often correlated with nutrition eo) = (Cunliffe 19144) but other factors must also be involved since the plates of Egyptian average specimens are sometimes massive. In smaller, weaker, poorly nourished individuals, the color is either pale or darker than usual, the scutal grooves are less distinct, the adanal shields are more linear and have more rounded junctures (similar to those of R. a endiculatus), the basis capi- tuli is more linear, the legs are See and other characters are less distinctly pronounced. However, a definite trace of the critical characters remains in all except exceedingly few, obviously misformed specimens. Females are equally as variable as males. The above remarks concerning scutal punctations also apply to the female except that the larger punctations are less uniformly in rows but are scattered even more indefinitely over the central area of the scutum. In areas where males are lightly punctate, interstitial punctations of females are frequently more dense. The elongate scutum, nor- mally five-sixths as wide as long, narrows acutely posterior of the eyes; halfway from the posterior margin of the eyes to the posterior angle the margin normally is obtusely angled, and at the posterior point there is a minute marginal expansion. Though difficult lucidly to describe, this slightly undulating scutal pattern forms a most distinctive picture after one has examined large series of specimens. This outline may be more generalized in poorly developed indivicuals, and its length-width ratio may be more equal, but it is often maintained even in runts and weak individuals. The usually pronounced lateral grooves of the scutum are additionally picked out by being inset with large punctations; they extend about three-fourths of the scutal length; though in lightly punctate specimens the lateral grooves are often less definite. In the cervical areas, shagreening or coarse puncta- tions may occur and the scapulae usually have a group of large punctations. [Females of this species cannot be keyed in Zunpt's (1950A) revision, since the wrong section of couplet 21 applies./ The immature stages have been described by Cunliffe (19144), Theiler (19Z3B), Cooley (1946), Feldman-Muhsam (1952A), others noted in REMARKS, above, and in numerous other reports. NOTE: For further references to variation, related species and subspecies, etc., see REMARKS, above. Be (ae 299 ear 300 298 dorsal and ventral views C, Q@, dorsal and ventral views Figures 297 and 298, Figures 299 and 300, RHIPICEPHALUS SIMPSONT an Specimens PLATE LXXXIII SL RHIPICEPHALUS SIMPSONI Nuttall, 1910. (Figures 297 to 300) THE CANE-RAT GLOSSY TICK L N 9 @ EQUATORLA PROVINCE RECORDS al Torit Thryonomys gregorianus subsp. Feb 2 5 “ei marsh rat" Feb (SGC) These records, from the east and west banks of Equatoria Prov— ince, are the only ones of this species from the Sudan. The Yei specimens in Sudan Government Collections were collected in 1911 by H. H. King, who had identified them as R. simus. DISTRIBUTION R. simpsoni has been reported from widely scattered localities throughout tes within the Ethiopian Faunal Region and probably occurs wherever its favorite host, the cane rat, does. It is com mon in Kenya and Uganda although it has not previously been re- ported from Kenya. WEST AFRICA: NIGERIA (Nuttall 1910. Simpson 1912B. Rageau 1953B). CENTRAL AFRICA: CAMEROONS (Rageau 1953A,B). BELGIAN CONGO (Schwetz 192%. Bequaert 1931). EAST AFRICA: SUDAN (Hoogstraal 19548). KENYA (Common on lesser cane rats, Choeromys g- gre orianus, at Subukia, Nakuru District, 6400 ft. alt.; Hoogstra egit). UGANDA (Mettam 1935. Bedford 1936. Theiler 1947. Numerous specimens seen in collections of Uganda Veterinary Service. See HOSTS below). TANGANYIKA (Reichenow 1941B). SOUTHERN AFRICA: NYASALAND (Wilson 1950B). MOZAMBIQUE (Santos Dias 1952). UNION OF SOUTH AFRICA (Bedford 1936. Theiler 1947). See HOSTS R. simpsoni has been reported only from cane (or "edible") rats, Thryonomys (= Aulacodus) spp. and Choeromys spp. (all authors). What appears to be an exceptional host is the giant forest rat, Cricetomys gambianus; specimens from this host from Uganda, A. D. Fraser lenis are in British Museum (Natural History) collections. BIOLOGY R. simpsoni appears to be almost entirely restricted to cane rats. The paucity of records is possibly due to rarity of host examination. Present evidence would indicate that this is one of the most host-specific of rhipicephalid ticks. Although R. suryson is closely related to R. sims, this latter species is seldom found on cane rats. DISEASE RELATIONS Unstudied. REMARKS Adults were figured and redescribed by Theiler (1947). The nymph was described by Santos Dias (1952G), who, incidentally, considered this species as a synonym of R. sims lon but who subsequently (1952G,H) recognized the obvious validity of R. simpsoni. Generally one may be suspicious that records of R. sims from cane rats actually refer to R. simpsoni, although two males of R. simus from a cane rat near Yirol as been seen, (E. T. M. Reid legit). All other collections, labelled as R. sims from cane oS that have been studied by the writer from various parts of Africa have proven to be R. simpsoni. = 127 = IDENT IFICATION Male: Size is usually small, from 2.3 mm. to 3.6 mm. long and from 1.5 mm. to 2.2 mm. wide. Typical males are easily dis- tinguished within the R. sims group by a combination of characters including broadly sickleshaped adanal shields; sparse, almost ob. solete, shallow, scutal punctations; distinct lateral and posterior grooves; short, converging cervical pits; pearshaped body; and slight dorsal process of coxa If. The several fairly large series of specimens in the present collection and other East African series that have been studied show considerable variation among individuals, though always one or more of the distinctive features of the species are retained. The posterior grooves may be very faint. In small, weak spec- jmens the lateral grooves may be more shallow than usual or they may be indicated by only a row of punctations. In specimens with exceedingly small adanal shields, the characteristic sickleshape is frequently reduced. The pointed dorsal projection of coxa I is very small and in a number of specimens it is reduced to merely a blunt hump. Female: This sex averages 3.4 mn. long and 2.0 mm. wide; it ranges from 2.5 mm. to 3.4 mm. long and from 1.5 mm. to 2.5 mn. wide. The scutum is about one-fourth longer than wide; its poste— rior margin is sinuous, with a slight medial protrusion; eyes are pale and flat; lateral grooves almost reach the posterior scutal margin; cervical grooves extend posteriorly for about half the scutal length. Scutal punctations are sparse and superficial; interstitial punctations are usually absent, but some fine ones may be present with a few larger punctations in lateral grooves. The great length of the scutum in relation to its width distin- guishes females from those of R. s. simus, R. s. sene alensis, and R. bequaerti. This character is an especially important one in specimens in which interstitial punctation is more apparent than is common for R. simpsoni. =) (20 = Figures 301 and 302, @, dorsal and ventral views Figures 303 and 304, g, dorsal and ventral views RHIPICEPHALUS SIMUS SIMUS an Specimens PLATE LXXXIV = f29) = RHIPICEPHALUS SIMUS SIMUS Koch, 1844. (Figures 301 to 304) THE GLOSSY TICK* L N Q ou EQUATORIA PROVINCE RECORDS 1 Torit MAN (feeding on) a Torit MAN (feeding on) 22 Torit MAN ee on) 2 2 Torit MAN (crawling on) t 3 Tort MAN (crawling on) 1 Torit MAN (feeding on) 1 Juba MAN (feeding on) 2 Juba MAN (crawling on) il Gilo Lemniscomys striatus massaicus a al Torit Lemniscomys striatus massaicus i! 2 2 3 Torit Burrows ~ L. striatus massaicus Mh Torit Lemniscomys macculus macculus 2 Torit Tatera benvenuta benvenuta ils) Torit Tatera venuta benvenuta ib fee 5 orit Burrows of T. b. benvenuta 2 Torit Mastomys natalensis 1smailliae Al. Tkoto Mastomys natalensis 1smalliae al Lotti Forest Praomys tullbergi sudanensis 5 Tkoto Ervicanthis niloticus jebelae iE Torit Arvicanthis niloticus isbelse LoVe 6 47 Flori Burrows of A. niloticus Jebelae 30 19 18 Torit Burrows of A. niloticus Spolas 1 31 28 Juba Burrows of A. niloticus yobelas Ik Ikoto Lepus capensis crawshayi 1 Torit Lepus sietoricc MLcrovis 9 Torit Panthera Leo Leo To = Torit Canis aureus soudanicus i Sl soricc Canis aureus soudanicus 6° °°7 “Torit Canis aureus soudanicus 2°73 — Torit Canis aureus ‘soudanicus (2) (2) *Also known as the black pitted_tick, the glossy brown tick, and the tailtuft brown tick (cf. Theiler 1952A,B). - 730 = NY or PCs SS * Torit Crocuta crocuta fortis Oct S26) Torit Mellivora capensis abyssinica Jan 2 2 Obbo Civettictis Saatte congica Apr iho Zo Atoreaisy, Civettictis civetta confica Feb 8 7 Lugurren Phacochoerus aethiopicus bufo Jan 4 5 Torit = scrofa sennarensis Apr 1 1 Nimle 1ppopotamus hibius amphibius May 1 Ae? Nimulle Hi otamis amphibius hibius Oct (SVS) i WGevelska a sohamieneperiom tiene = + (sce) 1 Kidepo aurotragus oryx pattersonianus Jan 1 Torit Taurotracus oryx attersonianus Feb 1 3% Kapoeta Hippotragus equinus bakeri Dec 4 Torit a r equinus bakeri Jan il Ikoto Rh a= guentheri smithii Dec 1 19 Boma Plains Syncerus ee aequinoctialis Dec Zee eae LOD Caffer aequinoctialis Jan 2 Holo Caffer aequinoctialis Maer i Lolianga catter aequinoctialis Jan 5 2 Kapoeta dogs Dec 8 14 Keyala domestic dogs Dec 225025 lorit domestic dogs Dec (3) G6 6 Torit domestic dog Jan 1 Torit domestic dog Mar 1 Torit domestic dog Apr Lee ereelorat domestic dog Jun dy Lorit domestic dog Nov Bey PyKatire domestic dog Oct 5 Loronyo domestic dog Jan 8 8 Juba domestic dog Jan i S*- i suba: domestic dog Dec eS 2 Juba domestic dog May (SGC) AL Nimule domestic dog Mar evel. Miado domestic dog Sep (SGC) 67 69 Kajo Kaji domestic dogs Dec (3) 65 2) Torit domestic pigs Mar *With 1 oR. sims senegalensis. This is the only collection seen from anywhere in Africa in which typical specimens of both subspecies have been found on a single host. - 731 = te =a +O ise) oy SES 3 O 44 io, 1 de al 2 ee 6 7 32 af TSS 6 17 2 L-'-"3 3 il ue ae Torit Torit Torit Torit Torit Katire Lolepori Loronyo Juba Juba Juba Loronyo Juba Juba Kajo Kaji Torit domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic domestic on grass Imatong Mts. ? host (8700 ft. alt.) pigs Apr pigs Nov (2) pigs Dec (4) pigs Jan cattle Dec cattle Oct cattle Dec cattle Jan cattle Jan cattle Dec horse Jan goat Jan sheep Jan sheep Dec (SGC) sheep Dec Aug (Weber 1943) DISTRIBUTION IN THE SUDAN King (1926) stated that R. s. sims occurs throughout Equatoria, Bahr El Ghazal, Upper Nile, Blue Nile, Kordofan, and Kassala Prov- inces and that it had at that time only recently established itself in Northern Province where proper conditions of humidity for its development prevailed around newly installed pumps and basin irri_ King apparently had no Darfur Province records for this gation. species. Sudan localities from which specimens of the glossy tick have been seen are: Bahr El Ghazal; Galual_Nyang Forest (hyena, black.legged mongoose, leopard, Lion, buffalo, and warthog; domestic dogs; old male