A Systematic Revision of the Genus Laelaps S. Str. (Acari: Mesostigmata) of the Ethiopian Region

Great Basin Naturalist

Volume 36 Number 2 Article 1

6-30-1976

A systematic revision of the genus Laelaps s. str. (Acari: Mesostigmata) of the Ethiopian region

C. Selby Herrin Brigham Young University

Vernon J. Tipton Brigham Young University

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Recommended Citation Herrin, C. Selby and Tipton, Vernon J. (1976) "A systematic revision of the genus Laelaps s. str. (Acari: Mesostigmata) of the Ethiopian region," Great Basin Naturalist: Vol. 36 : No. 2 , Article 1. Available at: https://scholarsarchive.byu.edu/gbn/vol36/iss2/1

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Published at Provo, Utah, by Brigham Young University

Volume 36 June 30, 1976 No. 2

A SYSTEMATIC REVISION OF THE GENUS LAELAPS S. STR. (ACARI: MESOS riGMATA) OF THE ETHIOPIAN REGION^

C. Selbv Herrin- and Vernon J. Tipton-

Abstract.— This paper presents the results of a systematic study of mites of the genus Laelaps s. str. collected from small mammals of the Ethiopian region. Specimens taken from approximately 100.000 small mannnals were examined from a wide variety of habitats and localities. The Ethiopian fauna of Laelaps mites includes 31 species. 4 of wliich are described as new: L. parasimillimus, L. tnyomys. L. malacomys. and L. aconiys. A rmmerical taxonomic analysis was made, the results of which were used in the preparation of a proposed classification of the African species of Laelaps. A key for identification of females is given, and females and males (where known) of all species are illustrated. Diagnostic characters are given for the female and male of each species. Collection data and, where pertinent, discussions of morphological characters and variability are i)rovided. Also included are discussions of host-parasite associations.

The objective of this paper is to present M. Lavoipierre. They were responsible a systematic revision of the genus Laelaps for the description of 21 Laelaps species s. str. (i.e., not inckiding species of and the publication of many new host and Echinolaclaps Ewing) of the Ethiopian locality records. Tipton (1960) treated region. There has been no recent pubHca- the genus Laelaps worldwide; however, tion which presents a sufficiently compre- 11 of the 32 species now known from hensive taxonomic review of this group Africa were described after this work. In of mites in Africa. Because of the great his book Arthropod Parasites of Verte- similarity as well as diversity among the brates in Africa South of the Sahara, Laelaps species in Africa, there has been Zumpt (1961) listed 22 species. The only a definite need for a complete, compre- keys to the identification of African spe- hensive revision of this group of mites. cies of the genus were those of Tipton This need is increased by the great di- (1960) and Taufflieb (1959). versity of Laelaps taxa found in the col- The concept of the genus Laelaps fol- lections from the Smithsonian African lowed in this paper is basically that of Ectoparasite Project. Tipton (1960). That is, we do not feel Several scientists have contributed that Laelaps and Echinolaelaps should be greatly to the knowledge of parasitic grouped together without at least separate Laelaps mites of the Ethiopian region. subgeneric status for each. Thus, this pa- Stanley Hirst (1912 to 1925) described as per deals only with Laelaps s. str. (sub- new seven species of African Laelaps, genus Laelaps) as recognized by Tipton which were included in Bedford's (1932, (1960) and does not include Echinolae- 1936) checklists of ectoparasites of Ethio- laps. The dorsal chaetotaxy signatures pian vertebrates. During the years be- followed in this paper are those of Hirsch- tween 1937 and 1954 Charles Radford mann (1957), and the morphological ter- published several papers dealing wdth new minologv is basicallv that of Evans and species and new host and collection rec- Till (1965). ords. In the 1950s and 1960s additional Following the discussion of taxonomy contributions were made by Drs. F. and classification analyses and the identi- Zumpt, R. Taufflieb, H. L. Keegan, and fication key to females, each species is

^Supported by the Smithsonian-African Ectoparasite Project through contract (DADA 17-73-C-3042) of the Medical Re- search and Development Command, Office of the Surgeon General, U.S. Army. -Center for Health iind Environmental Studies. Brigham Young University, Provo, Utah 84602.

113 114 GREAT BASIN NATURALIST Vol. 36, No. 2 treated as follows: synonymy, brief de- per: SAIMR = South African Institute scription of female and male (where for Medical Research; AMP =^ African known), summary of all collection rec- Mammal Project; USNM = United States ords (literature as well as collections of National Museum of Natural History, the African Mammal Project), and brief Smithsonian Institution; BMNH = Brit- discussion of differential diagnostic char- ish Museum (Natural History); Tauff- acters and host-parasite relationships. lieb = Roger Taufflieb, Dakar, Senegal. For the species described herein as new, A list of 159 characters was compiled the collection records are presented in for use in this study (Tables 2 and 3); more detail. of these, 33 were qualitative and 126 were For each species described as new to quantitative (measurements). Each speci- science, the holotype, allotype (where de- men selected for inclusion in the analyses scribed), and one or more paratypes are was examined and the value of each to be deposited in the U.S. National Mu- character recorded on data forms. The seum of Natural History, Washington, data were entered on computer punch D. C. Paratypes are to be deposited in cards preparatory to the com])uter anal- the collection of the South African Insti- yses. tute for Medical Research, Johannesburg, South Africa, and in the collections of Computational and Analytical Methods the authors. The computer analyses were j)erformed We acknowledge with gratitude the as- on the IBM 360 model 65 computer at sistance of the many people associated the Brigham Young University Computer with this study. Special thanks are given Center using the Niunerical Taxonomy to Dr. Henry W. Setzer for logistic sup- vSystem of Multivariate Statistical Pro- port and to Dr. Deane P. Furman for re- grams (NT- SYS) prepared by Dr. F. viewing the manuscript. We are grateful James Rohlf and associates of the State to the South African Institute for Medical University of New York at Stony Brook. Research (Dr. F. Zumpt) and the British The specific procedures used in the anal-

Museum, Natural History (Mr. K. H. yses were as follows: (1 ) transformation of Hyatt, curator), for the loan of type the basic data matrix by standardization specimens, and to Dr. R. Taufflieb, who (Sokal 1961); (2) computation of Pear- .sent specimens from his personal collec- son's product-moment correlation (Mich- tion. The Center for Health and Environ- *^ner and Sokal 1957) and Sokal's (1961) mental Studies provided the laboratory taxonomic distance to produce similarity si)ace and equipment used in this study. matrices; (3) analysis of each similarity Sheila E. Ford and Jerry N. Norton pre- matrix by the UPGMA cluster analysis, pared the illustrations. yielding a phonogram (a graphic presen- tation of phonetic resemblance for each Materials and Methods matrix; and (4) cophenetic correlations were computed to arrive at an estimate of Materials Utilized the degree of information transferred from Type specimens of most of the pre- the similarity matrices to the phenograms. viously described Loclaps species of Africa The purpose of these numerical taxonomic were obtained from various museums and analyses was to objectively evaluate the individuals for e.xamination and use in affinity or similarity between the taxo- the numerical taxonomic analyses. Where nomic units. The results of these analyses type specimens were not available, posi- were then used in making decisions re- tively identified representative specimens garding the validity of all previously de- from the ty[)e locality were obtained for scribed Laclaps sjiecies as well as new each species. Also, representative speci- taxa included in the analyses. The final mens of the various taxa found in {\\o propositi classification is based primarily collections of the Smithsonian-African oil this |)liase of th(^ investigation. Ectoparasite Project were included in th(> numerical taxonomic studies. The s[)eci- Ri',sui,Ts AND DiscrssiON mens (OTUs) included in the numerical Taxonojuic .Xnah ses taxonomic analyses are listed in Table 1. The following source abbreviations are Prior to the iuiriieri(.d t.ixoiiomic anal- used in Table 1 and elsevvluM-e in llic [)a- yses, a pi-oposed c lassification of tlu^ genu^ . .

June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 115

Table 1. Laelaps specimens included in the nnmericai taxonomic analyses.

Species Type Host Locality Source

L. transvaalensis Paratype Otornys irroratus Transvaal, So. Africa SAIMR L. keegani Arvicanthis niloticus Northern Nigeria SAIM3\ L. congoicola Paratype Oenornys hypoxanthus Brazzaville, Congo Taufflieb L. parasimillimus ii. sp. Holotj-pe DephoTuys defua Sou b re, Ivorj- Coast AMP L. simillirnus Paratype Thallotnys narnaquensis Transvaal, So. Africa SAIMR L. grenieri Paratype Lemniscomys strialus Brazzaville. Congo Taufflieb L. thamnomys Paratype Thanujornys rutilans Brazzaville, Congo Taufflieb L. kampalensis Paratype Lemniscomys striatus Kampala, Uganda SAIMR L. moucheti Paratype "rodents" Yaounde. Cameroons Taufflieb L. lavieri Paratype Mus bella Brazzaville. Congo Taufflieb L. nigeriensis Holotype Crocidura sp. Adu. Nigeria USNM L. fritzumpti Paratype Ratlus paedulcus Kalahara, So. Africa Taufflieb L. lavoipirrrei Paratype Lophuromys sikapusi Yaounde. Cameroons Taufflieb L. tillac Paratype Lemniscomys griselda Transvaal. So. Africa Taufflieb L. roubaudi Paratype Dasymys incomptus Brazzaville, Congo Taufflieb L. peregrinus Paratype Rhabdomys pumilio Transvaal. So. Africa Taufflieb L. peregrinus Rhabdomys pumilio (ORS) So. Africa AMP L. nuttalli S\nt>T5e Mus norvegicus Sierra Leone BMNH L. malacomys n. sp. Holotype Malacomys edwardsi Belekoam. Ivory Coast AMP L. algericus Syntype Mus algericus Tougoury, Algeria BMNH L. oraniensis Syntype "field mice" Oran. Algeria BMNH L. brandbergensis Paratope Petromyscus collinus Brandberg. SW Africa SAIMR L. zumpti Holotype Mus Irilon Njoro. Kenya USNM L. Tnyomys n. sp. Holotype Myomys daltoni Casa Manee. Senegal AMP L. liberiensis Holotype Epimys defua Liberia BMNH lamborni Syntype "Kapuku" Karonga, Nyasaland BMNH setzeri Paratype Praomys jacksoni Ora. Western Nigeria SAIMR brazzai Paratype Praomys sp. Brazzaville. Congo Taufflieb benoiti Paratope Mus bella Brazzaville. Congo Taufflieb aethiopicus S>Titype "rats" Wanga. Kenya BMNH vanxomereni S>Tit\'pe "rodent" So. Bugishu. Uganda BMNH acomys n. sp. Holotype Acomys spinosissineus Manicaland, Rhodesia AMP bocquieri Parat\T3e Chryssochloris Brazzaville. Congo SAIMR leucorrhina L. spinifer Hybomys sp. Congo Taufflieb L. paraspinosus Syntype Arvicanthis dorsalis So. Africa BMNH L. breviperitremus Paratvpe Acomys subspinosus Transvaal. So. Africa USNM L. kochi Arvicola terrestris Teheran Prov., Iran AMP

T.-VBLE 2. Quantitative characters (measure- 22. Length setae st. 4 (metastemal) ments) of females used in the numerical taxo- 23. Least width genital plate between coxae IV nomic analyses. 24. Greatest width genital plate 25. Total length genital plate Gnathosoma 26. Length genital plate from 1st genital setae 1. Greatest widtli at level of gnathosomal setae to posterior end 2. Length from base to palpal trochanter 27. Distance between 1st pair genital .setae 3. Lengtli of palps 28. Distance between 2nd pair genital setae 4. Length 2nd chelireral segment 29. Distance between 3rd pair genital setae

.5. Length chelae 30. Distance between 4th pair genital setae 6. Length distal hypostomal setae 31. Length 1st pair genital setae 7. Length medial hypostomal setae 32. Length 2nd pair genital setae 8. Length lateral hypostomal setae 33. Length 3rd pair genital .setae 9. Length gnathosomal setae 34. Length 4th pair genital setae 10. Distance between gnathosomal setae 35. Distance between genital plate and anal

1 1 Distance medial hypostomal setae to gnatho- plate somal setae 36 Greatest width anal plate 37. Length anal plate from anterior margin to Ventei postanal seta 12. Width sternal plate at level of coxae II 38. Distance between adanal setae 13. Median length sternal plate 39. Distance from anterior margin anal plate to

14. Distance between setae st. 1 adanal setae 15. Distance betw-een setae st. 2 40. Distance from adanal seta'^ to postanal seta 16. Distance between setae st. 3 41 Length adanal setae 17. Distance between setae st. 1 and st. 2 42. Length postanal seta 18. Distance between setae st. 2 and st. 3 43. Length short setae of unarmed venter

19. Length setae st. 1 44. Length longer setae of unarmed venter 20. Length setae st. 2 45. Length metapodal plates 21. Length setae st. 3 46. Width metapodal plates ..

116 GREAT BASIN NATURALIST Vol. 36, No. 2

Legs 116. 47. Greatest width coxa I 48. Midventral length coxa I 49. Length proximal seta coxa I 50. Length distal seta coxa I 5L Length anterior dorsal seta 1 of femur I 52. Length posterior dorsal seta 1 of femur I 53. Width genu I 54. Length tarsus I 55. Greatest width coxa II 56. Midventral length coxa II 57. Length anterior seta coxa II 58. Length posterior seta coxa II 59. Length tibia II 60. Width tibia II 61 Length tarsus II 62. Greatest width coxa III 63. Median length coxa III 64. Length anterior seta coxa III 65. Length posterior seta coxa III 66. Length genu III 67. Width genu III 68. Length tibia III 69. Length tarsus III 70. Greatest width coxa IV 71. Median length coxa IV 72. Length seta coxa IV 7^. Length trochanter IV 74. Width trochanter IV 75. Length femur IV 76. Length genu IV 77. Width genu IV 78. Length tibia IV 79. Length tarsus IV

Dorsum 80. Length of peritreme 81. Median length of dorsal plate 82. Greatest width of dorsal plate 83. Distance between setae r1 84. Distance between setae si 85. Distance between setae r2 86. Distance between .setae r5 87. Distance between setae i3 88. Distance between setae z1 89. Distance between setae i4 90. Distance between setae 7.2 91 Distance between setae i5 92. Distance between setae s6 93. Distance between setae SI 94. Distance between setae J2 95. Distance between setae J3 96. Distance between setae J4 07. Distance between setae Z4 08. Distance between setae S4 99. Distance between setae J5 100. Distance between setae Z5 101. Distance befvveeii setae si and i4 102. Distance between setae i4 and i5 103. Distance between setae 7.1 and z3 104. Distance between setae i5 and .13 105. Distance between setae J4 and Z5 106. Distance between setae .15 and posterior eiul of dorsal plate

107. Length of seta i1 108. I,engt}i of seta r1 109. Length of seta si 110. T,ength of seta i2 111. Length of seta s2 112. Length of seta r2 113. Length of seta r3 114. Length of seta r4 115. Length of seta s5 June 1976 HERRIN, TIPTON: AFRICAN LAEL/\PS 117

15. Position of adanal setae relative to anal ( 1 ) Setaceous orifice: (2) Spinelike orifice Peglike ( 1 ) Posterior to anal (3) (2) Slightly posterior to middle of anal 31. Form of preapical setae on tarsus III: orifice (0) All setaceous (3) At level of middle of anal orifice (1) One spinelike (4) Slightly anterior to middle of anal (2) One peglike orifice (3) Two spinelike (5) Anterior to anal orifice (4) Two peglike 16. Form of adanal setae: 32. Form of ventral seta of coxa IV: (1) Setaceous and slender (1) Setaceous (2) Spinelike and stout (2) Spinelike 17. Anterior extension of peritreme: (3) Peglike 33. Form of preapical setae on tarsus IV: ( 1 ) Anterior of co.xa I (2) Middle of coxa I (0) All setaceous (3) Posterior of coxa I (1) One spinelike (4) Anterior of coxa II (2) One peglike (5) Middle of coxa II (3) Two spinelike 18. Chitinization of anterolateral margins of dor- (4) Two peglike sal plate: (5) One spinelike and one peglike (1) Normal chitinization (6) Three spinelike (2) Heavily rliitinized (7) Three peglike 19. Number of setae on dorsal plate (8) Two spinelike and one peglike 20. General form of central and anterior dorsal (9) One spinelike and two peglike setae: (1) Setaceous Laelaps s. str. was prepared based upon (2) Spinolike. stout and short classical taxonomic methods (i.e., con- Legs sideration of unequal weight ^ven to a 21. Form of distal seta of coxa I: sinaller number of key characters). This (1) Setaceous (2) Spinelike proposed classification, as presented in (3) Peglike Table 4, defines three major groups 22. Form of proximal seta of coxa I: {similUmus group, nuttoUi group, and Setaceous (1) vansomereni primarily (2) Spinelike group) based upon (3) Peglike the form of the proximal and distal setae 23. Form of proximal ventral seta of trochanter I: of coxa I. The arrangement of taxa with- (1) Setaceous in the three groups is based on subjec- (2) Spinelike tive judgments after ha\ang examined 24. Relative length of pd 1 seta of femur I: (1) Subequal to ad 1 seta representative specimens of all taxa. No (2) About 1.5 times length of ad 1 seta subgroups were defined in this arrange- Two times or more length of ad 1 seta (3) ment. 25. Form of anterior seta of coxa II: (1) Setaceous Figure 1 summarizes the taxonomic re- (2) Spinelike lationships given by the UPGMA cluster (3) Peglike anaylsis of a standardized correlation co- 26. Form of posterior seta of coxa II: efficient matrix based an original (1) Setaceous upon (2) Spinelike (acute) data matrix of 37 OTUs and 159 charac- (3) Peglike (blunt) ters. The cophenetic correlation coefficient 27. Form of ventral proximal seta of trochanter for this phenogram (Fig. 1) was 0.669, II: (1) All setaceous which is not too high, yet higher than that (2) One spinelike for the phenogram of the taxonomic dis- (3) Two spinelike tance matrix. A phenon line drawn ver- 28. Form of preapical setae of tarsus II: tically across the phenogram at the 0.09 (0) All setaceous (1) One spinelike level defines eight clusters denoted as (2) One peglike A-H. It should be noted here that the (3) Two spinelike primary interest in the phenogram is in (4) Two peglike the grouping of taxa rather than the (5) One spinelike and one peglike (6) Three spinelike relative levels at which taxa and clusters (7) Three peglike link with each other. Also, it should be (8) Two spinelike and one peglike kept in mind that the vertical ordering (9) One spinelike and two peglike is significant, i.e., (10) Four peglike of taxa and clusters not 29 Form of anterior seta of coxa III: each cluster may be rotated on its hori- (1) Setaceous zontal axis by 180 degrees without alter- Spinelike (2) ing any relationships. (3) Peglike 30. Form of posterior seta of coxa III: Generally, the correlation between the |

2 118 GREAT BASIN NATURALIST Vol. 36, No.

0.84 -0.21 -0.06 0.09 0.34 0.54 —I -H —I I —I L. transvaalensis —

L. keegani

L. simillimus

L. parasimillimus n.sp.

L. congoicola

L. roubaudi

L. grenieri

L. lavoipierre

L. moucheti

L. bocquieri paraspinosus L. B L. kochi

L. malacomys n.sp

L. spinifer

L. kampale sis

L. tillae

L. peregrinus 1

L peregrinus 2

L. thamnomys

L. fritzumpti

L. lavieri

L. nigerlensis

L. nuttalli

L. liberiensis

L. lamborni

L. setzeri

L. benoiti

L. algerlcus

L. aethiopicus

L. brandbergensis

L. acomys n. sp. F

L. oraniensis

L. zumpti

L. brazzai G

L. myoniys n. sp.

L. vansonereni

L brevlperitremus

-0.21 0-24 0.39 0.84

rorrelatioii matrix Fig. 1. Phenogram obtained from UPGMA cluster analysis of standardized based on 159 diararters by 37 OTl' data matrix.

two classification arrangements is cjuite gram, but with two taxa (/.. laroipierrei good, es])ecially in the similarity between and L. roubaudi) of the nuttalli group in- taxa. The first six taxa (L. transvaalensis. cluded also. Of the eight taxa in cluster L. keegani, L. congoicola, L. parasimilli- A. /.. simillimus and L. parasimillimus are mus n. sp., L. simillimus. and L. grenieri) the most similar. Based on the numerical of the a priori defined simillimus group taxonomic analysis (phenogram) alone. formed the first cluster (A) of the pheno- one might be tempted to synonymi/.e these June 1976 HERRIN, TIPTON: AI-"RICAX I,.\1-,I.APS 119

Table 4. Preliminary proposed classification phonogram L. ruouchvti was included in Laelaps s. str. l)ase(I on classical of the genus chisU^r A and most similar to L. trans- taxonomic methods prior to the numerical taxo- raalcnsis. T.. tnalacornys n. sp., nomic analysis. which was originally placed in the inittaUi group, is simillimus group inc hided in cluster B of the phenogram L. transvaalcnsis '/unipt, lOiO L. kecpani Thiu-maii. 10i8 (= L. berlesi Kee- wilh L. spinifer, most likely because of the many peglike and sj)inelike leg and body setae, characteristics which are more typi- cal of taxa of the vansomereni group. It should be noted that /.. kochi of cluster B and L. algericus and L. oraniensis of clusters E and F were included in the numeri- L. mouclicti Taufflieh. 1959 cal taxonomic analysis, even though they L. lavicri Taufflieh. 1954 (= L. nigeriensis Keegan. 1962) are not indigenous to the Ethio})ian region, L. fritzumpti Tauffliel). 1964 because of their similarity to taxa from

luittalli group this region and because their distribution L. lavoipicrrei Taufflieh. 1954 includes northern Africa (Palearctic re- L. tillar Taufflieh. 1959 gion , which borders the Ethiopian re- L. roubaudi Taufflieh. 1954 ) L. percgrinus Taufflieh. 1959 gion on the north. L. nuttaUi Hirst, 1915 The four remaining taxa of the a priori L. malacomys n. sp. defined simillimus group (L. thamnomys, L. brandbergcnsis Taufflieh. 1959 L. kampalensis, L. lavieri, and L. fritz- L. zumpti Keegan. 1956 L. myomys n. sp. umpti) were divided between two closely L. liberiensis Hirst, 1925 (= /.. lamborni Hirst. related phenogram clusters (C and D). 1925) L. kampalensis appears in cluster C with L. setzeri Coffev. 1971 L. tillae and L. peregrinus. two taxa of the L.brazzai Taufflieh, 1962 L. benoiti Taufflieh, 1964 nuttalli group. Cluster D consists of the vansomereni group remaining three taxa, L. thamnomys, L. L. aethiopicus Hirst. 1925 fritzumpti, and L. lavieri. Prior to the L. vansomereni Hirst, 1923 numerical taxonomic analysis, L. lavieri L. acomys n. sp. and L. nigeriensis were determined to be L. bncquieri Taufflieh, 1962 L. spinifer Taufflieh. & Mouchet. 1956 synonyms, and this seems to be confirmed L. paraspinosus Tipton. 1960 ( = L. parvulus by their placement in the phenogram. Hirst. 1923) Cluster E contains six taxa of the a L. breviperiiremus CGaiTett & Strandtmann. priori defined nuttalli group plus L. aethio- 1967) picus of the vansomereni group. Prior to two taxa, bvit there are several quite dis- the numerical taxonomic analysis, L. lib- tinctive morphological key characters sep- eriensis and L. larnborni were determined arating them (see identification key). to be synonymous, and this seems to be The placement of L. lavoipicrrei and L. confirmed by their placement in the roubaudi in cluster A of the phenogram is phenogram. The close phenetic relation- based on overall jihenetic resemblance ship between L. liberiensis. L. setzeri, L. (i.e., 159 characters weighted equally) benoiti. and L. algericus was confirmed by rather than consideration of several key the rmmerical taxonomic analysis. The characters as in the classical taxonomic low level at which L. aethiopicus joins arrangement. In the phenogram L. mou- cluster E poses some question on its actual cheti is included in cluster B, which in- phenetic resemblance with the nuttalli cludes taxa of the a priori vanf^omereni group. group. Tn examining the basic data ma- Clusters F and G consist of all remaining trix, it appears that L. rnoucheti and L. taxa of the previously defined nuttalli bocquicri are phenetically similar in char- grou[) j)lus L. acomys n. sp., which was acters related to length of setae, especialh' originally placed with the vansomereni dorsal body setae. This was supported by group. These two clusters join together be- an additional numerical taxonomic anal- fore either joins with any other cluster, ysis in which the number of characters thus confirming the ])henetic relationship was reduced to 105, eliminating highly among the five taxa involved. The final

correlated characters, especially those re- cluster (H ) consists of L. vansomereni and lated to length of setae. In the resulting L. breviperitremus. In the previously 120 GREAT BASIN NATURALIST Vol. 36, No. 2 noted numerical taxononiic analysis based and setaceous proximal seta of coxa I, on 105 characters, L. vansomereni joined with but two exceptions: L. kampalensis L. aethiopicus prior to their inclusion in bears a setaceous seta both proximally and a particular cluster, thus giving some distally on coxa I, and L. aethiopicus bears validity to the a priori defined close a blunt, peglike seta both jiroximally and phenetic relationship between these two distally on coxa I. Major group III con- taxa. tains taxa bearing a blunt, peglike seta both proximally and distally on coxa I.

Systematics of the Genus Laelaps s. str. Table 5. Final proposed classification of the After critical study of the numerical genus Laelaps s. str. taxonomic analyses and close examination Major gioup I of as many specimens of each taxa as were Subgroup A available, a final proposed classification L. similUmus Zuinpt, 1950 of the genus Laelaps s. str. was prepared L. parasirnillimus n. sp. (Table 5). The taxonomic groupings in L. keegani Thurman. 1958 L. trnnsvaalensis Zumpt. 1950 this proposed classification are based pri- L. congoicola Taufflieb. 1959 marily on overall phenetic resemblance L. lavoipierrei Taufflieb, 1954 as determined by both classical and nu- L. grenieri Taufflieb. 1954 merical taxonomic evaluations. We found Subgrou]! B L. lavieri Taufflieb. 1954 no set of key characters which may be L. frilziimpti Taufflieb. 19fi4 used to completely and definitively sep- L. thnmnomys Taufflieb. 1954 arate all these groups, especially the sub- T.. rnnucheti Taufflieb. 1959 groups. The 31 taxa described in this paper Major group II Subgroup are treated in the same order as listed in A L. kampalensis Taufflieb. 1959 Table 5. following identification key The T.. lillae Taufflieb. 1959 reflects to some degree the phenetic rela- /,. peregrinus Taufflieb. 1959 tionships between most taxa as presented L. roubaiidi Taufflieb. 1954 Subgroup B in the proposed classification; however, r,. nuttalli Hirst. 1915 is it should be kept in mind that the key /,. aethiopicus Hirst. 1925 based on sets of diagnostic qualitative r.. liberiensis Hirst, 1925 characters, whereas the proposed classifi- /., setzrri Coffey. 1971 L. henoiti Taufflieb, 1964 is phenetic cation based more on overall Subgroup C

resemblance. L. hrandbergensis Taufflieb. 1 959 The taxa of the three major grou]is T.. zumpti Keegan. 1956 L. hrazzai Taufflieb, 1962 (I, II and III) of the proposed classifica- L. rnyomys u. sp. tion are separated primarily on the form L. malacomys n. sj). of the proximal and distal setae of coxa Major group III I. The taxa of major group I, except for Subgioup A L. vansomereni Hirst. 1923 L. lavoipierrei, may be distinguished by L. acomys n. sp. both coxa I setae being setaceous. The dis- Subgroup B tal seta of coxa I of L. lavoipierrei is very L. spinifer Taufflieb and Mouchow 1956 small but bhmt and peglike. The taxa of L. paraspinosus Tipton. 1960 T.. hoc(juirri Tauffliel), 1962 major group II may be basically distin- L. hreriperitrcmus (Gairett and guished by the blunt, peglike distal seta Strandtiuann. 1967)

Key to Species of Laelaps from Small Alannnals of Africa (Females)

1. Distal seta of coxa I setaceous 2 Distal seta of coxa I spinelike or peglike 1-

2(1). Tarsi II and III with all pnvjpical setae s(Ma((>ous oi- at most \\itli one spinelike ^ At least one blunt, peglike [)reapical seta on tarsi II and III K'

3(2). Smaller species, dorsal plate less than 575 /i long 1 Larger species, dorsal plate greater than 575 n long *• June 1976 herrin, tipton: African laelaps 121

4(3). Anal plate distinctly longer than wide; adanal sotae short, length no greater than length of anal orifice; proximal seta of coxa I long, almost twice as long as distal seta (Figs. 14-17) L. transvaalensis Zumpt Anal plate as wide as or wider than long; adanal setae distinctly longer than length of anal orifice; proximal seta of coxa I not unusually long 5

5(4). Posterior seta of coxa II long, setaceous or spinelike, never blunt and peglike; posterior seta of coxa III short and spinelike (Figs. 8-11) L. keegani Thurman Posterior seta of coxae II and III always blunt and peglike 6

6(5). Posterior margin of sternal plate moderately invaginated, at least to level of setae st. 3; first sternal setae long, extending distinctly beyond posterior margin of sternal plate (Figs. 20-21) L. congoicola Taufflieb Posterior margin of sternal plate only slightly, if at all, invaginated; first sternal setae shorter, not extending near to posterior margin of sternal plate 7