baboons infested with as many as two hundred adult specimens; orsing on men; SVS, HH). aw, 200 Te Sty. Lau (cattle; PN Ere = Jur River (hippopotams; SVS). “Fanjak (domestic dogs and cattle; SVS, HH). Wau (domestic pigs; SVS). Kenisa (numerous specimens from elephants; SGC). Several localities near Yirol (elephants; SVS). Yirol (hyena and cane rat; SVS). Upper Nile: Duk Fadiat (wild pig and warthog; SVS). Akobo Post tH on; Sac). Maban (cattle and goats; SVS). Pariak (cattle; SVS). Kaka (roan antelope; SGC). Bor (domestic dogs; HH, leopard; SGC). Malakal (domestic dogs; HH). Blue Nile: Roseires (cattle; SGC). Darfur: Zalingei (camels and horses; SVS). Kulme, Wadi Oribo (fox; SMNH). Kordofan: Tabanga (pigs; SGC). Talodi (cattle and pony; SGC). "Western Jebels" (cattle; svs). / Khartoum No records.7 Northern: Letti Basin (fox; SGC). Shendi (bull; SVS, sheep; SGC). These populations are probably quite restricted. DISTRIBUTION R. s. simus ranges throughout the Ethiopian Faunal Region. In West Africa, it is more or less widely replaced by the sub. species senegalensis. The Arabian range of the glossy tick, as mapped by the American Geographical Society (1954), should be limited to the mountains of the Yemen. {WEST AFRICA: Early records should be checked against R. simus senegalensis. The range of R. s. simus in West Africa Is im perfe SSS own. NIGERIA (Simpson 1912A¥,B. Unsworth 1949,1952. Mettam 1950). SIERRA LEONE (Neumann 1901,1911. Simpson 1913). TO GO (Neumann 1901,1911. Ziemann 1909). GOLD COAST (Simpson 1914. Beal 1920. Stewart 1934). IVORY COAST (Neumann 1901). PORTUGESE GUINEA (Tendeiro 1946A,B,1948,1951A,E,1952C ,D,E,1954: wherever *Specimens referred to as R. simus by Simpson (1912A, p. 325) are actually R. sims senegalensis. They were determined as R. simus falcatus by Nuttall a Warburton. These specimens are in British Museum (Natural History) collections. - 733 - illustrated the specimens appear to resemble the subspecies senegal. ensis rather than simus). FRENCH WEST AFRICA (Neumann 1911. Rousse. Tot 1953B. See also Sudan below. Theiler states (correspondence) that her extensive Senegal collections are all typical R. s. simus). NOTE: Tonelli-Rondellits (1938) reports from Sierra Leone, Togo, and Gold Coast are probably repetitions from Neumann .7 CENTRAL AFRICA: CAMEROONS (Neumann 1902A. Zumpt 1943A. Rageau 1951,1955A,B). FRENCH EQUATORIAL AFRICA (Rousselot 1951, 1953B). BELGIAN CONGO and RUANDA-URUNDI (Newstead, Dutton, and Todd 1907. Massey 1908. Nuttall and Warburton 1916. Roubaud and Van Saceghem 1916. Schwetz 1927C,1932. Bequaert 1930A,B, 1931. Zumpt 1943A. Bouvier 1945. Wanson, Richard, and Toubac 1947. Schoenaers 1951A,B. Rousselot 1953B. Theiler and Robin son 1954. Van Vaerenbergh 1954). EAST AFRICA: SUDAN / The synonymous R. shipleyi from "Soudan" (Neumann 1902A) refers either to the Sudan or to French West Africa, King (1908,1911,1926). Zumpt (1943A). Weber (1948). Hoogstraal (1954B)«/ ETHIOPIA (As R. simus, R. hilgerti, and R. erlangeri: Neumann 1902A,B,1911,1913,1922. Stella 19384, b39a,Bs19%5- Bost Roetti 1939. Zumpt 1943A. Charters 1946. D'Ignazio and Mira 1949). ERITREA (Stella 1939A,1940). FRENCH SOMALILAND (Hoogstraal 1953D). ITALIAN SOMALILAND (Paoli 1916. Franchini 1926,1927,192%,E. Veneroni 1928. Niro 1935. Stella 1938A,1939A,1949). BRITISH SOMALILAND (Drake-Brockman 1913B. Stella 1938A,1939A,1949). KENYA (As R. perpulcher and R. praetextatus; Gerstacker 1873. As "R. perpulcher from Mombas, Zanzibar™; Neumann 1904. As R. ecinctus; Neumann 1908,1922. Neumann 1911,1912,1913,1922. Neave 1912. Loveridge 1923D. Anderson 1924A,B. Lewis 1931A,B,C, 1932A ,B,1934,1943,19590. BrasseyEdwards 1932. Walker 1932. Daubney 1933,1934,1936B. Daubney and Hudson 1934. Kauntze 1934. Roberts 1935. Fotheringham and Lewis 1937. Mulligan 1938. "Kenya Vet. Serv." 1939A,B,1940 ,1947,1949,1951,1952. Lewis and Fotheringham 1941. Zumpt 1943A. Lewis, Piercy, and Wiley 1946. Dick and Lewis 1947. Weber 1948. Beaumont 1949. Heisch 1950B. Binns 1951,1952. van Someren 1951. Wilson 1953. Wiley 1953. Lumsden 1955). - 734 - UGANDA (A. Theiler 1910A. Bruce et al 1911. Neave 1912. Neumann 1922. Richardson 1930. Mettam 1932. Carmichael 1934. Wilson 1948B,C,1950C. Lucas 1954). TANGANYIKA (Neumann 1901,1907C,1910B,1911. Morstatt 1913. Loveridge 19234. Bequaert 1930A. Allen and Loveridge 1933. Moreau 1933. Evans 1935. Cornell 1936. Zumpt 1943A. J. B. Walker, unpublished, see various parts of HOSTS section below). SOUTHERN AFRICA: ANGOLA (Manetti 1920. Sousa Dias 1950. Santos Dias L950c). MOZAMBIQUE (Howard 1908,1911, larval iden tification in 1908 paper open to question. As R. ecinctus: Howard 1909B. Sant'tAnna 1911. Theiler 1943B. Zumpt 19Z3A. Santos Dias 1947B,1952D ,H,1953C ,19544,H,1955A). NORTHERN RHODESIA (Neave 1912. Le Roux 1934,1937,1947. Matthysse 1954. Theiler and Robinson 1954). SOUTHER! RHODESIA (Robertson 1904B. Edmonds and Bevan 1914. Bevan 1920. Jack 1921,1928,1937,1942. Lawrence 1942). NYASALAND (01d 1909. Neave 1912. De Meza 1918A. Davey and Newstead 1921. Zumpt 1943A. Wilson 1943,1945,1946,1950B). BECHUANALAND (Theiler, unpublished). SOUTHWEST AFRICA (Trommsdorff 1913,1914. See immature HOSTS below). UNION OF SOUTH AFRICA (Koch 1844. Neumann 1901,1912. Lounsbury 19034, 1904A,B,1905B,19060. Robertson 1904. A. Theiler and Stockman 1904. A. Theiler 1905B,1909B ,1911B,1912A,1921. Howard 1908. Galli-Valerio 1909B. Speiser 1909. A. Theiler and Christy 1910. Donitz 1910B. Moore 1912. Van Saceghem 1914. Breijer 1915. Bedford 1920,1926,1929,1932,1934,1936. Cowdry 1925B,C, 1926A,1927. R. du Toit, Graf, and Bekker 1941. Curson 1928. Cooley 1934. Bedford and Graf 1935,1939. R. du Toit 1942B,C, 1947. Zumpt 1943A. Zumpt and Glajchen 1950. Meeser 1952). OUTLYING ISLANDS: ZANZIBAR (Neave 1912. Aders 1917). SEYCHELLES (Desai 1952). MADAGASCAR: Neumann (1901,1911). Poisson (1927). Tonelli- Rondelli (1938). Buck (1948A) indicated that this species (""Haemaphysalis sims") is not established on Madagascar. Hoogstraal 535) ./ = 135 = ARABIA: YEMEN (Sanborn and Hoogstraal 1953. Hoogstraal, ms.). NOTE: The known distribution of this species in Arabia on the map of the American Geographical Society (1954) should be limited to the mountains of the Yemen. MISCELLANEOUS: Tonelli-Rondelli (1938) noted that Stella (1938D) had reported R. simus from ITALY and indicated that this report is certainly an error in identification; also that this species is absent from LIBYA / reported as present by Stella (19380) 7, ALGERIA, TUNISIA, and MOROCCO. Records for Mytilene, GREECE (Senevet 1920), repeated without further substantiation by Pandazis (1947), and records for TURKEY (Stefko 1917) are probably also erroneous or refer to introduced, non-established specimens. Neumann (1911) listed R. simus from EGYPT and Brumpt (1920) stated that East Coast fever exists in Egypt and might be carried by this tick. Mason (1922B) quoted this as an erroneous statement that R. simus occured in Egypt, to which Brumpt (1923, p. 43, footnote) replied that he had merely hypothesized this possibility on the basis of Neumann's record. Carpano (1936) reported micro- organisms in specimens of this tick (?imported or misidentified — HH) collected from carnivores in the Cairo zoological gardens. Actually, the glossy tick is not established in Egypt, as com firmed earlier by Mason (1922B). Records from TURKESTAN (Yakimov and Kohl-Yakimov 1911, Yakimov 1917,1922,1923) are based on material now considered as subspecies of R. sanguineus (cf. page 717). The "R. simus or R. sanguineus" ae = beers ~ _ from tortoises in IRAN (Michael 1899) probably refers to H. aegyptiun. Data from BORNEO (Neumann 1901) probably refers to R. sanguineus subsp. or to R. haemaphysaloides subsp. Christophers (1907C) reported R. sims from southern INDIA but, as Sharif (1928) says, this too is probably a misidentifica. tion. Patton's (1910) remarks concerning the transmission of Piroplasma gibsoni of India by a new species of tick related to R. simus have been elaborated in subsequent reviews to indicate that R. sims transmits this organism. =) 26 HOSTS Adult hosts of predilection are carnivores, pigs, buffalo, and other large or medium size game animals. Antelopes are usually second.choice hosts. Among domestic animals, dogs and pigs fre— quently are preferred. The incidence on cattle varies locally and may be either very high or very low even where the glossy tick is common. People are frequently attacked, especially in the vicinity of their dwellings. Although Matthysse (1954) con. siders adults to be parasites of medium size mammals, overall data indicate that host size is only one factor, the type of host being an equally important consideration. Larvae and nymphs feed chiefly on burrowing rodents, less commonly on other small animals. Adults Cattle: Factors influencing parasitism of domestic cattle by R. s. simus are still unknown. Although this tick is common throughout squatoria Province, its incidence on cattle is nil or low everywhere except in Juba District where the rate may run fairly high. In Central Sudan, the numbers on cattle are variable but never high. R. s. sims is one of the five species of this genus that occurs with any degree of frequency on Uganda cattle but in most districts the rate of infestation is low (Wilson 1950). In South Africa, the glossy tick "does not appear to thrive well on cattle" (Lounsbury 1904B). Theiler (correspondence) , however, reports that in cattle raising areas of South Africa these animals are the favorite host of the glossy tick. Wilson (1950B) considered this to be an uncommon Nyasaland tick because he took it on only 24 occasions during an extensive three year tick survey. Ten of these collections were from cattle. Earlier, Wilson (1946) doubted that females ever become fully engorged on cattle. Theiler (correspondence) has found that the glossy tick is more common in Nyasaland than Wilson believed. Rarity of cattle infestation here probably accounts for this discrepancy inasmuch as relatively few wild animals were examined. Matthysse (1954) considers this to be a rather important Northern Rhodesian cattle parasite, but does not provide a clear cut anal. ysis of the situation there. = Ee Low incidence of R. s. simus on cattle is not universal. It is not only common but numerous on cattle on the coastal plains of Kenya (Dick and Lewis 1947). Dogs: Throughout the range of R. s. simus, domestic dogs are frequently among its most common hosts and are mentioned by many authors. In Equatoria Province, dogs are infested by this parasite as frequently as they are by R. s. sanguineus, but the numbers of sims are considerably lower in all collections except those from Kajo Kaji. The yellow dog-tick, H. 1. leachii, is mich less common on dogs in the Sudan but in certain other areas of Africa this is one of their most important arthropod parasites. Kauntze (1934) and Roberts (1935) did not consider i. s. simus to be an important pest of dogs when studying bouton- neuse fever in Kenya, but Dick and Lewis (1947) found this common tick to be only slightly less numerous on dogs than R. s. sangui- neus in the coastal area of Kenya. At Nelspruit, South Africa, Lounsbury (1904A) collected more glossy ticks than kennel ticks on canines in the same place. According to Theiler (correspondence), R. s. simus and H. 1. leachii generally are more common on South Ifricen dogs than is R. s. sanguineus, and H. 1. leachii is usual. ly the most common of the three. “Matthysse (1954) Listed R. s. sims only once from Northern Rhodesian dogs, and noted R. appen diculatus, R. S. Sanguineus, and k. tricuspis more frequently. Pigs: Wherever I have encountered domestic pigs in the range of the glossy tick, a close association between the two and a high rate of infestation has been noted. No mention of similar situations is found in the literature. Wild warthogs and bushpigs (see below) are commonly infested. In South Africa, on the other hand, domestic pigs are amazingly free of ticks and more are recorded from bushpigs than from warthogs (Theiler, correspondence). Tendeiro's reports from Portugese Guinea (listed above) contain numerous references to R. s. simus and R. s. senegalensis on domes. tic pigs. The identity of West African specimens should be checked. Other domestic animals are more or less frequently reported as hosts, but the incidence is seldom if ever mentioned. In addition to the Sudan records given above, some of these are: Cat (Jack 1921,1942). = 190 = Horses (Neumann 1901,1902B,1911, Howard 1908, Massy 1908, Lewis LO31A, Bedford 1932B). Goats (Howard 1908, Lewis 1931A, Bedford 1932B, Wilson 1950B). (Howard 1908, Lewis 1931A,C ,1932A,1934, Daubney and Hudson 1934, Wilson 1950B). Camels (Neumann 1902B,1911). Chickens (Lucas 1954). Man More specimens of R. s. simus were taken on members of our parties in southern Sudan Than any other tick species. Several of these were engorging and others might have done so if they had not been removed. Galli-Valerio (19098), Jack (1942), and Lumsden (1955) noted incidental attacks on man. Veneroni (192 and Zumpt and Glajchen (1950) reported human paralysis after bites of the glossy tick. Others have said specifically that no spec- imens were found on people during surveys, for example, Kauntze's (1934) report on the Kilmani area near Nairobi. J. B. Walker (correspondence), however, reported five males and three females from man in tall grass country south of Arusha, Tanganyika. Wild Animals Primates: In Bahr El Ghazal, hundreds of specimens were collected from old male baboons, some of which harbored as many as 200 glossy ticks, chiefly in the axilla. In the same areas, baboons travelling in community groups were consistently free of ticks. In Kenya, we found specimens on Colobus monkeys (see also I. schillingsi, page 558), but, except for a note by Lumsden (1955), there is no reference in ene literature to similar col- lections. Theiler (correspondence) has a single record from Galago crassicaudatus agisymbanus from Zanzibar. Carnivores: These animals, along with some of the larger and thicker skinned herbivores of Africa, appear to be hosts of choice: Lions / All extensive African collections studied by the writer contain specimens from lions. Reported by Neumann - 739 . (1901 ,1907C ,1910B,1911,1922), Howard (1908), King (1926), Bedford (19328), Tewis (19328, 1934), Wilson (1946B,1950B), Weber (1948), Santos Dias (1952H, 1953¢), and others 7 Leopards (Loveridge 1923A, King 1926, Bequaert 1930A, Lewis oe Zumpt 1943A, Wilson 1950B. Numerous specimens in BMNH collections). Cheetah (Lewis 1932A,1934). Serval (Lewis 1931B,1932A). Various foxes (King 1926, Weber 1925, J. B. Walker, anpubliahed)& Various hyenas (Neumann 1922, Loveridge 1923A, Lewis 1934, Zumpt TOZ3A, J. B. Walker, unpublished). Various jackals (Neumann 1902B, Lewis 1931A,1932A,1934, Zumpt 19234). Various civets (Loveridge 1923A, Bequaert 1930A, Allen and Loveridge 1933, Zumpt 19434, Matthyase 1954). Genet (Bequaert 1930A). Hunting do 5 Lycaon pictus (Howard 1908, Van Saceghem 1914, Lewis I931A, Be B; J. B. Walker, unpublished). Ratel (Sudan records Aaa Marsh mongoose (Loveridge 1923A). The Onderstepoort collection, BM(NA), and the present collections contain numerous other specimens from these and similar carnivores. Antelopes: In Equatoria, the only records of this tick from antelopes are from two common elands, a tiang, a large—snouted dik.dik, and two roan antelopes. None others were found on the several hundred antelopes examined in Equatoria and Bahr EL Ghazal Provinces. Elsewhere in the Sudan, there is only a single record from a roan antelope at Kaka in Upper Nile Province; this was reported by King (1926). King also noted the gazelle as a host, but specimens are not now available. Simpson's (1914) records for a West African oribi and a reed buck quite possibly pertain to the subspecies senegalensis. Other published records from antelopes are: Duiker (Bedford 1932B). Grant's gazelle (Lewis 1934). Kudu (Bedford 1932, Lewis 1932A). Bushbuck (Lewis 1931A,1932A). Eland (Lewis 19314, ”1932A). Wildbeest (Lewis 1932A,1934). Sable antelope (Santos Dias *1950). Impala (Meeser 1952). Steinbuck (Lewis 1931B). On 49 Tanganyika Thomson's gazelles that yielded many ticks, R. s. simus was represented by only two females although this Subspecies was common on many other kinds of hosts from the same area. (J. B. Walker, unpublished). A single male was found on an eland. Hehe) = In Wilson's (1950B) list of nine kinds of hosts of this tick in Nyasaland, no antelopes are represented, and none were included among Howard's (1908) eleven kinds of hosts from southern Africa. The Onderstepoort collection contains only seven collections of glossy ticks from various antelopes throughout Africa (Theiler, correspondence). In addition to the kinds mentioned above, these are a topi from Uganda, a gemsbok from Southwest Africa, and an oribi from Southern Rhodesia. Other hosts: Elephants / In Equatoria Province these animals are either normally attacked by A. tholloni or are free of ticks; but in Bahr El Ghazal Province all elephants examined were heavily infected by R. s. simus and none harbored A. tholloni. Numerous specimens of R. s. simus have been examined in various collections from elephants in other parts of Africa. Others have been reported by Mettam (1932) 7. Buffalo / These animals are usually infested in the Sudan and frequently harbor numerous specimens of R. Ss. simus. Elsewhere they are also important hosts of this tick. Records have been published by Neumann (19070 ,1910B,1911,1922), Davey and Newstead (1921), Bequaert (1930A,1931), Lewis (1931B,C, 1932A,1943), Mettam (19325, and Santos Dias (1952D) 7. Rhinoceroses The black, or narrow-lipped rhinoceros is hardly an important ost because it is seldom numerous in nature, but where it occurs most individuals appear to be infested by the glossy tick. See Neumann (1922), Lewis (1932A), Zumpt (19434), J. B. Walker (un published). The excessively rare white, or square-lipped rhino_ ceros has been reported as a host by Breijer (1915) and Zumpt (19434); these_reports apply, however, only to the southern race/. Hi tamus / Specimens were found on the ears of three specimens examined in the Sudan; three other specimens were free of ticks. No other records are available 7. Pigs / The warthog and bushpig are frequently infested in the Sudan and elsewhere. See Neumann (1901,1907C ,1910B,1911), Howard (1908), Bequaert (1930A,1931), Lewis (1931B ,C ,1932A), Bedford (1932), Mettam (1932), Weber (1948), Wilson (19508) » and J. B. Walker CE lee Zebras / Reported by Neumann (1907 ,1910B,1911), Lewis (1931B,1932A), Mettam (1932), and Santos Dias (1952D) 7. Antbear or aardvark, Oryctero afer Neumann (1922), Bedford (1932), Lewis (1932A), Wilson (19Z6B, 950B), Matthysse (1954) 7. African porcupines J Neumann (1907C , 1910B,1911), Lewis (1932A), Matthysse 5 0. B. Walker (un. published) 7. Pangolin /Mettam (1932) 7. - Al. f Giraffe: Inasmuch as adults commonly parasitize so many large game animals, infestation of giraffes is to be anticipated. However, no published records have been found indicating that this is so. In Bahr El Ghazal Province, where this is a common tick and where numerous giraffes were examined, R. s. sims was not found on these animals.7 nie Exceptional hosts: Hedgehogs (Howard 1908, Van Saceghem 1914, aes 19328). Hares (Loveridge 1923A, Bedford 1932B, Lewis 1932A). In the laboratory, Lewis (1932A) found that adults feed readily on hares (see also immature stages below). Rabbit (Howard 1908). Cane rats (See also next paragraph. Sudan records include a single collection from Bahr El Ghazal and the Onderste— poort aetna contains three lots from Southern Rhodesia). Rodents (South African Otomys, Aethomys, and Rhabdomys; four records: Theiler, correspondence). ?Mistaken identity: Specimens from a cane rat (Bedford 1932B) were later found to be R. simpsoni and not R. simus (Bedford 1934). Neumann's (1922), Mettam's CIs32) and Loveridge'ts (1923A) specimens from cane rats should be checked against R. simpsoni. See HOSTS of R. simpsoni, page "Large gray cattle tick (?Rhipicephalus sinus Koch) (sic) attached to and completely blocking up the ear opening" of a lizard, Mabuia striata (Loveridge 1923D). This sounds like an Aponomma tick. Note Adults, found frequently in rodent nests in which the immature stages feed, are always newly molted, unmated, unfed individuals biding their time before venturing forth to find a host more at- tractive to them than rodents. Larval and Nymphal Hosts Rodent hosts: All of the numerous records of adults in the Onderstepoort collection are from larger size animals and most records of immature stages are from small, nest-inhabiting mammals (Theiler, correspondence). Our field experience in East Africa - 72 - and Arabia follows the same pattern. The details have been best worked out by Roberts (1935) (also noted by Kauntze 1934) who found the immature stages in the Nairobi area in nests of the following rodents: grass rats or kusu, Arvicanthis sp.3; groove—toothed rat, Otomys sp.; striped grass mouse, Lemniscomys sp.; four-striped age yee oars pumilio; and muLtimammate rat, Mastomys = Mus) coucha. In the Sudan, some larvae and many nymphs were taken on the same types of animals mentioned by Roberts and also on gerbils, Tatera b. benvenuta. Engorging specimens were taken on these animals trapped away from their nest. We also took a single nymph from two different kinds of hares in two localities but it seems that hares should be considered as exceptional hosts for immature stages. In Northern Rhodesia, Matthysse (1954) also reports larvae from hares. British Museum (Natural History) collections contain a nymph from another probably exceptional host, a ground squirrel Fermos— ciurus sp., collected by Karl Jordan in Southwest Africa. A break. down of collections of immature stages from South Africa in the Onderstepoort collection (Theiler, correspondence) shows that nine are from hares, two are from springhaas, five from hyraxes, one from ground Bee can 125 from to), deaiae genera of murid rodents: Aethomys (56), Rhabdomys (24), Lemniscomys, Mastomys, Thallomys, Arvicanthis, Rattus, and Mus; and forty from the ee cricetid rodents: Otomys GED); Myotomys, Parotomys, Tatera