7(6). Adanal setae rather short, not extending to base of postanal setae; unarmed venter bearing more than 10 })airs of rather short setae adjacent to genital and anal plates (Figs. 6-7) L. parasimillimus n. sp. Adanal setae longer, extending to or beyond base of postanal seta; unarmed venter bearing less than 10 pairs of medium-length setae adjacent to genital and anal plates 8

8(7). Metapodal plates rather narrow elongate; distance between 2nd genital setae distinctly less than distance between 3rd; sternal plate length /width ratio less than .75 (Figs. 2-3) L. similUmus Zumpt Metapodal plates irregularly oval, not so narrow and elongate; distance between 2nd genital setae greater than or equal to distance between 3rd; sternal plate length/ width ratio greater than .75 (Figs. 28-29) L. grenieri Taufflieb 9(3). Posterior margin of sternal plate only slightly invaginated medially; greatest width of genital plate at level of 3rd genital setae; distance between 1st genital setae equal to or less than distance between 4th genital setae (Figs. 43-44) L. thamnomys Taufflieb Posterior margin of sternal plate moderately invaginated, to or slightly beyond level of 3rd sternal setae; greatest wddth of genital plate at level of 2nd genital setae; distance between 1st genital setae distinctly greater than distance between 4th genital setae (Figs. 54-55) L. kampalensis Taufflieb

10(2). Dorsal plate with 38 pairs of rather small setae, especially more cen- trally located setae, setae px3 absent; anterior margin of anal plate rounded; medial hypostomal setae short, extending no further than

half fhstance to gnathosomal setae (Figs. 47-51) .... L. mouchcti Taufflieb Dorsal })late with usual 39 pairs of rather long setae; anterior margin of anal plate straight or slightly concave; medial hypostomal setae longer, extending distinctly further than half distance to gnathosomal setae 11

11(10). Peritreme longer, extending anteriorly to middle or posterior of coxa I; tarsi II and III each bear one blunt, peglike preapical setae, and

tarsus IV with no blunt preapical setae (Figs. 30-33) .. L. lavieri Taufflieb Peritreme short, extending only to level of middle of coxa II; tarsi II, III, and IV each bear two or more blunt, peglike preapical setae (Figs. 36-40) L. frhzumpti Taufflieb 122 GREAT BASIN NATURALIST Vol. 36, No. 2

12(1). Proximal seta of coxa I setaceous and elongate 13 Proximal seta of coxa I robust, short, and sjjinelike or peglike 25

13(12). Tarsi II, III, and IV with preapical setae setaceous or at most one

spinelilce - 14 Tarsi II, III, and IV with one or more blunt, peglike preapical .... setae 17

14(13). Distal seta of coxa I small, slender yet blunt and peglike; proximal seta of coxa I slender, setaceous; posterior margin of sternal plate

only slightly invaginated, no further than level of 3rd sternal setae .... 15 Distal seta of coxa I large, robust, blunt, and ]jeglike; proximal seta of coxa I long and somewhat enlarged, almost elongate spinelike; posterior margin of sternal plate moderately invaginated, distinctly beyond level of 3rd sternal setae 16

15(14). Distance between 1st genital setae distinctly less than distance between 4th, and distance between 2nd distinctly less than distance between 3rd; distal seta of coxa I very small, blunt, and peglike (Figs. 24-25) L. Icwoipierrei Taufflieb Distance between 1st genital setae distinctly greater than distance between 4th, and distance between 2nd greater than distance between 3rd; distal seta of coxa I not small (Figs. 56-57) L. tillae Taufflieb

16(14). Distance between 2nd genital setae subequal to distance between 3rd; smaller species, dorsal plate length less than 600 /x; peritreme extends anteriorly to level of middle of coxa I (Figs. 62-63) L. roubaudi Taufflieb Distance between 2nd genital setae distinctly greater than distance between 3rd; larger species, dorsal plate length greater than 600

/.(.; peritreme extends anteriorly to level of middle of coxa II (Figs. 60-61) L. peregrinus Taufflieb

17(13). Peritreme extends anteriorly to near middle of coxa I 18 Peritreme extends anteriorly to near middle of coxa II 19

18(17). Gnathosomal setae slender, medium length, and setaceous; all ventral setae of leg I slender, setaceous; adanal setae of moderate length (Figs. 66-70) L. mitlalli Hirst Gnathosomal setae long, robust, and almost spinelike; some ventral setae of leg I short, robust, and spinelike or peglike; adanal setae short (Figs. 119-123) L. myomys n. sp.

19(17) Seta pd 1 of femur I unusually long, nearly two times as long as ad 1 seta; greatest width of genital plate at level of 2nd genital setae; distance between 1st genital setae usually greater than distance between 4th (Figs. 98-102) L. hrandhergcnsis Taufflieb Seta pd 1 of femur I not unusually long, no more than 1.5 times as long as ad 1 .seta; greatest width of genital plate at level of 3rd genital setae; distance between 1st genital setae equal to or less than (hstauce between 4th 20

20(19). Adanal setae short, not reaching to base of postanal seta; posterior margin of sternal plate moderately in\ agiiialed, to or beyond level of ^rd sternal setae; distance between 2nd genital setae equal to distance between 3rd (Figs. 105-109) L. zumpti Keegan Adanal setae longer, extending to or beyond base of postanal seta; posterior margin of sternal plate oidy sliglitly invaginated or with moderate invagination medially between pair of posterior projections, not invaginated near- to lev(^l of ^rd sternal setae; distance June 1976 herrin, tipton: ai-rican laelaps 123

between 2nd genital setae distinctly less than distance between 3rd 21

21(20). Posterior margin of stenial plate only slightly invaginated, with rather small pair of posterior projections, if at all 22 Posterior margin of sternal plate with slight to moderate invagination hetweon pair of prominent posterior j)rojections 24

22(21). Tarsus II with three blunt, peglike j)reaj)ical setae; tarsus III with four to five blunt, peglikc setae, two of which are j)reapical; distal seta of coxa I more robust and enlarged; metapodal plates more elongate; smaller species (Figs. 126-130) L. malacomys n. sp. Tarsus II with only two blunt, peglike preapical setae; tarsus III with two to three blunt, peglike setae, one of which is preapical; distal seta of coxa I not so enlarged; metapodal plates more oval or triangular; larger species 23

23(22). All dorsal setae long to medium in length, setae J5 extending to or beyond posterior margin of dorsal plate (Figs. 77-81) L. liberiensis Hirst Anterior and all marginal setae long to medium in length, but posterocentral setae rather small, setae J5 short, not reaching even to level of setae Z5 (Figs. 84-88) L. setzeri Coffey

24(21). Sternal j)late distinctly wider than long, posterocentral dorsal setae shorter, setae .14 not reaching near to level of J5, and J5 extending no further than posterior margin of plate (Figs. 112-116) L. brazzai Taufflieb Sternal plate approximately as long as wide; all dorsal setae rather long, setae J4 extending almost to level of setae J5, and J5 extend-

ing beyond posterior margin of plate (Figs. 91-95) .. L. benoiti Taufflieb

25(12). Gnathosomal setae setaceous, never robust and spinelike or peglike 26 Gnathosomal setae robust, spinelike or peglike 28

26(25). First sternal setae long, extending beyond posterior margin of sternal plate, well beyond level of setae st. 3; seta pd 1 of femur I shorter than sternal setae; adanal setae slender, setaceous (Figs. 73-76)

- L. aethiopicus Hirst First sternal setae shorter, not extending to posterior margin of sternal plate or near to level of setae st. 3; seta pd 1 of femur I as long as or longer than sternal setae; adanal setae rather robust and spinelike 27

27(26). Anterior margin of sternal plate only slightly arched, posterior margin only slightly invaginated; anal plate rounded, slightly wider than long (Figs. 133-137) L. vansomereni Hirst Anterior margin of sternal plate strongly arched, posterior margin deeply invaginated; anal plate elongate, distinctly longer than wide (Figs. 140-143) L. acomys n. sp.

28(25). Lateral hypostomal setae robust, slightly recurred, and peglike; anterior seta of coxae II and III robust, spinelike or peglike; peritreme extends to anterior of coxa I; dorsal plate with only 31 pairs of mostly minute setae (Figs. 157-161) L. bocquieri Taufflieb Lateral hypostomal setae setaceous; anterior seta of coxae II and III setaceous; peritreme extends no further than anterior of coxa II; dorsal plate with at least 37 pairs of medium to large setae 29

29(28). Anal plate broadly triangular, considerably wider than long; adanal setae slender, setaceous; posterior margin of sternal plate irregular- 124 GREAT BASIN NATURALIST Vol. 36, No. 2

ly straight to very slightly iiivaginated; all setae of trochanters I and II setaceous; dorsal seta J5 very small (Figs. 146-149) L. spinifer Taufflieb and Mouchet Anal plate not unusually wide; adanal setae robust and spinelike or peglike; dorsal seta J5 longer, extending to or beyond posterior margin of dorsal plate 30

30(29). Dorsal plate with 37 pairs of setae, most medium length and setaceous, with posterior and lateral marginal setae long; all ventral plate setae rather long and setaceous; more than 50 setae on unarmed opisthosoma (Figs. 150-154) L. paraspinosus Tipton Dorsal plate with 39-40 pairs of setae, anterior two-thirds robust and spinelike, posterior one-third long and setaceous; sternal and first three genital setae short, robust, and spinelike; many less than 50 setae on unarmed opisthosoma (Figs. 164-165) L. breviperitremus (Garrett and Strandtmann)

Major Group I 3rd sternal setae; setae st. 1 of moderate length, reaching to level halfway be- This major division of the genus is tween setae st. 2 and st. 3. Anterior flap characterized by both proximal and dis- of genital plate not overlapping posterior tal setae of coxa I being setaceous, ex- margin of sternal plate; distance between cept for L. lavoipierrei in which the distal 1st genital setae slightly greater than dis- seta is very small and slender, yet blunt tance between 4th genital setae, and dis- and peglike. This group is further di- tance between 2nd genital setae slightly vided into two subgroups based primarily less than distance between 3rd genital on the form of the preapical setae of tar- setae; greatest width of genital plate at sus II. level of 3rd genital setae. Anal plate Subgroup A roundly triangular, as wide as or wider than long, wath anterior margins rounded; This subgroup consists of 7 taxa: L. si- adanal setae of moderate length, extend- millimus, L. parasirnillimus. L. keegani, L. ing to or slightly base of postanal transvaalensis, L. congoicola, L. lavoipier- beyond seta; adanal setae set at level near pos- rei, and L. grenieri. All taxa of this sub- terior of anal orifice. ven- group are characterized by having the pre- end Unarmed ter bears approximately nine })airs of apical setae of tarsus II setaceous or at setaceous setae, four pairs adjacent to most somewhat spinelike, but never blunt genital approximately and peglike. These taxa clustered quite and anal plate plus five pairs closely in the numerical taxonomic anal- near or on posterior lateral plates ysis. body margins; metapodal rather elongate. Peritreme extends to level of

middle of coxa I. Dorsal plate bears 39 Laelaps (Laelaps) simillimus Zumpt pairs of setaceous setae; most dorsal setae Figs. 2-5 of moderate length, length slightly less Laelaps simillimus Zumpt, 1950, S. Afr. J. Med. thati (Hstaiu e between adjacent setae; sub- Sci. 15:81 (Holotype: Johannesburg, Trans- terminal setae (J5) reaching at least to vaal, South Africa; South African Institute for Medical Research., .Johannesburg); Tipton, level of base of setae Z5. Twelve to 16 1960, Univ. Calif. Publ. Ent. 16(6):282; pairs of setae border dorsal o])isthosoma Zumpt, 1961. Publ. S.Afr. Inst. Med. Res. on soft integument. Both proximal and 9(1):29. distal setae of coxa I setaceous, |)roximal Description.— Female-. (Figs. 2-3) seta somewhat longer than distal seta;

Dorsal plate length 456 /., width 262 /x. setae pd 1 and ad 1 of femur I relatively Gnathosomal and hypostomal setae se- short and subequal in length; anterior taceous; medial hypostomal setae long, seta of coxae II and III and seta of coxa reaching almost to or slightly beyond base IV setaceous; posterior seta of coxae II f)f gnathosomal setae. Posterior margin of and III moderately robust, blunt, and sternal plate slightly invaginated, invagi- peglike; preapical setae of tarsi II and III nation reaching no fnrlhor than level of niosth' setaceous; however, one or two June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 125

Figs. 4-5. Laelaps simillimus Zumpt, male. (4) venter; (5) dorsum, scale = 100/u. 126 GREAT BASIN NATURALIST Vol. 36, No. 2

setae may be spinelike; all other leg South Africa (Pretoria); Zumpt Collection .(AMP) setae setaceous and normally (ieveloped. South Africa; 27 coll. (64 females. 2 males); AMP Male: (Figs. 4-5) Gnathosomal and Botswana; 1 coll. (1 female); AMP hypostomal setae setaceous, with medial Rhodesia; 1^ coll. (47 females, 1 male); hypostomal setae of moderate length but AMP not reaching to base of gnathosomal setae. Aethomys namaquensis South Africa (Transvaal); 32 females, Ventral setae, except adanal and postanal 2 males (type specimens); Zumpt, 1950 setae, of moderate length, each extending South Africa (Kamanjab); Tipton, 1960 in length beyond base of seta immediate- Lemniscomys griselda ly posterior; holoventral plate broad be- Rhodesia; 1 coll. (1 female); AMP South Africa; 1 coll. (1 female); AMP tween coxae II and III, narrowing con- Lophuromys aquilus siderably between coxae IV and expanded Congo-Leopoldville; 7 females; Taufflieb, considerably posterior to coxae IV; ex- 1964 panded area between genital setae and Angola (Dundo); 7 females; Taufflieb. 1962 Mastomys natalensis anal orifice bears five pairs of setaceous South Africa; 1 coll. (1 female); AMP setae; adanal setae of moderate length ex- South Africa (ORS); 15 coll. (21 females, tending slightly beyond base of postanal 2 males, 3 ny) ; AMP seta; adanal setae set slightly posterior Rattus sp.

South Africa (Transvaal); 1 female; to middle of anal orifice; postanal seta Taufflieb. 1964 somewhat more robust and longer than Rhabdomys pumilio adanal setae. Metapodal plates inappar- South Africa; 3 coll. (5 females. ent; unarmed venter bearing 12 to 14 1 male, 1 ny); AMP pairs of setae adjacent to holoventral plate. Unknown Rhodesia; 1 coll. (1 female); AMP Peritreme extends to level of middle or an- South Africa; 15 coll. (19 females. terior of coxa I. Dorsal plate bearing 39 1 male); AMP pairs of setaceous setae; length and position of setae as in female. Both proximal Remarks.— L. simiUimus closely re- and distal setae of coxa I setaceous, proxi- sembles most other taxa of subgroup A, mal seta about 1.5 times as long as distal differing in several distinguishing charac- seta; setae pd 1 and ad 1 of femur I rel- ters. It differs from L. parasiniiUinius in atively short with seta pd 1 somewhat bearing only a few medium-length setae longer than ad 1 ; anterior setae of coxae ventrally adjacent to the genital and anal II and III, posterior seta of coxa II, and plates, in the longer adanal setae, and in seta of coxa IV all setaceous; posterior seta the genital plate which is somewhat more of coxa II rather long and setaceous, slender posteriorly. L. shuillimus may whereas posterior seta of coxa III spine- be separated from L. grenieri by the nar- like; preapical setae of tarsi II and III rower more elongate metapodal plates, mostly setaceous; however, one or two on the smaller length width ratio of the ster- each may be spinelike; all other leg setae nal plate (less than .75), and by the dis- setaceous and normally developed. tance between the 2nd genital setae being distinctly less than that between the 3rd COLLF.CTION RECORDS genital setae. It may be easily separated Klephantulus intufi from L. keegoni by the blunt, jieglike South Africa; 1 coll. (1 female); AMP"* seta po.steriorly on coxae II and III, and Macroscelides proboscideus /.. significantly South Africa (ORS"*); 1 coll. (1 female); from transvaalcnsis by the AMP longer adanal setae, the broader anal plate, Tadarida midas and the generally more elongate dorsal South Africa; 2 (oil. (2 females); AMP Tatera afra plate. In L. congoicola the 1st sternal South Africa; Zumpt, 1961 setae are longer, reaching to or beyond Southwest Africa; Zumpt, 1961 the moderately invaginated posterior mar- Rhodesia; Zumpt. 1961 gin of the sternal plate. As noted jirevious- Tatera Irurngastrr ly, L. laroipirrn'i (Hffers in the very South Africa; 1 coll. (H females); AMP Aethnmys chryxophilux small, blunt distal seta of coxa I. South Africa; Zumpt. 1961 L. sifnillirnus has iieeii collected almost Southwest Africa; Zumpt, 1961 Rhodesia; Zumpt, 1961 exclusivelv from southern Africa, reach-

'AMP = African Mnnininl I'lojcd. ^ORS = Oiiingp Rivor Survey. June 1976 HERRIN. TIPTON: AI-RICAN LAELAPS 127 ing no further north than Angola and tance between 2nd genital setae distinctly Congo-LeopoldA ille. It is recorded from less than distance between 3rd genital a variety of small mammals, but pri- setae; greatest width of genital j)late at marily from species of Actliornys, and most level of 3rd pair of genital setae. Anal frequently from Aethomys chrysophilus. jilate roundly triangular in shape, approximately as wide as long with rounded margins; adanal setae very short and Laelaps {Laclaps ) parasiniiUinius. n. sp. small, length approximately equal to Figs. 6-7 width of anal orifice; adanal setae set at Holotype. female; Type locality: 10 mi. WNW level of posterior third of anal orifice; Ivory Coast; in U.S. National Soubre, Mu postanal seta rather small but somewhat seum, Washington. D.C. larger than adanals and somewhat more Description.— Female: (Figs. 6-7) robust. Unarmed venter bearing ap-

Dorsal plate length 405 /^i, width 244 /i. proximately 13 pairs of setae, medial 3 Gnathosomal and hypostomal setae se- pairs adjacent to genital and anal plates taceous; median hypostomal .setae of me- longer with lateral pairs much shorter, (liimi length, not reaching base of gna- some being almost spinelike; metapodal thosomal setae. Posterior margin of sternal plates small, elongate oval. Peritreme ex- plate very slightly invaginated medially; tends to level of middle or anterior of setae st. I of moderate length, reaching coxa I. Dorsal plate bearing 39 pairs of about one-third distance between 2nd and setaceous setae; most dorsal setae of me- 3rd sternal setae; sternal setae as well as dium length, length distinctly less than 4 pairs of genital setae of moderate length distance between adjacent setae; sub- and rather slender. Anterior flap of geni- terminal setae (5) rather small and slen- tal plate overlapping posterior margin der, reaching no further than level of base of sternal plate to or slightly anterior to of terminal setae; terminal setae consider- level of 3rd sternal setae; distance be- ably larger and more robust than all other tween 1st genital setae distinctly less than dorsal setae. Six to 8 pairs of rather small distance between 4th genital setae; dis- spinelike setae border dorsal opisthosoma

r- \

Figs. 6-7. Laelaps parasimUUmus n. sp.. female. (6) venter; (7) dorsum, scale = 100//. .

128 GREAT BASIN NATURALIST Vol. 36, No. 2 on soft integument. Proximal seta of coxa lapping posterior of sternal plate; dis- I of moderate length and robust, distal tance between 1st pair of genital setae seta of coxa I quite small and slender; slightly greater than distance between 4th setae pd 1 and ad 1 of femur I subequal in genital setae; in type specimens, greatest length; anterior seta of coxae II and III width of genital plate at level of 3rd and seta of coxa IV rather small, seta- genital setae; however, in specimens from ceous, although somewhat enlarged ba- Nigeria greatest width of genital plate at sally; posterior seta of coxae II and III level of 2nd pair of genital setae. Posterior rather small but robust and peglike; pre- margin of genital plate truncate as well as apical setae of tarsi mostly setaceous as anterior margin of anal plate; anal plate well as all other leg setae. width subequal to length; adanal setae Male: Unknown. slender reaching to base of postanal seta; adanal setae set at level near posterior Type material end of anal orifice; postanal seta robust, Dephomys defua slightly longer than adanal setae. Un- Ivory Coast (10 mi WNW Soubre); armed venter bearing about 16 pairs of female holotype and 1 deutonvmph (LWR 1471). setaceous setae, 4 pairs adjacent to genital Malacomys longipes and anal plates with others on posterior Ivory Coast (10 mi WNW Soubre); 2 lateral and lateral body margins; meta- females (LWR 1478). podal plates rather elongate and narrow. Remarks.— L. parasimillimus may be Peritreme extending to middle of coxa I. distinguished from all other closely related Dorsal plate bearing 39 pairs of setaceous taxa by the following combination of char- setae; most dorsal setae of moderate length, acters: more than 10 pairs of mostly small length equal to distance between adjacent setae ventrally adjacent to genital and setae; subterminal setae (J5) reach to or anal plates; rather wide genital plate at slightly beyond posterior margin of dorsal level of 3rd genital setae; short adanal plate. Approximately 18 pairs of setaceous setae; distal seta of coxa I much shorter setae border dorsal plate on soft integu- than proximal seta; small peglike seta ment. Both proximal and distal setae of posteriorly on coxae II and III; rather coxa I setaceous, with proximal seta 1.5 broad anal plate; and greater length of times as long as distal seta; setae pd 1 and sternal plate. ad 1 of femur I relatively short, subequal This new species has been collected in length, with ad 1 seta only slightly only from Ivory Coast on Dephomys longer; anterior seta of coxae II and III, defua and Malacomys longipes. posterior seta of coxa II, and seta of coxa IV setaceous; posterior seta of coxa III Laelaps (Laelaps) keegani Thurman rather small and peglike (bluntly spine-

like) ; tarsus II with one bluntly spinelike Figs. 8-13 preapical seta; all other leg setae seta- Laelaps berlesi Keegan, 1956, J. Egypt. ceous. Pbl. HIth. Assoc. 31 (6):264-265 (Holotype: Pyramids, Giza, Egypt; Male: (Figs. 12-13) Gnatho.somal and U.S. National Museum. Washington, hypostomal setae setaceous; medial hypo- D.C.). setae of length, Laelaps keegani Thurman, 1958, Ent. Soc. stomal moderate reaching Wash. 60 (2): 74. Papema, Furman, almost to base of gnathosomal setae. Ven- and Rothstein, 1970, Rev. Zool. tral setae, except adanal and postanal Bot. Afr. 81 (3-4): 330-336 (Host .setae, of moderate length and slen(ler, each locality) extending slightly beyond base of adja- Description.-— Female: (Figs. 8-11) cent posterior seta; holoventral plate

Dorsal plate length 505 //., width 301 //.. rather broad between coxae II and III, Gnathosonial and hypostomal setae se- greatly narrowing between coxae IV, and taceous; medial hypostomal sotae of mod- greatly expandecl posterior to coxae IV; erate length but not reaching base of gna- expan(l(nl area between gejiifa! setae and thosonial setae. Posterior margin of sternal anal orifice bears 5 pairs of setaceous plate only slightly invaginated; setae si. s(>tae; adanal setae rather short, apj)roxi- 1 of moderate length, reaching to point niately equal to length of anal orifice; halfway between .setae st. 2 and st. 3. An- adanal setae set at level near middle of terior flap of genital [)late slightly over- anal orifice; postanal seta approximately June 1976 HERRIN, TIPTON: AFRICAN L.AELAPS 129

Figs. 8-11. Laelaps keegani Thurman, female. (8) venter; (9) dorsum, scale = lOOju; (10) ventral view of tarsus II; (11) ventral view of tarsus III, scale = 50/i. twice as long as adanal setae and somewhat more robust and spinelike; most leg what more robust. Metapodal plates in- setae setaceous and normally developed; apparent, apparently fused to lateral ex- however, some may be shorter and some- tensions of holoventral plate; unarmed what spinelike. venter bears approximately 10 to 12 pairs of setaceous setae adjacent to holoventral Collection records Atelerix albiventris plate; peritreme extends to middle of cox- Upper Volta; 1 coll. (1 female); AMP ae I. plate bears pairs of seta- Dorsal 39 Crocidura sp. ceous setae; length and position of setae Upper Volta; 1 coll. (3 females. 1 approximately as in female. Both proxi- male, 1 ny.); AMP Tatera kempi mal and distal setae of coxa I setaceous, Dahomey; 1 coll. (1 female); AMP proximal seta somewhat longer than distal Arvicanthis niloticus seta; setae pd 1 and ad 1 of femur I Egypt (Giza); holotype female; Keegan, 1956 subequal in length; anterior setae of Ghana (Accra-Tema) ; Paperna. et al.. 1970 Nigeria; Zumpt Collection (AMP) coxae II and III, posterior seta of coxae Ghana; 1 coll. (1 female); AMP II and III, and seta of coxa IV seta- Ivorj' Coast; 10 coll. (19 females. 1 male); ceous, wdth posterior seta of coxa III some- AMP Vol. No. 2 130 GREAT BASIN NATURALIST 36,

Figs. 12-13. Laelaps keegani Thurman, male. (12) venter; (13) dorsum, scale = 100/u.

Nigeria; 23 coll. (62 females, 3 Laelaps {Laelaps) transvaalensis Zumpt males, 7 ny.); AMP Figs. 14-19 Senegal; 3 coll. (18 females); AMP Mus musculoides Laelaps transvaalensis Zumpt. 1950, S. Afr. J.

Ghana; 1 coll. (4 females, Med. Sci. 15: 77-82 (Holotype: Krugersdorp,

1 ny.); AMP Transvaal, South Africa; So. Afr. Inst. Med. Felis lybica Res.. Johaimesburg) ; Taufflieb, 1959, J. Ent. Soc. So. Afr. 22(2):404-408 (key); Tipton, Upper Volta; 1 coll. (1 female); AMP Unknown 1960, Univ. Calif. Publ. Ent. 16(6): 262-264, 283-284 (key, synopsis); Zumpt. 1961, Publ. Nigeria; 2 coll. (13 females); AMP So. Afr. Inst. Med. Res. 4(1): 30 (host, locality). Remarks.— L. keegani is quite distinc- tive in one key character which is invari- Description.— Female: (Figs. 14-17)

ant in all specimens examined in this Dorsal plate length 460 /i, width 317 /a. study; this character is the setaceous or Gnathosomal and hypostomal setae seta- spinehke posterior seta of coxae II and III, ceous; medial hypostomal setae long, rather than blunt, peghke setae as in all reaching to base of gnathosomal setae. other closely related taxa. In addition to Posterior margin of sternal plate invagi- this character, L. keegani may be sep- nated to or slightly beyond level of setae arated from certain other taxa by the St. 3; setae st. 1 of moderate length, reach- longer adanal setae and the broad anal ing to point halfway between setae st. 2 plate. and St. 3. Anterior flap of genital plate L. keegani was originally recorded from overlapi)ing |)osterior of sternal plate; dis- Giza, Egypt, on Arvicanthis niloticus tance between 1st genital setae slightly Subsequently, it has been collected from but distinctly further apart than 4th the countries of northwest Africa south genital setae; greatest width of genital of the Sahara and almost exclusively from plate at or slightly behind level of 2nd Arvicanthis niloticus. pair of genital setae. Anal plate distinctly June 1976 HERRIN, TIPTON: AFRICAN LAEL.\PS 131

Figs. 14-17. Laelaps transvaalensis Zumpt, female. (14) venter; (15) dorsum, scale = lOO/i; C16) ventral view of tarsus II; (17) ventral view of tarsus III. scale = 50/u.

longer than wide; adanal setae short, plate; distance between setae J4 greater length less than distance from adanal than that between setae .15. Ten to 12 setae to postanal seta; adanal setae set pairs of setae border dorsal opisthosoma at level of middle of anal orifice; oval on soft integument. Both proximal and reticulate pattern near anterolateral mar- distal setae of coxa I setaceous, with proxi- gins of anal plate. Unarmed venter bear- mal seta 1.5 times as long as distal seta; ing about 10 pairs of setaceous setae, 4 setae pd 1 and ad 1 of femur I subequal pairs adjacent to genital and anal plates in length, with ad 1 seta only slightly plus 6 pairs near or on posterolateral body longer; anterior seta of coxae II and III margins; metapodal plates elongate oval. setaceous and enlarged somewhat basally; Peritreme extends to middle of coxa I. posterior seta of coxae II and III robust, Dorsal plate bearing 39 pairs of setaceous peglike (bluntly spiniform); tarsi II and setae; central and lateral setae of moderate III each with one spinelike preapical seta; length (length equal to distance between all other leg setae setaceous, some may be adjacent setae); setae il, rl, r2, r3, r4, heavier than others but not distinctly J4 and J5 shorter; subterminal setae (J5) spinelike. reaching to posterior margin of dorsal Male: (Figs. 18-19) Gnathosomal and 132 GREAT BASIN NATURALIST Vol. 36, No. 2