and Gerbillus. Other reports of immature stage hosts are short-haired rats, meee sp. for larvae (Bedford 1932B) and Praomys jacksoni for nymphs during the dry season around Leopoldville (Wanson, Richard, and Toubac 1947), and an elephant shrew for a nymph "probably of this species" in Kenya (Lumsden 1955). Exceptional hosts of immature stages in the Onderstepoort collection (Theiler, unpublished) are three elephant shrews and one Crocidura shrew (insectivores) , one hartebeest, one mongoose, and one mierkat. Also included are three birds from the region of the Sabi-Lundi Function, Southern Rhodesia, the puff—back shrike, Dryoscopus cubla, the blue-breasted waxbill, Uraeginthus - 743 - angolensis, and the dusky-faced warbler, Tricholais scotops. It is difficult to determine whether so many kinds of Smeal hosts are the result of wider and more extensive field search or of ecological differences (see BIOLOGY below). Laboratory hosts: Lewis (1932A) observed that although nymphs and aaits fed readily on hares, larvae were more reluctant to do so. From this Lewis concluded that hares are less preferred hosts of larvae than they are of other stages, an inference that probably should be modified by other biological considerations. At Onderstepoort, both larvae and nymphs feed readily on guinea. pigs (Theiler, correspondence). Questionable remarks and conclusions: Lewis (1932A), after examining Roberts’ specimens from rodent burrows, concluded that larvae feed less readily on rodents than do nymphs. Supporting data were not provided but, unless we are still unaware of some unique phase in the life cycle of this tick, it appears that in East Africa, at least, larvae and nymphs attack the same host. Data from collections made throughout the entire year will be necessary before other conclusions can be drawn. Lewis (1932A) noted a number of larger mammals, from the size of porcupines to rhinoceros, as nymphal hosts, and Stella (1939B) indicated guineafowl. Reidentification of pertinent material is indicated. The Onderstepoort collection (Theiler, correspondence) contains a single collection of nymphs from a red hartebeest in Natal. Lounsbury (1904A) "wholly failed in attempts to rear larvae on dogs, not one of many thousands applied having fed to reple- tion", but later (19060) succeeded in doing so. The ox was considered an unsuitable host for larvae but "nymphs and adults do not appear to dislike cattle". Around Lourenco Marques, Sant'Anna (1911) noted, larvae of R. Ss. sims are encountered and so is a human disease, possibly Ehat now called boutonneuse fever, following tick bites. Sub sequent reviewers have elaborated this remark to indicate that larval simus bite man, and the mre enthusiastic have quoted this report as stating that the larvae transmit boutonneuse fever. - 744 - BIOLOGY Life Cycle Laboratory studies indicate that R. s. simus undergoes a three. host type of life cycle. Under experimental conditions, Theiler (unpublished), Lewis (1932A), and Lounsbury (1905) have found the life cycle periods to be as follows: PERIOD DAYS OC. TEMPERATURE Theiler Lewis Lounsbury Lewis Oviposition to hatching 20. 24 Bilt 21.27 Larval prefeeding period 3 7 Larva feeds 2 5 3} 2 Premolting period &12 11 6 23. 26 Nymphal prefeeding period aA il Nymph feeds se ibt 5 3 Premlting period ABS 21% 27 14 21.25 Adult prefeeding period t t/ Female feeds Tm 2h, dé 9 Preoviposition period BEs6 6 23—26 eo abbh Note: Theiler rearings at 24°C. to 26°C. and approximately 80% R.H. Hares were used as hosts for all stages in Lewis! experiments, though not with great success. The ease with which these observa. tions could be repeated using normal hosts and normal conditions of temperature and humidity found in rodent burrows suggests an interesting study for collecting comparative data. Field observations indicate that both immature stages feed on nest-inhabiting rodents. The nymphal-adult molt occurs in the same nest, and adults remain in the nest for some time before seeking larger hosts. As discussed below, a considerable amount *2 months in winter. - 745 - of additional field study is required to answer many questions concerning the life cycle of R. s. sims. This tick and H. leachii are the only ubiquitous African species that in their immature and adult stages, respectively, feed first on nest— inhabiting rodents and then on larger animals. More rigid field observations, besides being a most pleasurable occupa. tion, should be easily accomplished. In the following paragraph the question of where the female oviposits is raised. In this connection, Howard's (1909B) ob. servation of an unengorged male and female of R. simus (= R. ecinctus) mating on the leaf of an Acacia thorn tree on the Zambesi River is of special interest (although the possibility of misidentification of these specimens must be considered). Almost invariably, rhipicephalid ticks mate on the host, as do most other ixodids. Howard's note indicates the necessity of further research to determine whether R. s. simus possesses a unique type of mating and egg laying, and, if’ so, whether this is a constant or an exceptional phenomenon, and whether it is associated with an ability of larvae to seek out their pre— ferred habitat and hosts rather than waiting for a passing rodent. Mating observed by J. B. Walker (correspondence) in Kenya has been of the ordinary rhipicephalid type, on the host. Walker also says that engorged females that have already dropped from the host will mate with males that have been feeding on the same host and then removed and placed with these females. This might be merely a mating act and not initial or essential fertiliza. tion (HH), for generally, it appears, female engorgement is not complete or normal unless copulation has been effected. Ecology From Roberts! (1935) studies in the Nairobi area it appears that the immature stages of R. s. simus prefer slightly subsurface rodent nests rather than deeper nests of the same and of other kinds of rodents. The grass rat, Arvicanthis, is possibly the most important immature stage host. It 1s not known whether larvae actually seek out the nest and attack the animal there, or whether they attach to a rodent wandering in search of food and are then carried to any nest that their host might be ine Seiko habiting. It would be contrary to all previous observations on ixodids to assume that the female selects the situation in which she oviposits, although this possibility must be investigated. By way of comparison, immature stages of H. leachii in the Nairobi area prefer deeper nests of Mastomys coucha. In the Njoro area of Kenya and in the Sudan, however, we have taken larvae and nymphs of these two ticks species in both shallow and deeper nests. Arvicanthis nests are usually within a foot of the surface of the ground, but sometimes they are two or three times as deep. The nest is reached by a network of a few or many tun. nels, each with a small exit among vegetation. The round nests, composed of lesser or greater amounts of moist grass and leaves, appear to be occupied for several generations. Slight rises, such as borrow heaps, mounds beneath bushes, or pathsides are favorite burrowing sites. These rodents frequently nest and search for food in close association with human activities. Habitats of some of the chief South African hosts of im mature stages (see HOSTS above) differ widely. Aethomys nama. uensis frequents rock crevices and piles of stones ae A. chrysophi lus is more terrestrial and lives in sheltered bush on the plains, among rocks, or in burrows under bushes or rocks. Otomys lives in holes in the ground or in selfmade shelters in matted vegetation; those in the Karroo construct these shelters from large piles of weeds while others utilize small grass or weed nests in marshes or among rocks. Rhabdomys hides in holes in the ground and its pathways run through dense vegetation. It is evident, therefore, that the usual ecological niches of hosts of immature stages in East Africa differ from those in southern Africa. In southern Africa, Theiler (unpublished) finds that the glossy tick occurs from the eastern tall grass veld (Port Eliza. beth) northwards through subtropical overgreen and deciduous tree and thorn forest into northern Transvaal, Southern Rhodesia, and Mozambique. In these subtropical stretches, the heavy rain. fall areas of Natal are comparable with the coastal plains of Kenya, and the dry, warm conditions in Kruger National Park and northern Transvaal are comparable with central Kenya (see next paragraph). Records indicate this tick to be less common west = 747 = of the Drakensberg escarpment, but to occur up to 10,000 feet elL evation in Basutoland. It is common in Highveld with good annual rains, heavy frost, and snow in winter but seldom recorded from dry Highveld. In the mixed grass veld of the middleveld, with ten to 25 inches of rainfall annually and cold, sharp winter frosts, it is almost entirely absent, though it does occur in southern Transvaal middleveld. It is numerous in the Bankenveld and Limpopo highlands and also in Bushveld regions of Transvaal. In Karroo areas it dies out in areas with less than ten inches of rainfall annually. Records from Southwest Africa are only from the northern, more moist areas. In Southern Rhodesia, it is especially common in eastern and northern areas. In the hot, more or less humid, coastal lowlands of Kenya, R. Ss. simus is especially common (Wiley 1953). Dick and Lewis (192.7) consider this to be the most abundant and widely distri- buted tick in the Kenya coastal lowlands and Wilson (1953) also notes its frequency there in areas where R. pravus and A. gemma are found (cf. pages 681 and 274). In the arid Northern Prov— ince of Kenya, the glossy tick is less common than elsewhere in the Colony, but it does occur anywhere under a variety of condi- tions, whether these be hot and arid, cold, damp high altitudes, or hot, moist coastal lowlands (Wiley 1953). Theiler (1943B), supported by subsequent remarks by Santos Dias, noted that R. s. simus is not only particularly abundant but actually the Tost ubiquitous tick in some parts of Mozambique. Study of data for the Somalilands, in the coastal areas just north of Kenya, suggests that R. s. simus is common only under local conditions in these less humid areas. From details published by all investigators concerning the Belgian Congo it would appear that in most parts of the colony this tick is decidedly less common than it is in southern Sudan. In the Sudan, R. s. simus is common everywhere in the south and at least frequent in Southcentral areas. It becomes more localized and uncommon with the approach of semidesert conditions. Many areas in which it occurs have a long, severe dry season but rainfall of twenty to almost fifty inches annually during the wet season. King (1926) noted that although this tick is absent in the desert areas of Northern Province, populations had estab. lished themselves there when local conditions of humidity were modified after pump and basin irrigation was introduced. lO Altitudinal distribution in Kenya is from sea level to 11,000 feet (Lewis 1932A). Lewis (1931C) considered this species as one relatively little affected by conditions influenced by altitude. In RuandaUrundi, the glossy tick ranges to about 5600 feet al. titude (Schoenaers 1951B). Neumann's (1907C,19108) Tanganyika records include specimens from altitudes up to about 6000 feet. In Arabia this tick is known only from the temperate, watered 3500 foot to 7000 foot range of the Yemen mountains but not from drier and hotter lowlands (Hoogstraal, ms.). The seasonal cycle of R. s. simus is not well known. During the colder months of the year, Wilson (1946) found no adults in Northern Province, Nyasaland but his collections were mostly from cattle, which are not common hosts there. Engorged females were found during the wet season (Wilson 1950B). In the Sudan newly molted adults were collected in rodent nests during the dry sea son and engorged adults on larger hosts chiefly during the dry season. In Northern Rhodesia, Matthysse (1954) found adults on cattle chiefly during the rainy season, but some also in dry periods. The emergence of adults and the length of time they normally remain in their host's nest at different seasons of the year should be investigated. On cattle, adults of R. s. sims are usually found on the tail switch, feet, and anus (Theiler 1943, Wilson 1948B, Matthysse 1954). On does they feed almost anywhere, as noted by Theiler (1943B) and Wiley (1953). Our Sudan specimens from hippopotamus were all found on the hosts! ears. Neumann's thirty specimens from a Kenyan rhinoceros were taken from the host's inguinal area. Adults awaiting a host show "a marked predilection for tall grass overhanging paths, but are common everywhere", including in human habitations, where they were associated with boutonneuse fever by Dick and Lewis (1947). They are also found in houses in Somaliland (Drake-Brockman 1913B, Veneroni 1928), where they bite children and cause paralysis. The close association between the glossy tick and man is easily understandable. Adults are common on domestic dogs and less frequently, under local conditions, on cattle. The immature stages feed on rodents that frequently nest under vegetation bordering garden plots and cultivated fields, beside roads and paths, near buildings, or in the vicinity of streams where people congregate. Many of these rodent hosts - 749 - are also attracted to man's agricultural activities and the adult ticks frequent tall grass in search of hosts. All these factors bring man and tick closer together. When dragging for ticks in Nyanza Province of Kenya, Lewis (1931C ) obtained adult glossy ticks only in shady woodlands while the other species, R. appendiculatus, A. variegatum, and H. leachii were obtained from open country with grass and bushes. In Moreauts (1933) study of the food of the tickbird, Buphagus erythrorhynchus (Stanley), in Tanganyika, fifteen specimens = R. S. simus were found in the stomachs of four of 58 birds examined. enya, van Someren (1951) examining the stomachs of twelve of the same kind of bird found four females in two of them. He found seven adults in three out of seven stomachs of B. a. africanus. DISEASE RELATIONS MAN: Boutonneuse fever (Rickettsia conorii). Paralysis (?toxin). CATTLE: East Coast fever (Theileria pa rva). A uae ee (Anaplasma marginale). Redwater (Babesia srme _ PIGS : Piroplasmosis (Babesia trautmanni). SHEEP: Not a vector of Nairobi sheep disease (virus). {CARNIVORES: Statements that R. simus may transmit Babesia sibsoni among jackals and dogs and By felis” among pumas are based on errors in tick identification and on errors in quoting original reports. REMARKS Misshapen specimens have been reported by Santos Dias (19478, 19558). In the latter paper, a gynandromorph is also described and illustrated. Symbiotes have been reported by Cowdry (1925C,1926A,1927) and reviewed by Jaschke (1933). Dinnik and Zumpt (1949) reported on larval integumentary sense organs in relation to those of nymphs and adults as well as of other species. The "R. simus group" of Zumpt (1942A) consists of the species R. sims with two subspecies Sims and senegalensis, and of several Telated species, R. longicoxatus, R R. lunulatus, R. tricuspis, R. distinctus, Kk. p. planus, and Rk. p. com Tanatus, R. simpsoni, and R. re1chenow é: fee =R. zump i Santos Dias 13505). These are ‘all Ethiopian Faunal Region species. Zumpt also included the Indo- Malayan species R. ih. haemaphysaloides and R. h. pilans, but I then. hesitate so to consider The outstanding diagnostic criterion of the R. simus group is a scutal pattern of few large, fairly deep punctations arranged in about four more or less regular, individually characteristic, long- itudinal rows. Among these, interstitial punctations are usually inconspicuous. Adanal shields of males of each species are dis— tinctive. Female scutal patterns may be more variable and those of certain closely related species or subspecies may be difficult to differentiate. Actually, R. s. sanguineus conforms to this group criterion and might well be consiaered aS a member of the R. sims group. It would then also be much easier to associate the Indo-Malayan species, mentioned above, with this group. Theiler (correspond ence) writes that she agrees with this view. Santos Dias (1%2C) has considered this group differently. IDENT IF IC AT ION Males: This sex is easily recognized and extremely few spec— imens ever cause doubt as to their identity. The group criterion of four more or less regular rows of rare scutal punctations is always definite, but their distribution and depth are somewhat variable. Although interstitial punctations are usually faint or obsolete, a few specimens show them mre distinctly, though = (LS never enough to confuse the picture. Posteromedian and paramedian grooves are absent; rarely they are just barely discernible, but never pronounced. This latter fact is an important distinction between R. s. sims and those specimens of R. s. sanguineus that otherwise closely approximate R. s. sims in appearance. The scutum is arched and usually shiny black, less commonly reddish brown or reddish yellow, with definite, deep lateral grooves and clear festoons. The central festoon may protrude with engorge— ment. A hump of coxa I projecting dorsally frequently is large enough to resemble a pointed projection, but closer inspection indicates its bluntness. Ventrally, the shape of the adanal shields, with their rounded external and internal margins, is usually characteristic; but in some specimens, in which they are abnormally narrow and with a deeply concave inner margin, they approach the sickleshape of the subspecies senegalensis. Size is extremely variable, running from minute (1.9 mm. Long - rarely) to well over 5.0 m. long. Female: The outline of the black (rarely reddish) scutum is subeircular with a broadly rounded, slightly sinuous posterior margin; the length.width ratio is approximately equal or slightly wider than long. Punctations are normally rare but may be some. what heavier and more distinct than in the male. Interstitial punctations are rare and inconspicuous, or absent; rarely they are somewhat pronounced and slightly confuse the typical picture of this species. Lateral grooves are of variable distinctness and length; frequently they are extended posteriorly by a few punctations; in some individuals they are much reduced. The converging and thence diverging cervical grooves usually clearly extend to or almost to the posterior margin, but may be obsolete on the posterior third of the scutum. Lateral fields usually contain two or three punctations; eyes are flat. Size is variable with tremendously engorged individuals reaching 15.0 mm. in length. Most females are easily keyed but those with reduced lateral grooves are apt to be confusing. Females of the subspecies sims and senegalensis are quite similar. The nymph and larva have been described and illustrated by Theiler op) = TERR = Figures 305 and 306, <, dorsal and ventral views Figures 307 and 308, 9, dorsal and ventral views RHIPICEPHALUS SIMUS SENEGALENSIS an Specimens PLATE LXXXV - 753 - RHIPICEPHALUS SIMUS SENEGALENSIS Koch, 1844. (Figures 305 to 308) THE SCIMITAR-SHIELDED GLOSSY TICK et Zz +0 Qa EQUATORIA PROVINCE RECORDS 5 5 Nimle Lycaon pictus somalicus Jul (SVS) 1 Kapoeta* Hippotragus equinus eri Dec i 17, Boma: yneerus caffer aequinoctialis Dec Plains** 2 Holo Syncerus caffer aequinoctialis Mar 3 Mongalla Guncerus caffer aequinoctialis -— (SGC ) 2. Kheirallah Smeerus caffer aequinoctialis Mar (scc ) 1 lLugurren frecectecacs aethiopicus bufo Jan 10 15 #£Sunat Phacochoerus aethiopicus bufo Apr 3 4, Kheirallah Phacochoerus aethiopicus bufo ~ (SC) 2 Bundle domestic dog Mar (SGC) (Lado) 4 4&4 Magwe on grass May (SVS) 4 4 Mvolo on grass =. (ScC) 1 3 Mvolo on grass Jul (Svs) Specimens in the Sudan Government collection noted above were collected by H. H. King in 1910 end 1911. They had been identified as R. sims and as R. falcatus. DISTRIBUTION IN THE SUDAN Bahr El Ghazal: 16’, lo from buffalo, Guar, Galual_Nyang Forest, © June 1953, P. J. Henshaw legit. Io, and 5a and 18 99 *Collected with 36% and lo R. sims sims. This is the only col lection seen from anywhere In Africa in which typical specimens of both subspecies have been found on a single host. The host was obviously a migrant. %*For comment, see BIOLOGY below. - 754 = from two warthogs, same locality as above, 7 June 1953, and 8 January 1954, collected by E. T. M. Reid and P. Blasdale. Nw merous other specimens collected in the same area in July by the same persons, but without host data. lc, » recently molted clinging to grass (with Dermacentor rhinocerinus), 25 miles west of Yirol, 23 November 195Z, E. T. M. Reid legit. 