Figs. 18-19. Laelaps transvaalensis Zumpt, male. (18) venter; (19) dorsum, scale = 100^. hypostomal setae all setaceous, with me- seta nearly two times as long as distal dial hypostomal setae two times as long seta; setae pd 1 and ad 1 of femur I sub- as gnathosomal setae. Ventral setae, ex- equal in length; anterior seta of coxae cept adanals and postanal, of moderate II and III setaceous and enlarged basally; length, each extending in length beyond posterior seta of coxa II elongate and se- base of seta immediately posterior by taceous, whereas posterior seta of coxa about one-third its length; holoventral III short and ])eglike; tarsi II and III plate rather narrow between coxae IV each with one preapical spinelike seta, and considerably expanded posteriorly; ex- and all legs with some ventral short spine- panded area between genital setae and like setae. anal orifice bearing 5 pairs of setaceous CoLI,ECTION RECORDS setae; adanal setae short, length less than Crocidura sp. distance between adanals and postanal South Africa; 1 coll. (1 female); seta; postanal seta spinelike and at least Zumpt, 1961 two times as long as adanals. Metapodal Mncroscelides proboscideus plates elongate oval; unarmed venter bear- South Africa (ORS); 1 female; AMP Cryplotiiys hotlrntotus ing 2 pairs of setae adjacent to holoventral South Africa; 1 coll. (6 females); AMP plate plus about 5 pairs on posterolateral Prtionius typicus margins of body. Peritremo extends to South Africa (ORS); 1 female; AMP level of middle of coxa I. Dorsal plate Tatera Icucogastfi bearing 39 pairs of setaceous setae; length South Africa; 1 female; AMI' and position of setae as in female. Soft Aethonns chrysophilus

South Africa; 7 coll. ( 10 fenuiles. 7 integument of opisthosoma bearing about males); AMP 9 pairs of setae. Both proximal and distal Aclhonirs ritimacfuensis setae f)f coxa I setaceous, with j)roximal South .Africa (Transvaal); 1 -f coll. June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 133

Mastomys natalensis plate slightly invaginated, invagination South Africa; 1 female; Zumpt. 1961 reaching to level of third sternal setae; Myomys daltoni setae st. 1 long, reaching well beyond pos- Senegal; 1 female; AMP Rhabdomys pumilio terior margin of sternal plate. Anterior South Africa; 1+ coll.; Zum[)t. 1961 flap of genital plate overlapping posterior South Africa; 5 coll. (6 females, 4 margin of sternal plate slightly; distance males); AMP 1st genital setae slightly greater Saccostomus campestns between

South Africa; 1 female; AMP than distance between 4th genital setae, Otornys angoniensis distance between 2nd genital setae South Africa (ORS); 1 coll. (15 females, slightly greater than distance between 2 males); AMP 3rd genital setae; greatest width of genital Otomys irroratus South Africa (Transvaal); holotype and plate at le^el slightly anterior to 3rd geni- 4 females; Zumpt, 1950 tal setae. Anal plate roundly triangular, South Africa (Transvaal); 1+ coll.; Tipton. width equal to length, with anterior mar- 1960 gins rounded; adanal setae rather short, South Africa; 3 coll. (13 females); AMP Unknown host extending no further than base of post- South Africa (ORS); 2 coll. (5 females); anal seta; adanal setae set at level of pos- AMP terior end of anal orifice. Unarmed venter South Africa; 12 coll. (23 females, bearing about 8 to 10 pairs of setaceous 11 males); AMP setae, 4 pairs immediately adjacent to Botswana; 1 coll. (2 females) ; AMP genital and anal plates plus approximately Remarks. L. transvaalensis be — may 4 to 6 pairs near or on posterior lateral easily separated from other taxa by the body margins; metapodal plates oval to of plate, i.e., narrower anal form the anal elongate-oval, length about twice width. plate with short adanal setae and pair of Peritreme extends to level of anterior of dark areas laterally. Other distinguishing coxa I. Dorsal plate bearing 39 pairs of characters are as follows: dorsal plate setaceous setae; most dorsal setae of me- rather broad relative to length; peritremes dium length, length equal to or slightly posteriorly than usual for the wider greater than distance between adjacent genus; moderately emarginated posterior setae; subterminal setae (J5) reaching to of sternal plate; unusually long margin or slightly beyond posterior margin of dor- proximal seta of coxa I; and rather broad sal plate; terminal setae (Z5) much long- genital plate posteriorly. er than other adjacent setae. About 12 This species is recorded almost exclu- pairs of setae border dorsal opisthosoma sively from South Africa from a great on soft integument. Both proximal and variety of small mammal hosts. One col- distal setae of coxa I setaceous, proximal lection is recorded from Botswana and seta distinctly longer than distal seta; one, possibly erroneous identification, setae pd 1 and ad 1 of femur I subequal from Senegal. collections speci- More and in length; anterior seta of coxae II and mens have been collected from species of III and seta of coxa IV slender, setaceous; Otomys than from any other host. It is posterior seta of coxae II and III robust, anticipated that future records will reveal blunt and peglike, with posterior seta of a wider geographic distribution in much coxa II somewhat longer than that of southern Africa than the currently avail- coxa III; no robust, blunt preapical setae able records indicate. on tarsi II. III. or IV; however, one or two spinelike preapical setae may be pres- Laelaps (Laelaps) congoicola Taufflieb ent on tarsi II and III; most other leg Figs. 20-23 setae setaceous and normally developed.

Laelaps congoicola Taufflieb, 1959. J. Ent. Soc. Male: (Figs. 22-23) Gnathosomal and S. Afr. 22(2):397-398 (Holotype: Brazzaville. hypostomal setae setaceous, with medial Congo; Museum d'Histoire Naturelle, Paris). hyy)ostomal setae of moderate length, ex- Description.— Female: (Figs. 20-21) tending almost to base of gnathosomal

Dorsal plate length 538 /t, width 349 p.. setae. Ventral setae, except adanal and Gnathosomal and hypostomal setae seta- postanal setae, relatively long, each ex- ceous; medial hypostomal setae relatively tending well beyond base of seta im- long, reaching almost to base of gnatho- mediately adjacent; holoventral plate somal setae. Posterior margin of sternal rather broad between coxae II and III. 134 GREAT BASIN NATURALIST Vol. 36, No. 2

1 21

Figs. 20-21. Laelaps congoicola Taufflieb, female. (20) venter; (21) dorsum, scale = 100/i.

Figs. 22-23. Laelaps congoicola Tauffliol), male. (22) venter; (23) dorsum, scale == 100/i. June 1976 HERRIN. TIPTON: AFRICAN LAELAPS 135

narrowing considerably between coxae IV, DivscRiPTioN.— Female: (Figs. 24-25)

and greatly expanded immediately pos- Dorsal plate length 495 fx, width 315 ix. terior to coxae IV; expanded area between Gnathosonial and hypostomal setae seta- genital setae and anal orifice bearing 5 ceous; medial hy])ostomal setae of mod- pairs of setaceous setae; adanal setae rel- erate length, reaching almost to base of atively short, not extending to base of gnathosonial setae. Posterior margin of postanal seta; adanal setae set slightly sternal plate only very slightly invagi-

posterior to level of middle of anal orifice; nated medially; setae st. 1 of moderate postanal seta considerably longer than length, reaching to level approximately adanal setae and rather robust. Metapodal halfway between setae st. 2 and st. 3. plates inapparent, apparently fused to Anterior flap of genital plate overlapping lateral extensions of holoventral plate; un- posterior margin of sternal plate at least armed venter bearing 6 to 9 pairs of seta- to level of 3rd sternal setae; distance be- ceous setae, about 3 or 4 pairs set quite tween 1st genital setae subequal to dis- close to holoventral plate laterally. Peri- tance between 4th genital setae, distance treme extending to middle or anterior of between 2nd genital setae slightly less than coxa I. Dorsal ])late bears 39 pairs of seta- distance between 3rd genital setae; great- ceous setae; length and jjosition of setae as est width of genital plate at level of 3rd in female. Both proximal and distal setae I)air of genital setae. Anal plate roughly of coxa I setaceous, with proximal seta triangular in shape, as wide as long, with slightly longer than distal seta; setae pd 1 anterior margins concave or invaginated; and ad 1 of femur I subequal in length; adanal setae of moderate length, extending anterior seta of coxae II and III, posterior slightly beyond base of postanal seta; seta of coxa II and seta of coxa IV all seta- adanal setae set at level near posterior ceous; posterior seta of coxa III robust and end of anal orifice; postanal seta rather spinelike; no blunt preapical setae on robust and slightly longer than adanal tarsi II, III, or IV; however some pre- setae. Unarmed venter bearing 10 to 14 apical setae may be spinelike; most other pairs of setaceous setae, 4 pairs immedi- leg setae setaceous and normally de- ately adjacent to genital and anal plates veloped. plus approximately 6 to 8 pairs near or on posterior lateral body margin; metapodal Collection records plates elongate, considerably longer than Oenomys hypoxanthus wide. Peritreme extends to level of mid- French Congo (Brazzaville); 35 females (type specimens); Taufflieb. 1959 dle or anterior of coxa I. Dorsal plate bearing 39 pairs of setaceous setae; most Remarks. most distinguishing — The dorsal setae of medium length, length characters of L. congoicola. separating it almost equal to distance between adjacent from other closely related taxa, are the setae; subterminal setae (J5) reaching al- long St. 1 setae and the moderate posterior most to posterior margin of dorsal plate. invagination of the sternal plate, i.e., Nine to 12 pairs of setae border dorsal setae st. 1 extends to or beyond posterior opisthosoma on soft integument. Proximal margin of sternal plate. Other diagnostic seta of coxa I setaceous and of moderate characters are as follows: relatively broad length, distal seta of coxa I quite small, oval dorsal plate; inedium-length adanal blunt and peglike; setae pd 1 and ad 1 of setae; blunt, peglike seta on posterior of femur I subequal in length; anterior seta coxae II and III; and setaceous proximal of coxae II and III and seta of coxa IV and distal setae of coxa I. setaceous; posterior seta of coxae II and This taxon has been reported only from III robust, blunt, and peglike; no robust, Oenomys hypoxanthus in French Congo blunt preapical setae on tarsi II, III, or (Brazzaville). IV; however often with one spinelike seta on coxae II and III; most other leg setae setaceous and normally developed. Laelaps (Laelaps) lavoipierrei Taufflieb (Figs. Gnathosomal and Figs. 24-27 Male: 26-27) hypostomal setae setaceous; medial hypo- Laelaps lavoipierrei Taufflieb. 1954. Ann. Parasit. stomal setae of moderate length, reaching 29(4) :4+0 (HoloU-pe: Brazzaville. Congo; base of gnathosomal setae. Ven- Institut Pasteur. Paris); Tipton, 1960. Univ. almost to Calif. Publ. Ent. 16(6) :274. tral setae, except adanal and postanal 136 GREAT BASIN NATURALIST Vol. 36, No. 2

Figs. 24-25. Laelaps lavoipierrei Taufflieb, female. (24) venter; (25) dorsum, scale = 100;u.

Figs. 26-27. r.arlfi/)\- lar()if)irrrri Taiifflicl). male. (26) venter; (27) dorsum, scale =-- lOO/i. Tune 1976 HERRIN, TIPTON: AFRICAN LAELAPS 137 setae, rather long an(] slender, each ex- In this one character alone it resembles tending well beyond base of adjacent j)OS- taxa of major group II, but on the basis terior seta; holoventral })late rather broad of Dxcrall morphological characters it is between coxae II and III greatly narrow- placed in major group I near L. congoi- ing between coxae IV, and greatly ex- cola and L. grenieri. Another character panding posterior to coxae IV; expanded which may be used to separate L. lavoi- area between genital setae and anal orifice pierrei from the latter two taxa is the bearing 5 pairs of setaceous setae; adanal straight to concave shape of the anterior setae of medium length, extending to or margin of the anal plate, rather than a slightly beyond base of postanal seta; rounded, convex margin. adanal setae set at level near posterior This mite is parasitic primarily on end of anal orifice; postanal seta slightly r.ophurnmys sikapusi in northwest Africa longer than adanal setae and slightly more south of the Sahara. Single collections have robust. Metapodal plate inapparent, a])- been recorded from several other small parently fused to lateral extensions of mammals. holoventral plate; unarmed venter bearing approximately 8 to 10 pairs of setaceous Tjielaps (Laelaps) grenieri Taufflieb pos- setae, those more marginal and more Figs. 28-29 terior somewhat longer. Peritreme extends TMelaps grenieri Taufflieb. 1954. Aiui. Parasit. to level of coxa I. Dorsal plate bearing 39 29(4) :439 (Holotype: Brazzaville, Congo; pairs setae; length posi- of setaceous and Institut Pasteur, Paris); Tipton, 1960. Univ. tion of setae approximately as in female. Calif. Publ. Ent. 16(6): 270. Both proximal and distal setae of coxa I Description.— Female-. (Figs. 28-29) setaceous, proximal seta somewhat longer Dorsal plate length 515 /x, width 307 /<,. than distal seta; setae pd 1 and ad 1 of Gnathosomal and hypostomal setae seta- femur I subequal in length, both rather ceous; medial hypostomal setae of mod- short; anterior seta of coxae II and III, erate length, reaching at least half dis- posterior seta of coxa II, and seta of coxa tance to gnathosomal setae. Posterior mar- IV all setaceous; posterior seta of coxa III gin of sternal plate irregularly straight to short and spinelike; several preapical setae very slightly invaginated; setae st. 1 rel- of tarsi II and III rather robust and spine- atively long, reaching almost to level of like; most other leg setae setaceous and sternal setae. Anterior flap of genital normally developed; however, some may 3rd plate not overlapping posterior margin be shorter and somewhat spinelike. of sternal plate; distance between 1st geni- Collection records tal setae slightly greater than distance Hipposideros baetus betw^een 4th genital setae, distance be- Ivory Coast; 1 coll. female); (1 AMP tween 2nd genital setae slightl}' greater r.ophuromys sikapusi tlian distance between 3rd genital setae; Congo (Bi^azzaville) ; 3 females (type specimens); Taufflieb. 1954 greatest width of genital plate at level of Cameroon (Yaounde); Taufflieb. 1962 2nd pair of genital setae. Anal plate Ghana; 26 coll. (59 females, 3 males. roughly triangular in shape, as wide as 5 ny.); AMP long, with anterior margin almost straight; Ivory Coast; 21 coll. (104 females. 14 males, 3 ny.); AMP adanal setae of moderate length, extend-

Nigeria; 11 coll. (57-|- females. 1 male, ing somewhat beyond base of postanal 1 ny.); AMP seta; adanal setae set at level of posterior Mastomys natalensis end of anal orifice. Unarmed venter Ivorv' Coast; 1 coll. (1 female. 1 male); AMP bearing 6 pairs of setaceous setae, 4 pairs Upper Volta; 1 coll. (1 female); AMP immediately adjacent to genital and anal Mus tjiusculoides plates plus 2 pairs near or on posterior Ghana; 1 coll. (1 female); AMP lateral body margin; metapodal plates Praomys tullbergi elongate-oval. Peritreme e.xtends to level Ivory Coast; 1 coll. (1 fejnale. 6 ny.); AMP of anterior of coxa I. Dorsal plate bearing Uranomys ruddi 39 pairs of setaceous setae; most dorsal Ghana; 1 coll. (3 females); AMP setae of moderate length, length equal to Remarks.— L. lavoipierrei ma}' be easily or slightly greater than distance between separated from all other taxa by the very adjacent setae; subterminal setae (J5) small, blunt, peglike distal seta of coxa I. reaching onlv to level of base of terminal 138 GREAT BASIN NATURALIST Vol. 36, No. 2

Figs. 28-29. Laelaps grenieri Taufflieb. female. (28) venter; (29) dorsum, scale = 100;u. setae; terminal setae rather long and Lemniscomys barbarus slender. Fourteen to 18 pairs of setae bor- Ghana; 2 coll. (2 females); AMP Upper Volta; 2 coll. (10 females. der dorsal opisthosoma on soft integument. 1 ny.); AMP Both proximal and distal setae of coxa 1 Lemniscomys macculeus setaceous, proximal seta considerably Ivory Coast; 2 coll. (8 females); AMP Lemniscomys striatus longer than distal seta; setae pd 1 and ad Congo (Brazzaville) ; 6 females (type 1 of femur I sube(}ual in length; anterior specimens); Taufflieb. 1954 seta of coxae II III seta of and and coxa Ghana; 1 coll. (3 females); AMP IV setaceous; posterior seta of coxae II Ivorv Coast; 8 coll. (29 females, and III robust, blunt, and peglike; no ro- 1 male); AMP Nigeria; 11 coll. (41 females. bust, blunt preapical setae on tarsi II, 1 ny.); AMP III, or IV; however, some preapical setae Togo; 8 coll. (19 females); AMP may be spinelike; most other leg setae Lophuromys sikapusi setaceous and normally developed. Nigeria; I coll. (7 females); AMP Mas musculnidcs

Male: Unknown. Congo (Brazzaville); 1 coll. (1 female): Taufflieb, 1954 Myomys daltoni

COLLF.CTION RECORDS Ghana; 1 coll. (1 female); AMP Tadarida major Praomys titUbergi Nigeria; 1 coll. (1 female); AMP Upper Volta; 1 coll. (2 females); AMP I rnnomys oweni Talera Kempi Senegal; 2 coll. (2 females); AMP Dahomey; l-j- coll. (1-^- female); I hanomys riiddi Zumpt coll. (AMP) Ivorv Coast; 3 coll. (13 females. Dasymys incomptus 1 ny.); AMP Congo (BrazzaviihO ; Taufflieb, 1962 Unknown Hybomys trivirgatus Dahomey; 1 coll. (2 females); AMP Ivorv Coast; 4 coll. (26 females. Ivorv Coast; 2 coll. (6 females.

4 ny.); AMP 1 ny.); AMP Tune 1976 HERRIN, TIPTON: AFRICAN LAELAPS 139

Remarks.— There is no one character setae. Anal i)late triangular in general which may be used to distinguish L. shape, longer than wide, and with anterior prenieri from all other taxa of this sub- margin slightly invaginated; adanal setae group as in the case of L. keeganL L. of moderate length, extending to base of transvaalcnsis. L. congoicola, and L. lavoi- postanal seta; adanal setae set at level of pierrci. This taxon may be sej)arated from posterior end of anal orifice. Unarmed L. simiUimns and L. parasirriillirnus by venter bearing approximately 14 pairs the greatest width of the genital phite at of setaceous setae, 1- pairs adjacent to the level of the 2nd genital setae rather genital and anal plates plus ap})roximately than at the level of the 3rd, the meta- 10 j)airs near or (m posterior body mar- podal plates more irregularly oval rather gins; metapodal plates oval, slightly longer than elongate, and the greater length than wide. Peritreme extending to level width ratio of the sternal plate (greater of posterior of coxa I. Dorsal plate bearing than .75). 39 pairs of setaceous setae; most dorsal L. grcnicri has been recorded from a setae of medium length, length usually variety of different small mammal hosts equal to distance between adjacent setae; in northwestern Africa south of the subterminal setae (J5) reaching at least Sahara; however, it is parasitic primarily to level of terminal setae and possibly on several species of Lemniscomys. to posterior margin of dorsal plate. Twelve to 14 pairs of setae border dorsal opistho- Both proximal Subgroup B soma on soft integument. and distal setae of coxa I setaceous, proxi- This subgroup is composed of only four mal seta somewhat longer than distal T.. L. taxa: lavieri. L. fritzumpti. thamno- seta; setae pd 1 and ad 1 of femur I mys. and L. moucheti. The first two and subequal in length; anterior seta of coxae the fourth taxa bear at least one blmit, 11 and III and seta of coxa IV setaceous; peglike setae preapically on tarsi II and posterior seta of coxae II and III robust, III, whereas tarsi II and III of L. thamno- blunt, and peglike; tarsi II and III each mys bear all setaceous setae. This latter with one robust, blunt preapical setae species is placed with subgroup B be- plus one spinelike seta; all other leg seta cause of its overall phenetic similarity setaceous and normally developed. to the other three species. Male: (Figs. 34-35) Gnathosomal and hypostomal setae setaceous, with medial Laelaps (Laclaps) lavieri Taufflieb hypostomal setae somewhat longer than Figs. 30-35 gnathosomal setae but not reaching to Laclaps lavieri Taufflieb, 1954, Arm. Parasit. base of gnathosomal setae. Ventral setae, :442 (Holotype: Brazzaville, Congo; 29(4) except adanal and postanal setae, rela- Institut Pasteui-. Paris); Tipton. 1960. Univ. tiveh' long, extending well beyond base Calif. Publ. Ent. 16(6) :273-274. T.aelaps nigeriensis Keegan, 1962. J. Parasit. of seta immediately posterior; holoventral 48 (4): 621 -622 (Holotype: Adu, Nigeria; plate broad between coxae II and III, United States National Museum, Washington. narrowing considerably between coxae IV D.C.). and greatly expanded posterior to coxae Description. — Female: (Figs. 30-33) IV; expanded area between genital setae

Dorsal plate length 600 /x, width 419 /<,. and anal orifice bearing 5 pairs of seta- Gnathosomal and hypostomal setae setaceous setae; adanal setae relatively short, ceous; medial hypostomal setae relatively extending no further than base of postanal long, reaching almost to base of gnatho- seta; postanal seta much longer than somal setae. Posterior margin of sternal adanal setae. Metapodal plates inapparent, plate very slightly invaginated medially. api)arently fused to lateral margin of holo- Anterior flap of genital plate not overlap- ventral plate. Unarmed venter bearing 9 ping posterior margin of sternal plate; dis- or 10 pairs of setae immediately adja- tance between 1st genital setae subequal to cent to holoventral plate, plus 8 to 10 distance between 4th genital setae, and pairs on posterior lateral body margin. distance between 2nd genital setae sub- Peritreme extends to level of posterior or equal to distance between 3rd genital middle of coxa I. Dorsal plate bearing 39 setae; greatest width of genital plate near pairs of setaceous setae; posterior dorsal or slightly anterior to level of 3rd genital setae relatively long, length considerably 140 GREAT BASIN NATURALIST Vol. 36, No. 2

Figs. 30-33. \Laelaps lavieri Taufflieb, female. (30) venter; (31) dorsum, scale 100/i; (32) ventral view of tarsus II; (33) ventral view of tarsus III, scale = SO/t.

greater than distance between adjacent tively short and spinelike; tarsi II and setae; subterminal setae (J5) extending III each with two or three rather robust, well beyond posterior margin of dorsal spinelike to peglike preapical setae; other plate and about half the length of the leg setae mostly setaceous and normally terminal setae; both proximal and distal developed. setae of coxa I setaceous, with proximal COLLKCTION RECORDS .seta considerably longer and robust more Crocidura sp. basally than distal seta; setae [kI 1 and Nigeria (Adu); 2 femali's; Koegan, 1962

ad 1 of femur I subequal in lenglh; an- Srli'isnrt'.r grmmcus terior seta of coxae II and III, posterior (iliaiia: I coll. (1 fiMiialc. 1 ma!.-); AMP seta f)f coxa II, ajid seta of coxa IV all 1 1 ) psifimilhdi' nionslrosus setaceous; [jostcrior seta of (oxa III rela 'Ivory Coast; 1 coll. (1 female): AMP Tune 1976 IIERRIN, TIPTON: AFRICAN L.^EL.^PS 141

Figs. 34-35. Laelaps lavieri Taufflieb. male. (34) venter; (35) dorsum, scale 100/i.

Nycteris urge Mus haussa Upper Volta; 1 coll. (1 female, Nigeria; 2 coll. (5 females.

1 male); AMP 1 male); AMP Hipposideros coffer Alus minutoides coll. females. Ivory Coast; 2 (3 Ghana; 1 coll. (1 female); AMP 1 male, 2 ny.); AMP Ivory Coast; 1 coll. (1 female); AMP Hipposideros crclops Rhodesia; 3 coll. (18 females,

Ivory Coast;' 1 coll. (1 female); AMP 1 male); AMP Tatera leucogaster South Africa (ORS); 7 coll. (9 males); Soutli Africa; 2 coll. (2 females); AMP AMP South Africa (ORS); 1 coll. musculoides (1 female); AMP Mus Aethomys chrysophilus Congo (Brazzaville); 4 females (.type specimens); Taufflieb. 1954 Rhodesia; 1 coll. (1 female); AMP South Africa; 1 coll. (2 females); AMP Ghana; 13 coll. (46 females. South Africa (ORS); 2 coll. 16 males, 5 ny.); AMP

(2 females) ; AMP Ivory Coast; 23 coll. (62 females. Lemniscomys macculeus 3 males. 53 ny.); AMP

Ivory Coast; 1 coll. (3 females); AMP Nigeria (Adu); 1 coll. (1 female); Lemniscomys striatus Keegan. 1962 Togo; 2 coll. (3 females); AMP Nigeria; Zumpt collection (AMP) Lophuromrs sikapusi Upper Volta; 3 coll. (4 females); AMP Ghana; 1 coll. (10 females); AMP Mus setulosus Mastomrs natalensis Ghana; 18 coll. (80 females. Ghana; 1 coll. (1 female); AMP 6 males. 2 ny.); AMP South Africa; 1 coll. (1 female. Ivorv Coast; 23 coll. (127 females, 1 male); AMP \\ males. 10 ny.); AMP South Africa (ORS); 4 coll. ( 6 females); AMP Myomys daltoni collection (AMP) Mus sp. Ghana; Zumpt Angola (Dundo); 2 females; Taufflieb. 1962 Praomys lullbergi Cameroons (Yaounde); Taufflieb. 1962 Ghana; 5 coll. (12 females.

Congo (Leopoldville) ; 3 females. 7 males. 2 nv.); AMP

1 male; Taufflieb. 1962 Ivory Coast; 1 coll. (1 female); AMP ;

142 GREAT BASIN NATURALIST Vol. 36, No. 2

Saccostomus campestris to posterior margin of sternal plate. An- South Africa (ORS); 1 coll. (1 female); terior flap of genital plate overlapping AMP posterior margin of sternal plate only Ictonyx striatus slightly if at all; distance between 1st South Africa (ORS); 1 coll. ( 1 female); AMP genital setae subequal to distance between Genetta villiersi 4th genital setae, distance between 1st 1 coll. females); AMP Ivory Coast; (2 genital setae may be slightly less; distance Unknown between 2nd genital setae slightly less Botswana; 3 coll. (3 females, 2 males) AMP than distance between 3rd genital setae; Ghana; 1 coll. (10 females, greatest width of genital plate at level of 5 males, 8 ny.); AMP 3rd pair of genital setae. Anal plate Togo; 1 coll. (2 females) ; AMP roundly triangular, width equal to length, Remarks.— L. lavieri resembles L. fritz- anterior margins straight; adanal setae umpfi in overall morphological characters rather long, extending distinctly beyond but ma}^ be separated by the longer peri- base of postanal seta; adanal setae set at treme (extends anteriorly to middle or level of posterior end of anal orifice. Un- posterior of coxa I) and the presence of armed venter bearing aproximately 10 only one blunt, peglike preapical seta on pairs of setaceous setae, 4 pairs adjacent tarsi II and III rather than two or more to genital and anal plates plus approxi- on each tarsi II-IV. This latter character mately 6 pairs near or on posterior lateral may also be used to separate L. lavieri body margins; metapodal plates elongate- from L. thamnomys, as well as the overall oval. Petritreme extending anteriorly to bod}^ size; i.e., L. thamnomys is consider- level of middle of coxa II. Dorsal plate ably larger (length of dorsal plate great- bearing 39 pairs of setaceous setae; most er than 575 jx) . Both L. lavieri and L. dorsal setae relatively long, length equal fritzumpti may be separated from L. to or slightly greater than distance be- moucheti by the longer dorsal setae as tween adjacent setae; subterminal setae opposed to quite short dorsal setae in the (J5) reaching almost to posterior margin latter, and by the presence of dorsal setae of dorsal plate. Fifteen to 20 pairs of setae px 3 which is absent in L. moucheti. Also bordering dorsal opisthosoma on soft the medial h3qpostomal setae are distinc- integument. Both proximal and distal tiA'ely longer than in L. moucheti. setae of coxa I setaceous, with distal seta L. lavieri has been collected from many slightly shorter than proximal seta; setae small mammal hosts throughout Africa, pd 1 and ad 1 of femur I subequal in southern Africa as well as northwest length; anterior seta of coxae II and III Africa; however, it is parasitic primarily and seta of coxa IV setaceous; posterior on species of Mus. More collections are seta of coxae II and III robust, blunt, and recorded from this group of hosts than peglike, posterior seta of coxa II somewhat from all other small mammal hosts com- more elongate than that of coxa III; tarsi bined. II and III each with three rather robust, blunt jjreapical setae; tarsus IV with one

Laelaps (Laelaps) fritzumpti Taufflieb or two longer blunt, ])reapical setae; all ij ' Figs. 36-42 other leg setae setaceous and normally developed. Laelaps fritzumpti Taufflieb, 1964, Z. f. Para- siten. 24:305-308 (Holotype: Nosob River, Male: (Figs. 41-42) Gnathosomal and Kalahari, South Africa; South African Insti- hypostomal setae setaceous, with medial tute for Medical Research. .Johannesburg). hypostomal setae of moderate length but Description.^— Female: (Figs. 36-40) not reaching to base of gnathosomal setae.