10% and 2290 (some of which intergrade with R, simus simus) (cf. IDENTIFICATION below), from elephant near Kenisa, May 1953, E. T. M. Reid legit. Sudan Government collections contain other Kenisa area specimens from elephants, collected in 1911, which show similar intergrada tion with R. s. sims (cf. REMARKS below). Mr. Reid has sent me numerous specimens from three elephants shot near Yirol; these are typical R. s. simus. DISTRIBUTION R. simus senegalensis is a West and Central African tick with scattered loci in more humid areas of East Africa as far south as northern Nyasaland. WEST AFRICA: NIGERIA (Unsworth 1952. AsR. Sims: Simpson Bera p. 525; det. by Nuttall and Warburton as R. simus falcatus; see p. . As R. simus longoides: Unsworth 199, am 1950, Gambles 1951). TRACY MST AIOE (Koch 1844. eee 1951, 1953B. As R. simus longoides: Villiers 1955). SIERRA LEONE, IVORY COAST, GOMD C OD Cok TOS. (Zumpt 1943A*). PORTUGESE GUINEA (Tendeiro 19528, Ci519535 1954) . CENTRAL AFRICA: CAMEROONS (Zumpt 1943A*. Rageau 1951,1953A, B). FRENCH EQUATORIAL AFRICA (Zumpt 1943A*. Rousselot 1951). BELGIAN CONGO and RUANDA_-URUNDI (?As R. simus shipleyi: Bequaert 1930B,1931. Zumpt 1943A*. Rousselot 19538. Theiler and Robinson 1954. Van Vaerenbergh 1954. See HOSTS below). EAST AFRICA: SUDAN (In part as R. simus and as R. falcatus: King 1920. Hoogstraal 1954B,C). UGANDA and TANCANYIKA (Hoogstraal 1954C. J.B. Walker, unpublished; see HOSTS below). *A1] Zumpt (1943A) records are under R. simus longoides subsp. nov. which Zumpt (1950A) later synonymized under der R. simus senegalensis. - 755 - SOUTHERN AFRICA: NYASALAND (As R. falcatus: several spec- imens, Nuttall tot 1LO99A in BMNH, with numerous R. longus, from Chitipa Valley, Dowa District, 1910, J. B. Davey legit). NOTE: Koch's (1844) record from Egypt, subsequently quoted by numerous authors, is either based on a mistaken locality label or on a specimen of R. s. Sanguineus. Koch based his original description on females, only, ae Senegal (French West Africa) and Egypt. See REMARKS below. HOSTS Like the subspecies sims, R. simus senegalensis attacks a variety of larger game and domestic ani s. The immature stages probably feed on rodents, but no data concerning their host pref- erences are available. Domestic animals: Cattle (Zumpt 1943A*, Rousselot 1951, 1953B, Onsworth 1952, Rageau 1953B, Hoogstraal 1954C**). Horses (Nuttall lot 182 in BMNH, see NIGERIA above. Zumpt 19/3A*). Pigs (Rousselot 1951,1953B, Rageau 1953B). Sheep (Rousselot 1951,1953B). Dogs (Rousselot 1951,1953B, and Uganda and Sudan records above). Wild animals: Bushpig (Zumpt 1943A*). Warthogs (Nuttall lot 1099A, see NYASALAND above. Rousselot 1951,1953B. Sudan records above. Uganda specimens in BMNH). [ ?Giant eland (Bequaert 1930B,1931, see BELGIAN CONGO above) 7. Wildebeest (Walker , uripabl ished). Bongo (Rageau 1951). Roan antelope (Sudan record above). Buffalos (Hoogstraal 1954C**, Van Vaeren. berczh 1954, Villiers 1955, Theiler, unpublished. Walker, un. published. Sudan records above). Forest dwarf buffalo (Tendeii ro a RCC Elephants (Common hosts in mee El Ghazal; recorded above). Hunting dog, Lycaon pictus (Sudan record above). Lion (Congo specimens in MCZ ail collections). *All Zumpt (1943A) records are under R. sims longoides subsp. nov. which Zumpt (1950A) later synonymized under R. simus senegalensis. *#¥#My 1954C report of domestic cattle should be wild buffalo ("Bos caffer" on label). SoS NOTE: Rousselot (1951,1953B) records specimens from the cane rat, ee swinderianus. These specimens should be checked against R. Simpsoni. In the same reports he states that the short haired rat, Praomys jacksoni, is a host in the Belgian Congo. Un. less this note —— Oo parasitism by an immature stage, this animal would be a most unusual host. Santos Dias (19520,1953, 1954) states that one nymph along with many males has been taken from a forest dwarf buffalo. BIOLOGY Life Cycle It cannot be determined whether the life cycle data presented by Rousselot (1953B, p. 92) under R. simus senegalensis concern this subspecies or the subspecies Sims; other remarks under the same heading refer obviously to the subspecies sims. Unfortunate ly, no clues to the life cycle of R. simus sene alensis in nature are available. Rousselot (1953B, p. OL) claims that this is a three—-host subspecies. Ecology This is a tick of West African higher rainfall areas. Popu. lations that range into East Africa appear to be confined to animals found in forests, in more heavily vegetated savannah, and in the vicinity of lakes. The Boma Plains buffalo on which some Sudan specimens were taken was probably a migrant, for the Boma Plains are too arid for many months of the year to allow this tick to survive. According to Unsworth (1952), in Nigeria the subspecies senegalensis "appears to have approximately the same distri- bution as... sims, but it is not so common’. DISEASE RELATIONS It is claimed that specimens of R. simus senegalensis nat— urally infected with Q fever (Coxiella bur burnetit) have been found in Portugese Guinea. el (ey ae REMARKS Koch (1844) based his original description on females only. His material was reported as from Senegal and Egypt. It is most likely that the Egyptian record is due to a mistaken locality label. Just how the Senegal specimen has been associated with what is today called R. simus senegalensis has not been deter. mined. A female specimen of the R. simus group from Koch's time would be difficult to identify with any degree of certainty, especially to subspecies. Neumann (1911) synonymized senegalensis under R. simus. Zumpt (1943A,1950A) described what is now con- sidered as R. simus senegalensis. A number of the Sudan and Tanganyika collections listed here. in were sent to Santos Dias for identification, along with some specimens of R. longus. The R. simus senegalensis material was determined by him as k. aon us and the Re ions material was labelled R. capensis pseudolongus. Zumpt (1943A,1950A) warns his readers that heavily punctate R. simus senegalensis may superficially resemble R. longus. The R. Longus of ae Dias (1953D) appears to be what Zumpt con. ‘Siders a heavily punctate R. simus senegalensis. A further note is necessary concerning collections listed above as simus.senegalensis intergrades from elephants near Kenisa, Bahr El Ghazal Province. These are comparatively small, brownish specimens, with simus scutal punctation, posteromedian and para- median grooves absent or very faintly indicated, and adanal shields showing every degree of variation from the most typical simus type to the most typical senegalensis type. These series nicely corro- borate Zumpt's treatment of senegalensis as a subspecies of simus. IDENT IFICATION Male: Within the R. sims group, as described in the key ana under R. simus, males of the subspecies senegalensis are referred to a group With sickleshaped adanal shields. The scutal outline is definitely wider in relation to length than in most specimens of R. s. simus and the scutal surface is flat, not arched as in SB isis the subspecies simus. The posteromedian groove is long and narrow, the paramedian grooves are shorter and wider; these three grooves vary in depth and distinctness from specimen to specimen but are never deep or strikingly apparent. The distribution of large, scutal punctations is an important criterion in distinguishing this tick. On the anterior three-fifths of the scutum, large or moderate size punctations are arranged in four irregular rows of three to six puncations each. On the posterior two-fifths, there are from six to twenty large punctations in one and a half to three irregular, closely grouped rows on either side of the posteromedian groove. Scattered about the paramedian grooves are four to eight scattered large punctations. The interstitial punctations are always very shallow but they may vary in dis. tinctness from almost absent to large enough (though still super_ ficial) initially to confuse the basic pattern of large puncta. tions as described above. On close examination, however, this pattern is easily discernible. The lateral grooves are usually deep and long. One or three median festoons may protrude when engorged. The average size is that of the largest R. s. simus and the color, while usually jet black, may also be brownish, especially in smaller specimens. Female: This sex is difficult to distinguish from that of the subspecies simus, but in general its larger size, similarity of scutal punctation in comparison with that of the male, and its association with the male refers most specimens to the sub. species senegalensis. 5 (YS Figures 309 and 310, o&, dorsal and ventral views Figures 311 and 312, 9, dorsal and ventral views RHIPICEPHALUS SULCATUS udan Specimens PLATE LXAXVI = 100s RHIPICEPHALUS SULCATUS Neumann, 1908. (Figures 309 to 312) THE GROOVED BROWN TICK DISTRIBUTION IN THE SUDAN Bahr El Ghazal: 2&t' from an oribi, Ourebia ourebi subsp., near Tonj, 16 March 1953, N. A. Hancock efit. 180% 309 from chest of leopard, Panthera pardus subsp. and lo crawling on leg of man, Alel, 36 miles south of Yirol, 18 March 1953, E. T. M. Reid legit. These specimens are in the HH collection. These are the only definite records of R. sulcatus from the Sudan. Those reported by King (1926) have proven, upon examination of his specimens by Dr. Theiler, to be R. s. san. guineus. A single male was R. supertritus. DISTRIBUTION (Known Correct References) As now known, the range of R. sulcatus extends from the Congo and southwestern Sudan to Nyasaland. Whether this dis tributional picture is more restricted than this tick's range in nature remains to be seen. CENTRAL AFRICA: “CONGO" / Type locality as stated by Neumann (1908). According to Bequaert (1931) this probably refers to French Equatorial Africa (French Congo) but Theiler and Robinson (1953B) refer it to the Belgian Congo./ EAST AFRICA: SUDAN 5 Hoogstraal (1954B). Material iden tified by Dr. G. Theiler. TANGANYIKA /A few actual 99 R. sulcatus among R. s. sangui- neus collection reported as R. sulcatus by Zumpt (19Z2B); see Theiler and Robinson (1953B).7 ZayOle= SOUTHERN AFRICA: NORTHERN RHODESIA (Theiler and Robinson 1953B,1952). NYASALAND (9 from Chitala, Theiler and Robinson 19538). Known Incorrect References FRENCH WEST AFRICA: Rousselot (1951) reported specimens of "Re senguinc7s sulcatus", which, as illustrated (1953, figure 37), represent a ypical o and 9g of R. s. sanguineus. BELGIAN CONGO: Bequaert's (1931) "R. sulcatus" is R. longus ane to Zumpt (1942B) and confirmed by Theiler and Robinson 1953B). SUDAN: The material reported by King (1926) as R. sulcatus has been identified by Dr. Theiler as R. s. es (Sudan Government collections). The specimen from a dog a risk. supertritus a TANGANYIKA: Zumpt (1942B) in part. See Known Correct References above. Que stionable References FRENCH WEST AFRICA: Adults provisionally identified, from goats at Dakar (Theiler and Robinson 1953B). CAMEROONS: Rageau (1953A,B). UGANDA: Two doubtful o9 mentioned by Theiler and Robinson (1953B). Mettam (1932) recorded a puff adder as host. BELGIAN CONGO: Schwetz (1927); specimens need checking. NORTHERN RHODESIA: Adults from hares, civet, and genet (Matthysse 1954). SOUTHERN RHODESIA: Jack (1921,1928,1942); specimens need checking. MOZAMBIQUE: As R. sanguineus punctatissimus: (Santos Dias 1952H). SOUTHWEST AFRICA: Questionable 9 mentioned by Theiler and Robinson (1953B). UNION OF SOUTH AFRICA: Cooley (1934); specimens need checking. HOSTS (From Known Correct References Only) Hare (Lepus saxatilis) and two leopards (Theiler and Robinson 1953B). Leopard (aumpt 1042B). An oribi and a leopard, also one crawling on leg of man (Sudan records above). ru Oe te BIOLOGY Life Cycle The developmental periods under summer laboratory conditions have been reported by Theiler and Robinson (1953B) to be: PERIOD DAYS Preoviposition to larval hatching 44 Larva feeds Ss} we) 5) Postfeeding period 8 to 18 Nymph feeds Dito 6 Adults did not attach. The laboratory hosts were guinea pigs (Theiler, correspondence). Ecology Unstudied. DISEASE RELATIONS Unstudied. REMARKS Zumpt (1942B) included R. sulcatus in the R. capensis group, which is represented in the Sudan also by R. longus (see page 665) and Rk. supertritu 4 The original specimens of R. sulcatus are in collection num. ber 1439 of the Veterinary School of Toulouse. Santos Dias (1952H) considers R. sulcatus Neumann, 1908, to be a synonym of R. sanguineus punctatissimus Gerstacker, 1873, which Zumpt (1950A) considers to be a synonym of R. Ss. sanguineus. Inasmuch as the original description of R. sanguineus punctatis— simus is not sufficiently detailed satisfactorily to settle this question, and the type specimens have not been examined by Santos =) /OSm= Dias, the more conservative approach of Zumpt for this difficult problem is utilized here. Theiler and Robinson (1953B) stress the difference between this species and R. capensis Koch, 1844, and R. serranoi Santos Dias, 1950, two ticks not known to occur in the Sudan. In our territory, R. sulcatus is easily differentiated from all species but R. s. sanguineus in its coarser forms. In northern Sudan and usually in Central Sudan, R. s. sanguineus is usually not coarse enough to be confused with R. eet but in southern areas both species may superficially resemble each other. AL though Neumann's original description of R. sulcatus, together with its illustration, was excellent, the perplexing variation in R. s. sanguineus and the previously unrecognized fact that the latter species has a pattern of large punctations, no matter how densely punctate it may be, has long confused the real iden tity of R. sulcatus. We have spent dozens of hours trying to differentiate among hundreds of collections of R. s. sanguineus before the arrival of R. sulcatus specimens from Dr. Thotlers reared series. Afterwards, no question concerning their specific differences remained. If one turns a questionable male obliquely to the light, punctations of coarse R. s. sanguineus will be observed to be relatively shallow, mostly nondiscrete and sloping. Most im portant, a few more or less clearly defined rows of widely spaced, large, deeper, punctations will be noted among the shal. lower interstitial punctations. In R. sulcatus no row formation of large punctations is present, large punctations are numerous and deep, and medium size punctations are also deep. Specimens, if well preserved, should be briskly rubbed with damp tissue paper until their surface shines in order best to distinguish these characters. In females, the relative isolation of large punctations among small or medium size ones usually differentiates R. s. sanguineus from R. sulcatus in which large and medium size punctations are indiscriminately scattered. The actual relatively greater length of the sulcatus scutum, which gives a first impression of being even longer than it is, easily distinguishes most specimens. Some variable R. s. sanguineus females may so closely approximate this elongate Scutal appearance that care must be used when apply-— ing this character. al Obras Material referred to as R. sulcatus by Enigk and Grittner (1953), in their remarks on the breeding and biology of ticks, should be re-examined for identity. IDENTIFICATION The diagnosis presented below is modified from Theiler and Robinson (1953B). Male; Size small, averaging 2.5 mm. long and about 1.5 m. wide (Neumann's specimens from 2.9 mm. to 3.4 mm. long and from 1.5 mm. to 1.8 m. wide). Scutum narrow anteriorly, widening posterior of level of eyes, surface slightly convex, shiny. Lateral grooves discrete and pronounced, may contain large punc. tations; in some specimens extended to include penultimate fes_ toon. Posteromedian and paramedian grooves widely elongate, wrinkled. Punctations deep, medium to large size, dense, some confluent; fewer laterally but some on festoons, scapulae, and lateral folds. Eyes flat, may be bounded by a few large puncta tions. Coxa I may have a dorsal, wnpointed hump. Basis capituli one and a half to two times as wide as long, with pronounced lateral angles at anterior third of length (Neumann's original, frequently reproduced illustration of this species shows the basis capituli extremely wide and pointed, most probably the artist's impression, drawn foreshortened, gained from a downward. pointing capitulum). Adanal shields typically like those of large R. s. sous but in some specimens they may be more elongately rounde the ee: margin emarginate (as is common in smaller R. s. Sanguineus); an approach to the sickleshape is seen in a Sia b rane erey, Female; Medium size, unengorged about 3.00 mm. long and 1.75 mm. wide (Neumann; 3.0 mm. x 1.5 mm.). Scutum flat, shiny, rich brown, emargination wide; as broad as long or slightly longer than broad, but appearing long due to the pronounced lateral grooves and the somewhat tapering, sinuous posterior margin (Neumann; 1.5 mm. x 1.25 m.). Cervical pits short, deep; cervical grooves short and inconspicuous. Lateral grooves pronounced, with external ridge, and picked out with irregular punctations which may be confluent. Eyes large, flat, flush with surface, may be bounded dorsally by a few large confluent =. Ope punctations. Punctations medium to large, deep, somewhat un evenly densely scattered, sometimes confluent, extending onto lateral folds. Basis capituli about three times as broad as long; cornua short, blunt points; lateral angles fairly sharp at midlength; surface not so heavily punctate as in male; porose areas circular, a little more than their own diameter apart. Palpi slightly longer than basis capituli and twice as long as broad, segments 2 and 3 as long as broad, segment 1 visible dorsally. The larva and nymph have been described and illustrated by Theiler and Robinson (1953B). = “/00l= Figures 313 and 314, @, dorsal and ventral views Figures 315 and 316, 9, dorsal and ventral views RHIPICEPHALUS SUPERTRITUS an Specimens PLATE LXXXVII Sethe RHIPICEPHALUS SUPERTRITUS Neumann, 1907(B). (Figures 313 to 316) THE EAST-AFRICAN SHAGREENED TICK iN 0) Sc, EQUATORIA PROVINCE RECORDS iL 2 Kajo Kaji Syncerus caffer aequinoctialis Jan 1 Juba cerus catter aequinoctialis Nov (SVS) 1 "Alangw yneerus Catfer aequinoctialis - (SCC) Acholi" The male in the Sudan Government Collection, collected by H. H. King, had been identified by him as R. falcatus. The collecting locality of this specimen may now be just inside the Sudan or just over the Uganda border. DISTRIBUTION IN THE SUDAN Upper Nile: Sudan Government collections contain a male spec imen, Sees from a dog at Bor by H. H. King, which had been identified and reported by him (1926) as R. sulcatus. DISTRIBUTION R. supertritus is an uncommon East African tick that ranges into REL Africa and into the northern part of southern Africa. CENTRAL AFRICA: CAMEROONS (Schulze 1941). BELGIAN CONGO (Neumann 19075,1011. Massey 1908. Nuttall and Warburton 1916. Bequaert 1930B,1931. Zumpt 1942B). EAST AFRICA: SUDAN (In part as R. sulcatus: King 1926. Hoogstraal 19528). ERITREA (HH collection from Nacta). UGANDA (Nuttall lot 2396 in BMNH collections, from Gomba). KENYA (Lewis 19314,C, 1933,1934). TANGANYIKA (Zumpt 1942B). Sis SOUTHERN AFRICA: ANGOLA (As R. coriaceus: Nuttall and Warbur. ton 1908 and Warburton 1912. Sousa Dias 1950. Santos Dias 1950). NORTHERN RHODESIA (Theiler and Robinson 1954). SOUTHERN RHODESIA (Jack 1921,1928,1942). NYASALAND (As R. coriaceus: Nuttall and Warburton 1908, Old 1909. Warburton 1912. Neumann 1908 ,1911. Neave 1912. Zumpt 1942B. Theiler 1947. Wilson 1950B. Hoogstraal 19540). MOZAMBIQUE (Santos Dias 1950B,1952D,1952H,1953B). HOSTS R. supertritus is an uncommon parasite of larger wild animals and appears to feed only occasionally on domestic animals. Hosts of immature stages are unknown. Domestic animals: Horse (Neumann 1907B,1911, Massey 1908). Dog (Sudan record above). Cattle (Nuttall and Warburton 1916, Matthysse 1954, Theiler and Robinson 1954). Antelopes: Eland (Bequaert 1930B,1931). Zambesi eland (Santos Stes-19538). Kudu (Jack 1942, Santos Dias 1953B, Hoog- straal collection from Eritrea. Hartebeest (Zumpt 1942B, Wilson 1950B). lLichtenstein's hartebeest (Santos Dias 1952H,1953B). Sable antelope (Theiler 1947, Wilson 19508). Other mammals: Buffalo (Theiler 1947, Wilson 1950B, Santos Dias 1950B,1052D,1953B, Sudan records above). Rhinoceros (Lewis 1933). Warthog (Hoogstraal 19540). Lion (Lewis 1934). Zebra (Zumpt 1942B). BIOLOGY This interesting tick seems to be rare wherever it occurs. Its life cycle is unknown. It sometimes occurs in large numbers on single animals or herds. For instance, in Matthysse's (1954) study of ticks in Northern Rhodesia, this tick was found only once on cattle, but the collection consisted of eighteen males and fifteen females. Sys DISEASE RELATIONS Unstudied. REMARKS The haller's organ of R. supertritus has been illustrated by Schulze (1941). Zumpt (1942B) includes R. supertritus in the R. capensis group (see R. longus, page 667). IDENT IFICATION This rugose species has been described best by Theiler (1947). Variations are within the general remarks below. Male: This species is usually large, from 3.3 mm. to 5.3 m. long and from 1.3 mm. to 3.3 mn. wide, and usually black. It is easily recognized by a combination of characters including a pointed dorsal projection of coxa I; scutal punctations that are large, dense, and closely spaced or contiguous; and conspicuous reticulation or shagreening of the cervical areas and of the posterior grooves. The posteromedian groove is longer and nar- rower than the paramedian grooves; the lateral grooves are wide and deep. One or three median festoons protrude upon engorgement. The narrowly elongate adanal shields have a moderately convex outer margin and an almost straight or slightly concave inner margin; these margins meet at a pointed or a rounded anterior and posterior juncture and the shields are more ovoid than tri- angular in shape. Female: Conspicuous shagreening or reticulation of the cer- vical areas and of the lateral grooves also distinguishes females of this species. The punctations are coarse and rugose, adjacent or contiguous centrally. The dark brown scutum, which is about as wide as long, has flat eyes at about midlength and a sinuous or gradually rounded posterior margin. Its pronounced lateral grooves are impunctate and extend to the posterior margin; the cervical grooves are deep and converging anteriorly, superficial and diverging posteriorly. SO 318 320 Figures 317 and 318, &, dorsal and ventral views Figures 319 and 320, 9, dorsal and ventral views RHIPICEPHALUS TRICUSPIS an specimens PLATE LXXXVIII Ae RHIPICEPHALUS TRICUSPIS Donitz, 1906* (= R. LUNULATUS Neumann, 19078). (Figures 317 to 320) THE TRICUSPID GLOSSY TICK DISTRIBUTION IN THE SUDAN Bahr Bl Ghazal; All from Galual-Nyang Forest, 1953; E. T. M. Reid or N. A. Hancock, collectors: 2%’, 299 from tiang, Damaliscus korrigum tiang, April. lo from warthog, Phacochoerus aethiopicus Subspp., 4 September. 1c from oribi, Ourebla ourebi subspp., fi June. lo from giraffe, Giraffa camelopardalis subspp., 17 August. 