Dorsal plate length 634 /x, width 417 /-. Ventral setae, except adanal and post- Gnathosomal and hypostomal setae seta- anal setae, rather long, each extending ceous; medial hypostomal .setae of mod- in l(>ngth well beyond base of seta im- erate length, not reaching to base of metHately ])osterior; holo\eiitral plate Ji gnathosomal setae. Posterior margin of rather narrow between (oxae IV, and ' sternal plate slightly invaginated, invagi- greatly exjianded innnediately jiosterior nation not reaching to level of 3rd sternal to coxae IV; expanded area between geni- , setae; setae st. 1 rather long, reaching to or lal setae and anal orifice bearing 5 pairs almost to level of 3rd sternal setae but not of setaceous setae; adanal seta(^ relatively Iiiiio 1976 HERRIN, TIPTON: AFRICAN LAELAPS 143

Figs. 36-40. Laelaps fritzumpti Taufflieb, female. (36) venter; (37) dorsum, scale = 100/t; (38) ventral viev^' of tarsus II; (39) ventral view of tarsus III; (40) ventral view of tarsus IV, scale = SOfi. 144 GREAT BASIN NATURALIST Vol. 36, No. 2

iFigs. 41-42. Laelaps fritzumpti Taufflieb, male. (41) venter; (42) dorsum, scale = lOOju. short, reaching no further than base of three spinelike preapical setae, one ])air postanal seta; postanal seta considerably on each tarsi may be blunt; all other leg longer than adanal setae and enlarged setae mostly setaceous and normally de- somewhat basally. Metapodal plates fused \ eloped. to lateral margins of holoventral plate Collection records posterior to coxae IV. Peritreme extends FArphantuliis m varus to level of middle of coxa II. Dorsal plate Soutli Africa (ORS); 2 coll. (2 females); bearing 39 pairs of setaceous setae; most AMP FJephanlulus rapes/ r is dorsal setae rather long, length consider- South Africa (ORS); 10 coll. (11 f.Muales); ai)ly greater than distance between adja- AMP cent setae; subtermirial setae (.15) some- Macroscelides proboscideus

South Africa. (ORS); 1 coll. (3 females); what longer than normal, extending dis- AMP tinctly beyond posterior margin of dor- Desmodillas naricalaris sal plate. Unarmed venter bearing 6 to 8 South Africa (ORS); 8 coll. (12 females); pairs of setae adjacent to holoventral plate. AMP Soft integument of opisthosoma bearing Gcrbillus pacha South Africa (ORS); ^1 coll. (Vl- fenuilos. 8 to 10 pairs of setae. Both proximal and 1 male); AMP distal setae of coxa I setaceous, proximal Tatcra brandsi seta considerably longer than distal seta; South Africa (ORS); 2 coll. (2 females): setae ad 1 and pd 1 of fenuir I sube(|ual AMP Talcra Icacopasler in length, ad 1 .seta slightly shorter; an- !• South Africa (ORS); coll. i !(> females); terior seta of coxae I and II, posterior .seta AMP of coxa II, and seta of coxa IV all seta- Pr/rnnnsias coll in us

South Africa; I coll. (1 fenuUo) ; AMP ceous; posterior seta of coxa III shorter and South Africa (OHS); 7 coll. (7 females); .spinelike; tarsi II and III cadi with two or AMP June 1976 HERRIN, TIPTON: AFRICAN Lu-^ELAPS 145

Aetliomys sp. Congo; Institut Pasteur, Paris); Tipton, 1960, South Africa (Cape Province); 13 Univ. Calif. Publ. Ent. 16(6): 283. females; Taufflieb. 1964 Aethomrs chrysophilus Description. — Female: (Figs. 43-44)

Rhodesia; 1 coll. (1 female); AMP Dorsal plate length 599 /a, width 364 /j,. South Africa (ORS); 2 coll. (63 Gnathosomal and hypostomal setae seta- females. 1 male); AMP ceous; medial hypostomal setae of mod- Aethomrs namaqitensis erate length but not to Botswana; 1 coll. (12 females): AMP reaching base of Botswana (northern); Taufflieb. 1964 gnathosomal setae. Posterior margin of South Africa (ORS); 79 coll. (534 sternal plate only very slightly invagi- females. 2 males) ; AMP nated; setae st. 1 rather long, reaching South Africa (Transvaal); 2 females; Taufflieb. 1964 almost to level of 3rd sternal setae. An- Mastomrs natalensis terior flap of genital plate slightly over- South Africa (ORS); 3 coll. lapping posterior margin of sternal plate; females); AMP (3 distance between 1st genital setae only Thallomys sp. South Africa (Cape Province); 2 slightly less than distance between 4th females; Taufflieb. 1964 genital setae, distance between 2nd genital Thallomys paedulcus setae slightly less than distance between South Africa (Cape Province); 17 ^rd genital setae; greatest width of genital females (type specimens); Taufflieb. 1964 South Africa (ORS); 12 coll. plate at level of 3rd pair of genital setae. (52 females); AMP Anal plate roundly triangular, as wide as Rhabdomys pumilio long, with anterior margins rounded; South Africa (ORS); 10 coll. adanal setae of moderate length, extend- (12 females); AMP Saccostomus campestris ing slightly beyond base of postanal seta; South Africa (ORS); 2 coll. adanal setae set near posterior end of anal (8 females); AMP orifice. Unarmed venter bearing approxi- Otomys sp. mately 10 pairs of setaceous setae, 4 pairs South Africa (Cape Province); 1 female; Taufflieb, 1964 immediately adjacent to genital and anal Otomys irroratus plates plus approximately 6 pairs near or South Africa (Orange); 1 female; on posterior lateral body margins; meta- Taufflieb. 1964 podal plates oval, slightly longer than Paratomys brandsi w^ide. Peritreme extends to level of middle South Africa CORS); 1 coll. (1 female); AMP of coxa I. Dorsal plate bearing 39 pairs of Unknown setaceous setae; most dorsal setae of me- South Africa; 1 coll. (1 female); AMP dium length, length only slightly greater South Africa (ORS); 11 coll. than distance between adjacent setae, if as (14 females); AMP long; subterminal setae (J5) reaching no Remarks.— L. fritzumpti may be distin- further than posterior margin of dorsal guished from all other taxa of the sub- plate. Twelve to 14 pairs of setae border group by the three blunt, peglike preapical dorsal opisthosoma on soft integument. setae on tarsi II and III, and by the shorter Both proximal and distal setae of coxa I peritreme (extends only to middle of setaceous, proximal seta considerably long- coxa II) . Several other diagnostic charac- er than distal seta; setae pd 1 and ad 1 of ters are the relatively large genital and femur I subequal in length; anterior seta anal plates, adanal setae almost as long of coxae II and III and seta of coxa IV as postanal seta, and rather long dorsal setaceous; posterior seta of coxae II and setae. III robust, blunt, and peglike; no blunt, This species is very abundant on many robust preapical setae on tarsi II, III, or small mammal hosts throughout southern IV; however, some preapical setae may be Africa. It has been collected most fre- spinelike; most other leg setae setaceous quently from Aethomys species, Thallo- and normally developed. mys species. Rhabdomys pumilio, Ger- Male: (Figs. 45-46) Gnathosomal and bullus paeba, and Elcphantulus rupestris. hypostomal setae setaceous; medial hypostomal setae of moderate length, reaching Laelaps (Laelaps) thamnomys Taufflieb almost to base of gnathosomal setae. Ven- Figs. 43-46 tral setae, except adanal and postanal

Laelaps thamnomys Taufflieb, 1954. Ann. setae, rather long and slender, extending

Parasit. 29(4) : 444-44-6 (Holotype: Brazzaville. well beyond base of adjacent posterior 146 GREAT BASIN NATURALIST Vol. 36, No. 2

T /

Figs. 43-44. Laelaps thamnornys Taufflieb. female. (43) venter; (44) dorsum, scale = 200/i.

Figs. 45-46. Laelaps ihaninoniys Taiifflii-l), male. (45) venter; (46) dorsuin. scale 100/t. June 1976 HERRIN, TIPTON: AFRICAN L.AELAPS 147

seta; holoventral plate rather broad be- plate at level of 3rd genital setae rather tween coxae II and III, greatly narrowing than at level of 2nd, and distance be- between coxae IV, and greatly expanded tween 1st genital setae equal to or less posterior to coxae IV; expanded area be- than distance between 4th, rather than the tween genital setae and anal orifice bear- reverse. ing 5 pairs of setaceous setae; adanal This taxa is recorded primarily from setae of medium length, extending to or I'/iarnnornys rutilans in northwest Africa slightly beyond base of postanal seta; south of the Sahara. adanal setae set at level slightly posterior to middle of anal orifice; postanal seta Laclaps (iMclaps) moucheti Taufflieb approximately twice as long as adanal Figs. 47-53 setae, aTid somewhat more robust. Meta- T.aelaps moucheti Taufflieb, 1959, J. Ent. See. podal jilates inapparent, apparently fused So. Afr. 22(2): 398-399 (Holotype: Yaounde, to lateral extension of holoventral plate; Cameroun; Museum d'Histoire Naturelle, unarmed venter bearing approximately Paris). 12 to 14 pairs of setaceous setae adjacent Description.— Female: (Figs. 47-51) to holoventral plate. Peritreme extending Dorsal plate length 484 ^, width 302 /x. to middle of coxa I. Dorsal plate bearing (rnathosomal and hypostomal setae seta- 39 pairs of setaceous setae, length and ])o- ceous. Medial hypostomal setae relatively sition of setae approximately as in female. short, reaching no more than half dis- Both proximal and distal setae of coxa I tance to gnathosomal setae. Posterior mar- setaceous, proximal seta somewhat more gin of sternal plate somewhat invaginated, robust and longer than distal seta; setae invagination reaching no further than ad 1 and pd 1 of femur I subequal in level of 5rd sternal setae; setae st. 1 of length, pd 1 seta slightly longer; anterior medium length, reaching to level halfway seta of coxae II and III and seta of coxa between 2nd and 3rd sternal setae. An- IV of moderate length and setaceous; posterior flap of genital plate overlapping terior seta of coxa II of moderate length posterior margin of sternal plate only and rather robust, with posterior seta of slightly; distance between 1st genital coxa III short, robust, and spinelike; most setae slightly greater than distance be- leg setae setaceous and normally devel- tween 4th genital setae, distance between oped; however, some may be shorter and 2nd genital setae slightly greater than rather spinelike. distance between 3rd genital setae; Collection records greatest width of genital plate at level of Mus musculoides 2nd pair of genital setae. Anal plate Togo; 1 roll. (1 female); AMP roundly triangular, almost as wide as Praomrs tullbergi long, with anterior margins rounded; Togo; 2 roll. (2 females); AMP of Thamnoniys rutilans adanal setae moderate length, extending Congo (Brazzaville): 6 females (type to base of postanal seta; adanal setae set specimens); Taufflieb. 1954 at level of posterior end of anal orifice.

Ivory 1 coll. Coast; ( 6 females); AMP Unarmed venter bearing approximately Togo; 8 coll. (66 females. 1 male. 14 to 16 pairs of setaceous setae, 4 pairs 1 ny.); AMP immediately adjacent to genital and anal Remarks.— As noted previously, L. plates plus approximately 10 to 12 pairs thamnomys differs from other taxa of near or on posterior lateral body margin; subgroup B in the form of preapical metapodal plates oval to elongate-oval. setae on tarsi II and III, i.e., setaceous Peritreme extends to level of middle of rather than blunt, peglike. Also, it is a coxa I. Dorsal plate bearing 38 pairs of rather large species with the dorsal plate rather small setaceous setae, setae px3

exceeding 575 //. in length. In these two absent; most dorsal setae relatively characters L. tliarnnomys is quite similar short, length no greater than half distance to L. kampalcnsis, which is placed in ma- between adjacent setae; subterminal setae jor group II, subgroup A. because of over- (J5) smaller than all others, and terminal all phenetic similarity. The former differs setae (Z5) considerably longer than other from the latter in the following characters: adjacent setae. Ten to 12 pairs of setae only slightly invaginated posterior margin border dorsal opisthosoma on soft integu- of sternal plate, greatest width of genital ment. Both proximal and distal setae of

I 148 GREAT BASIN NATURALIST Vol. 36, No. 2

dorsum, scale = lOO/x; (49) Figs. 47-51. Laelaps moucheti Taufflicb, female. (47) venter; (48) = ventral view of tarsus II; (50) ventral view of tarsus III; (51) ventral view of tarsus IV, scale 50/t. June 1976 HERRIN, TIPTON: AFRICAN LAEL.'^PS 149

Figs. 52-53. Laelaps moucheti Taufflieb, male. (52) venter: (53) dorsum, scale 100/i.

coxa I setaceous, with proximal seta con- relatively short, length not extending to siderably longer than distal seta; setae base of postanal seta; adanal setae set ad 1 and pd 1 of femur I subequal in near level of middle of anal orifice; post- length; anterior seta of coxae II and III anal seta considerably longer than adanal and seta of coxa IV setaceous; posterior setae. Metai)odal plates inapparent, ap- seta of coxae II and III rather robust, parently fused with lateral extension of blunt, and peglike; tarsi II and III each holoventral plate; unarmed venter bear- with one robust, blunt preapical setae; ing 16 to 18 rather small, slender setae.

1 all other leg setae mostly setaceous and Peritreme extending to middle of coxa I. normally developed. Dorsal plate bearing usual 39 pairs of setaceous setae; length and position of Male: (Figs. 52-53) Gnathosomal and setae as in female. Soft integument of hypostomal setae setaceous, medial hypost- opisthosoma bearing 10 to 12 pairs of omal setae relatively short, not reaching setaceous setae. Both proximal and distal more than half distance to gnathosomal setae of coxa I setaceous, with proximal setae. Ventral setae, except adanal and seta considerably longer than distal seta; postanal setae, of moderate length, each setae pd 1 and ad 1 of femur I subequal extending in length somewhat beyond in length; anterior seta of coxae II and base of seta immediately posterior; holo- III and seta of coxa IV slender and seta- ventral plate rather broad between coxae ceous; posterior seta of coxae II and III II and III, rather narrow between coxae spinelike; no blunt preapical setae on IV, and greatly expanded immediately tarsi II, III or IV; however, some pre- posterior to coxae IV; expanded area be- apical setae may be spinelike; most other tween genital setae and anal orifice bear- leg setae setaceous and normally devel- ing 5 pairs of setaceous setae; adanal setae oped. 150 GREAT BASIN NATURALIST Vol. 36, No. 2

Collection records ceous proximal seta on coxa I. L. kam- "rodents" palensis differs in having a setaceous dis- Cameroon (Yaounde); 10 females (type spec- tal seta on coxa I, and L. aethiopicus bears imens); Taufflieb, 1959 blunt, peglike setae both proximally and Remarks.— L. moucheti possesses distally on coxa I. Both of these taxa are several diagnostic characters which placed in this major group because of their separate it from other taxa of subgroup overall phenetic similarity to taxa within B (major group I): unusually short dor- this group. sal setae, particularly posterocentrally; Subgroup dorsal setae px3 absent, thus dorsal plate A bears only 38 pairs of setae; medial hy- The four species of this subgroup (L. postomal setae short, extending no further kampalensis. L. tillae. L. peregrinus, and than half distance to gnathosomal setae. L. rouhaudi) are characterized by tarsi This species is reported only from the II and III bearing setaceous preapical "type" collection which is from "rodents" setae; no blunt, peglike setae are present in Cameroon (Yaounde) . It has not been on the tarsi of any of them. This is in con- found in any collections of the African trast to all other taxa of major group II Mammal Project to date. which bear at least one blunt, peglike preapical seta on tarsi II and III. As noted above, L. kampalensis differs from Major Group II the other three species of this subgroup

This major group contains 14 taxa, as in having a setaecous distal seta on coxa I. opposed to major group I which consists of 11 and major group III which has 6. Laelaps (Laelaps) kampalensis Taufflieb With but two exceptions (L. kampalensis Figs. 54-55 and L. aethiopicus) , taxa of this major Laelaps kampalensis Taufflieb. 1959. J. Ent. Soc. are characterized the presence group by So. Afr. 22(2): 402-403. (Holotype: Kampala. of a blunt, peglike distal seta and a seta- Uganda; Museum d'Histoire Naturelle. Paris).