200’, lo from domestic horse from ae > 20 June. R. tricuspis is not known from elsewhere in the Sudan but probably also occurs on the west bank of Equatoria Province. DISTRIBUTION Re ees is scattered throughout Africa, within the Ethiopian Faunal Region. WEST AFRICA: NIGERIA (Unsworth 1949,1952. Gambles 1951). TOGO (As R. glyphis: Donitz 1910A). SIERRA LEONE (Zumpt 1943A). FRENCH WEST (Rousselot 1951,1953B). PORTUGESE GUINEA (Tendeiro 1951E,1952B,C ,E,1953,1954). GOLD COAST (Stewart 1935). *Dr. G. Theiler has kindly made an extensive study of the morpho- logical variation, distribution, ecology, and taxonomy of this species especially for this work. See also her review of R. tricuspis (1947, pp. 292-298). ot (12m CENTRAL AFRICA: *CAMEROONS (Unsworth 1952. Rageau 1953A,B). *#FRENCH EQUATORIAL AFRICA (Rousselot 1953B). *BELGIAN CONGO (Neumann 1907B. Massey 1908. Schoenaers 1951A. Nuttall and Warburton 1916. Schwetz 1927B. Bequaert 1930B,1931. Fain 1949. Theiler and Robin son 1954. Santos Dias 1954D. Van Vaerenbergh 1954). EAST AFRICA: SUDAN (Hoogstraal 1954B). TTALIAN SOMALILAND (Paoli 1916. Stella 1940). UGANDA (Neave 1912. Mettam 1932. Wilson 19500). KENYA (Lewis 1931¢. Binns 1951). TANGANYIKA (Donitz 1910B. As R. glyphis: Donitz 1910A. Zumpt 1943A). = SOUTHERN AFRICA: ANGOLA (Gamble 1914. Sousa Dias 1950. Specimens collected by Wellman seen in BMNH; see also HOSTS below). MOZAMBIQUE (Santos Dias 1950D,1952 ,D,H,1953B). NORTHERN RHODESIA (Theiler 1947. Matthysse 1954. Theiler and Robinson 1954). ¥*NYASALAND (Neave 1912. Warburton 1912. Theiler 1947. Wilson 1950B). *BEC HUANALAND (Donitz 1906,1910B. Theiler 1947). SOUTHWEST AFRICA (DOnitz 1910B). *UNION OF SOUTH AFRICA (Howard 1908. Bed ford 1920,1926,1932B. Theiler 1947). HOSTS Domestic animals and many larger game animals serve as hosts for R. tricuspis. Immature stage hosts in nature are unknown. Domestic animals: Horses (Neumann, 1907B, Massey 1908, Zumpt 1943A, Sudan records above). Cattle (Donitz 1910A,B, Nuttall and Warburton 1916, Schwetz 1927B, Zumpt 1943A, Theiler 1947, Wilson 1950B, Binns 1951, Rousselot 1951,1953B, Schoenaers 1951A, Santos Dias 1953B, Matthysse 1954, Van Vaerenbergh 1954). Sheep and “Theiler (correspondence) has seen additional material from these territories. = V3 <= goats (Nuttall and Warburton 1916, Theiler 1947, Wilson 1950B). Pigs (Gamble 1914, Tendeiro 1951E,1952B). Dogs (Gamble 1914, Nuttall and Warburton 1916, Wilson 1950B, Santos Dias 1952, Matthysse 1954). Wild animals: Lion and serval (Wilson 1950B). Leopard (Zumpt 19438). "wild dog" (Lewis 1931C). Buffalo (Wilson 1950B, Santos Dias 1953B). Forest dwarf buffalo (Tendeiro 1951E,1952, C ,E,1953,1954). Zebra, Lichtenstein's hartebeest, and sable antelope (Santos Dias 1953B). Oribi (gwape) and steinbuck (Wilson 1950B). Duikers, various (Wilson 1950B, Santos Dias 1953B). Waterbuck and bushbuck (Mettam 1932). Reedbuck (Wil- son 1950B, Santos Dias 1950D,1953B). Oribi, tiang, and giraffe (Sudan records above). Bushpig (Donitz 1910A, Zumpt 1943A, Santos Dias 1950D,1953B). Warthog (Wilson 1950B, Santos Dias 1953B, Sudan record above). Hedgehog (Neumann 1911). Hares (Theiler 1947, Wilson 1950B). Fruit bat (Specimens collected by i Jordan, now in BMNH; probably a decidedly exceptional host). BIOLOGY Life Cycle R. tricuspis has been reared by Theiler (correspondence) for the morphological studies reported by her (1947). The life cycle details will be published subsequently. Ecology In order to determine whether there might be ecological or other distributional factors between R. tricuspis and what has been reported as R. lunulatus, Theiler (correspondence) has checked all locality records against data for vegetation type, rainfall, extent of dry season, and relative humidity. She finds that both range indiscriminately from (1) dry forests of the Rhodesian highlands through (2) highgrass-low tree savannah or Guinea.southern Congo savannah, and (3) acacia desert grass savannah of the Sudan and northern Kalahari to (4) tallgrass subtropical evergreen and deciduous tree and thorn forest. =~ Th = In South Africa (Theiler 1947), R. tricuspis occurs in the warmer areas with thorn trees, from the semiarid bushveld of the Kalahari to the moister bushveld of the lowveld of northern and eastern Transvaal and of Natal. It is absent in the Karroo scrubveld, the open grassveld of Orange Free State, and the middleveld and highveld of Transvaal. Dr. Theiler's present study indicates that R. tricuspis appears to be resistant to a wide range of humidity and aridity. It may be found in heavy rainfall areas with 42 inches annually and in areas where there is as much as seven months of drought. Santos Dias (1950D, 1952) records of R. tricuspis and lunw latus" are from areas with the same kinds of vegetation | and drought periods and the same average rainfall. Donitz's (1910B ) specimen from Little Namaqualand probably fell from its host when the Trekboers were migrating with their sheep from Bush. manland and the southern Kalahari into Namaqualand in search of rains and pasturage. In Northern Rhodesia, Matthysse (1954) found adults of the tricuspid brown tick mainly in the tail brush of cattle but also on the feet, anus, and ears. Most specimens that I have seen have been from the tail brush. They appear to be present chief ly during the rains but also in the dry season. DISEASE RELATIONS MAN: Specimens from Portugese Guinea have been reported to be free of Q fever (Coxiella burnetii). PIGS: This tick is "possibly a vector of porcine piro- plasmosis (Babesia trautmanni)". REMARKS R. tricuspis was described by Donitz (1906) from the Kala. hari of Bechuanaland. The following year, Neumann (19078) des. cribed R. lunulatus from the Congo, and in 1910(A) Donitz des. cribed R. glyphis from Togo and Tanganyika. Donitz's papers SE were long overlooked by earlier workers, and obvious specimens of R. tricuspis were identified as R. lunulatus. Zumpt (1943A ,1950A) accepted the differing descriptions of R. tricuspis and R. junulatus as referring to separate species. Fle “Synonymized R. glyphis under R. lumulatus but did not see specimens of R. tricuspis. Theiler (L047) reared R. tricuspis and observed that the range of variation in the Fj generation from a single female included characters ascribed to R. tricuspis as well as to R. lunulatus. For the purposesof the present work, Theiler has restudied her material and confirmed her earlier ob- servations. Santos Dias (1950D ,1952C) described differences, which Theiler has found to be intraspecific, for R. tricuspis and R. lunulatus. a It might be indicated that in Theiler and Robinson (1954) the term R. lunulatus is used only for literature references of that name, but does not infer that these authors consider R. lunulatus to be a valid species. a In the following description, the range of variation in Theiler's material is noted. Only typical specimens were des— cribed in her 1947 paper, in which R. tricuspis was compared with R. sims simus. IDENTIFICATION Male: The somewhat linear arrangement of the few large, deep Scutal punctations identifies R. tricuspis in the R. simus group within which it stands out easily by the curiously sinuous posterior margin of the adanal shields that are projected more or less as a spur at the inner or outer juncture, or at both junctures. In typical specimens, these two pointed spurs and the pointed, heavily sclerotized accessory anal shields give this area of the tick a tricuspid appearance. The spurlike outer juncture is pronounced but the inner point is frequently more rounded and shorter. Lateral grooves, usually well indicated, rarely may be much reduced or indicated by only @ line of adjacent punctations. The posterior grooves are typically distinct but in some specimens tend to disappear. [ Note that in specimens of R. simpsoni from a single host the same variation in scutal grooves - 776 - occurs. Small rhipicephalid species that crowd into a small area of the host frequently show these modifications ./ Small or fine interstitial punctations are present, thouch they may be faint. Coxa I bears a pointed dorsal projection that typically is promi_ nent but in some specimens is smaller, though still pointed. This reddish to black tick usually has a pearshaped body and measures up to 3.5 mm. or even 4.3 m. long. Female: Usually small size (up to 4.5 m. long and 2.3 m. wide) and association with the male distinguishes this sex from that of R. s. sims. Clearcut characters to distinguish these two females are difficult to define in view of the frequent reduction of the lateral grooves in Rk. s. sims. In most spec. imens of R. tricuspis the shieldshaped Scutum contrasts with the subcircular scutum of R. s. simis. The lateral grooves are characteristically short (shorter than in R. s. sims) and contain four to six closely adjacent punctations. Larvae and nymphs have been described by Theiler (1947) and compared with those o R. s. sims. - 777 RHIPICEPHALUS ?SP. (not illustrated) ia, NS One EQUATORIA PROVINCE RECORDS Zam alie Imatong Heterohyrax brucei hoorstraali Feb Tee el Imurok Heterohyrax brucel hoogstraali Feb REMARKS . These nymphs were originally identified as those of R. muhlensi (Hoogstraal 19543). Subsequent study of the difficult problem of immature rhipicephalid identification indicates that they are of a different species and most closely resemble R. maculatus Neumann, 1901, (Theiler, correspondence) an ornamented species from southeastern Africa that is not known from the Sudan. Note (pase 637) that it is quite possible that these spec— jmens are the immature stages of the species referred to herein as . ?distinctus. = TIS = ANIMALS AND OTHER SOURCES FROM WHICH TICKS HAVE BEEN COLLECTED The following lists are a résumé of Sudanese tick host records in the present collection. The fauna of Torit District in Equatoria Province has been most thoroughly studied, that of Eastern and Juba Districts, to the east and west of Torit District respectively, has also received considerable attention though not to the extent of that in Torit District. These three Districts comprise that part of Equatoria Province lying east of the Nile. The west bank of Equatoria Province remains poorly known and will undoubtedly pro- vide a rich source of new data to future workers. In Bahr El Ghazal Province, the area from Yirol to Wau and northwards has been fairly well studied; the remainder of this Province has been surveyed but should receive more attention. Upper Nile Province is the least studied of this group of three Provinces with tropical African savannah landscape and big game animals. Some East African tick species presently unknown in the Sudan may occur on animals in this area. Hosts from which ticks have been collected in the remaining Provinces, comprising desert scrub, semidesert, and desert zones of the Sudan, are mostly domestic animals. Kassala and Northern Provinces are less well represented in these collections than Darfur and Kordofan Provinces. Equatoria Province data are more representative of the over all picture of host parasite relationships than those of other Provinces and are, therefore, reviewed in greater detail than data for other Provinces. Certain significant negative data are also included. - 779 - $19;9WO|!) XIXXXT GLVId MNvd LSVa SHILITVOOT ONILOMTION AONTAOUd VIUOLYNCE Tze emsty off otf (vrONVeV») (WNH9) VONVON alt trsetee DUIO\ 018 fy N Hd 10GNY oS VIdOIHLS ae oGE 19» obuojol BINDS, “a vnoLoonoa: (VNV WYN D RAS ‘exe : VANIY - 10yo}50N ms \ ‘ younwje ( LOIM am SvH VONIGIO STEED buo © oksjuog ! aN “ y; fier a ! j LIN DAO. — 3 es0buos0y* EAP DEL) Ie} va nt n “ : # __\) ppunioge gM oiio7s-7 —¢ ou0109 < - % \ _0depIy) y OUT. \ ¢ = \ \ : X oj 06u06y * \ 0je0doy \ * \ « p0ley “ SiW LIdv7 ‘ vi 1400/07