Figs. 54-55. Laelaps kampalensis Taufflieb, feniah'. (54) venter; (55) dorsum, scale 100/i. June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 151

Description.— Female-. (Figs. 54-55) to L. tillae; however, it may be distin- Dorsal plate length 659 ^i, width 445 /i. guished from the latter by the significant- Gnathosomal and hypostomal setae seta- ly larger size (dorsal jjlate greater than ceous; niecHal hypostomal setae of mod- 575 /I long), and by the setaceous distal erate length but not reaching base of seta of coxa I rather than a small, blunt, gnathosomal setae. Posterior margin of peglike seta. In the above noted characters sternal plate invaginated to or slightly as well as certain others L. kampalensis is beyond level of setae st. 3; setae st. 1 of similar to L. tkamnomys; yet it differs in moderate length, reaching to point half- the following notable characters: posterior way between setae st. 2 and st. 3. Anterior margin of sternal plate distinctly more flap of genital plate not overlapping pos- in^aginated; greatest width of genital plate terior of sternal plate; distance between at level of 2nd genital setae rather than 1st genital setae distinctly greater than at 3rd; and distance between 1st genital distance between 4th genital setae; great- setae distinctly greater than distance be- est width of genital jilate between 2nd and tween 4th. 3rd genital setae; distance between 2nd L. kampalensis is recorded from the genital setae subequal to that between 3rd type collection which is Lemniscomys stri- genital setae; posterior margin of genital atus in Uganda, except for a single col- plate betw'een 4th pair of genital setae lection from an unknown host in Togo. straight to slightly invaginated. Anal plate Thus, at present very little is known of length subequal to width, anterior margin the actual host and geographic distribution convex; adanal setae relatively long, ex- of this species. tending well beyond base of i)ostanal seta; adanal setae set near posterior end of Laelaps (Laelaps) tillae Taufflieb anal orifice; postanal seta somewhat longer Figs. 56-59 than adanal setae. Unarmed venter bearing about 14 pairs of setaceous setae, 8 Laelaps tillae Taufflieb, 1959. J. Ent. So. Afr. 22(2):403-404 (Holotype: Kruger National pairs distinctly ventral adjacent to genital Park, Transvaal, South Africa; Museum d'His- and anal plates, with other pairs more toire Naturelle, Paris). marginal; metapodal plates elongate-oval. Description.— (Figs. Peritreme extends to middle of coxa I. Female: 56-57) Dorsal plate bearing 39 pairs of setaceous Dorsal plate length 644 /x, width 416 /t. setae; most dorsal setae of moderate length, Gnathosomal and hypostomal setae seta- length equal to distance between adjacent ceous; medial hypostomal setae of medium setae; subterminal setae (J5) reaching length, reaching slightly further than half to posterior margin of dorsal plate. Four- distance to gnathosomal setae. Posterior margin of sternal plate slightly in- teen to 1 7 pairs of setae border dorsal plate on soft integument. Both proximal vaginated; sternal setae st. 1 relatively and distal setae of coxa I setaceous, with long, reaching to or slightly beyond level proximal seta slightly longer than distal of 3rd sternal setae, but not to posterior plate. seta; setae pd 1 and ad 1 of femur I margin of sternal Anterior flap of genital subequal in length, with ad 1 seta only plate overlapping posterior margin slightly longer; anterior seta of coxae II of sternal plate only slightly; distance be- and III and seta of coxa IV setaceous; pos- tween 1st genital setae distinctly greater terior seta of coxae II and III robust and than distance between 4th genital setae, peglike; all leg setae setaceous; however, and distance between 2nd genital setae one or two preapical setae of tarsi II and distinctly greater than distance between III may be somewhat more robust than 3rd genital setae; greatest width of genital other setae. plate at level of 2n(l pair of genital setae. Male: Unknown. Anal plate triangular in general shape, slightly longer than wide, with anterior Collection records margin straight; adanal setae of moderate Lertiniscomys striatus length, extending to or slightly beyond Uganda; 8-f females (type specimens); Taufflieb. 1959 base of postanal seta; adaiial setae set at Unknown level of posterior end of anal orifice. Un- Togo; 1 coll. (1 female); AMP armed venter bearing approximately 10 Remarks.— In overall morphological pairs of setaceous setae, 4 pairs adja- characters L. kampalensis is most similar cent to genital and anal plates plus ap- 152 GREAT BASIN NATURALIST Vol. 36, No. 2

Figs. 56-57. Laelaps tillae Taufflieb, female. (56) venter; (57) dorsum, scale = 200^i.

Figs. 58-59. Laelaps tillae Taufflieb, innle. (58) venter; (59) dorsum scale 100/t. June 1976 HERRIN. TIPTON: AFRICAN LAELAPS 153 proximately 6 pairs near or on posterior tal setae of coxa I setaceous, with proxi- lateral body margins. Metapodal plates mal seta somewhat longer than distal seta; rather elongate. Peritreme extends to lev- setae pd 1 and ad 1 of femur I subequal el of middle of coxa I. Dorsal plate bear- in length, with seta pd slightly longer; ing 39 pairs of setaceous setae; most dorsal anterior seta of coxae II and III, posterior setae of medium length, length equal to sota of coxae II, and seta of coxa IV or slightly greater than distance between setaceous, coxa IV seta somewhat shorter; adjacent setae; setae J4 somewhat shorter posterior seta of coxa III rather short, than adjacent anterior setae, and sub- robust, and spinelike; no robust, blunt terminal setae (15) reaching no further peglike preapical setae on tarsi II, III, than posterior margin of dorsal plate; or IV; however, some preapical setae terminal setae (Z5) approximately three and other leg setae may be shorter and times as long as subtenninal setae. Twelve spinelike. to 15 pairs of setae border dorsal opistho- Collection records soma on soft integument. Proximal seta Aethomys chrysophilus of coxa I setaceous, distal seta rather small, South Africa; 2 coll. (3 females); AMP blunt, and peglike; setae ad 1 and pd 1 Lemniscomys sp. of femur I subequal in length; anterior South Africa (Transvaal): 7 females; seta of coxae II and III and seta of coxa Taufflieb. 1959 Lemniscomys griselda IV setaceous, coxa IV seta somewhat Rhodesia; 1 coll. (1 female); AMP smaller; posterior seta of coxae II and III South Africa; 1 coll. (1 female, rather robust, blunt, and peglike; no ro- 1 ny.); AMP bust, blunt preapical setae on tarsi II, South Africa (Transvaal); 29 females; Taufflieb, 1959 and 1964 III, and IV; however, some preapical setae Mastomys natalensis spinelike; setae may be most other leg South Africa; 4 coll. (4 females, setaceous and normally developed. 4 males) ; AMP

xSouth Africa (Transvaal) ; 22 females Male: (Figs. 58-59) Gnathosomal and (type specimens); Taufflieb, 1959 hypostomal setae setaceous ; medial hy- Rhabdomys pumilio postomal setae long and slender, extend- South Africa; 1 coll. (1 female); AMP Saccosfomus campestris ing to base of gnathosomal setae. Ventral Rhodesia; 1 coll. (1 female); AMP setae, except adanals. rather long, each Unknown extending in length well beyond base of South Africa (ORS); 1 coll. (1 female); adjacent posterior setae; holoventral plate AMP rather broad betw^een coxae II and III. Remarks.^— As noted previously, L. quite narrow^ between coxae IV, and mod- tillae is phenetically quite similar to L. erately expanded posterior to coxae IV, hampalcnsis in overall morphological char- but not greatly exjianded as in some other acters but differs primarily in the smaller species; expanded area between genital size (dorsal plate less than 575 /x long) setae and anal orifice bearing 5 pairs of and in the presence of a small, blunt, setaceous setae; adanal setae relatively |)oglike seta distally on coxa I. L. tillae short, extending no more than half dis- differs from the other two taxa of sub- tance between postanal seta; adanal setae group A by the smaller, more slender set at level near middle of anal orifice; peglike distal seta of coxa I, the more postanal seta considerably longer than slender, setaceous proximal seta of coxa adanals, and rather robust and spinelike. 1. and the slight invagination of the pos- Metapodal plates elongate-oval, at least terior margin of the sternal plate. twice as long as wide; unarmed venter L. tillae has been collected only from bearing approximately 10 pairs of setae. southern Africa on a half dozen different 2 immediately adjacent to holoventral hosts; however, it is reported most fre- plate and anal region plus about 8 pairs quently from Mastomys natalensis and on posterior lateral body margins. Peri- Lemniscomys species. treme extending to middle or anterior of coxa I. Dorsal plate bearing 39 pairs Laelaps (Laelaps) peregrinus Taufflieb of setaceous setae; length position and ap- Figs. 60-61 proximately as in female. Soft integument Laelaps peregrinus Taufflieb. 1059. .1. Ent. So. of opisthosoma bearing about 8 to pairs 10 Afr. 22(2): 401 -402 (Holotype: Pilgrims Rest, of setaceous setae. Both proximal and dis- Transvaal. South Africa; Museum d'Histoire )

154 GREAT BASIN NATURALIST Vol. 36, No. 2

Figs. 60-61. Laelaps peregrinus Taufflieb, female. (60) venter; (61) dorsum, scale = 200/i.

Naturelle de Paris); Taufflieb, 1964, Z. f. to 10 })airs distinctly ventral with others Parasiten 24:307. more marginal; metapodal plates elongate- Description. — Female-. (Figs. 60-61 oval. Peritreme extending to middle or posterior of coxa I. Dorsal plate bearing Dorsal plate length 653 /a, width 438 //.. Gnathosomal and hypostonial setae seta- 39 pairs of setaceous setae; most setae of ceous; medial hypostomal setae long, but moderate length, length equal to distance not reaching to base of gnathosomal setae. between adjacent setae; subtemiinal setae Posterior margin of sternal plate invagi- (J5) reaching slightly beyond posterior nated slightly beyond level of seta st. 3; margin of dorsal plate; approximately 20 setae st. 1 rather long, reaching almost to setae bordering dorsal o])isthosomal on posterior margin of sternal plate. Anterior soft integument. Proximal seta of coxa I flap of genital plate rather narrow and setaceous but somewhat robust, distal seta not overlapping })osterior of sternal plate; usually blunt, peglike (may be robust distance between 1st genital setae much and spinelike), and about half the length greater than distance between 4th genital of proximal seta; setae jkI 1 and ad 1 of setae; distance between 2nd genital setae fenuir I subequal in length, with ad 1 only distinctly greater than distance between slightly longer; anterior seta of coxae II 3rd genital setae; greatest width of genital and III and seta of coxa IV setaceous; plate at level of 2n(l i)air of genital setae. posterior seta of coxae II and III bluntly Anal plate sHghtly wider than long; s|)in(>like; all leg setae setaceous; however, adanal setae of moderate length, extend- some j)reapical setae of tarsi ^la^ be some- ing to base of postanal seta; adanal setae what robust. set at level near posterior end of anal Male. Unknown. orifice; postanal seta considerably larger CoLLKCTION RI'.COROS than adanal setae. Unarmed venter bear- Arl/ioriiys chryso])hiliis ing about 17 [)airs of s(>ta(eous setae, 6 South .XfritiK I roll. feiualo); AMP June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 155

Rhabdomys pumilio concave or invaginated; adanal setae rel- South Africa (Transvaal); 1 female atively long, extending beyond base of (typo specimen); Taufflieb, 1959 postanal seta; adanal setae set at level South Africa (Cape Prov.); 1 female; Taufflieb. 196+ near posterior one-third of anal orifice.

South Africa; 1 coll. {\ female); AMP Unarmed venter bearing 7 or 8 pairs of South Africa (Somerset East. Cape setaceous setae, 4 pairs immediately ad- Prov.); 1 coll. (1 female); AMP jacent to genital and anal plates plus 4 or Otomrs sp. 5 pairs near or on posterior lateral body South Africa (Transvaal) ; 1 female; Tauff lieb. 1959 margins; metapodal plates irregularly oval, slightly longer than wide. Peritreme Rem.\rks.— L. peregrinus and L. rou- extending to level of middle or anterior baudi are quite similar to each other, both of coxa I. Dorsal plate bearing 39 pairs differing from L. tillae in the much mor(^ of setaceous setae; most dorsal setae of robust setae of coxa I. and in the deeper medium length, length equal to or slightly invagination of the posterior margin of greater than distance between adjacent the sternal plate. L. peregrinus may be setae; subterminal setae (J5) reaching distinguished from L. rouhaudi by the fol- slightly beyond posterior margin of dorsal lowing key characters: genital plate dis- |)late. Fourteen to 16 pairs of setae border- tinctly more expanded posteriorly with ing dorsal opisthosoma on soft integument. greatest width at level of 2nd genital Proximal seta of coxa I of medium length, setae; peritreme extends anteriorly only robust, and spinelike, distal seta of coxa to level of middle of coxa II; and larger I short, robust, and peglike; setae ad 1 idiosoma, dorsal plate length greater than and pd 1 of femur I subequal in length; 600 /.... anterior seta of coxae II and III and seta L. peregrinus has been collected to date of coxa IV setaceous, with coxa IV seta only from the country of South Africa, rather small; posterior seta of coxa II almost exclusively from Rhabdomys blunt, peglike but longer, posterior seta pumilio. of coxa III short, robust, and peglike; one preapical seta of each tarsi II and III Laelaps (Laelaps) roubaudi Taufflieb spinelike; most other setae setaceous and Figs. 62-65 normally developed.

T.aelaps roubaudi Taufflieb, 1954, Ann. Parasit. Male: ('Figs. 64-65) Gnathosomal and 29(4) ;437 (Holotype: Brazzaville. Congo; hypostomal setae setaceous; medial hy- Institut Pasteur, Paris); Tipton. 1960, Univ. postomal setae of moderate length, reach- Calif. Publ. Ent. 16(6) :281. ing almost to base of gnathosomal setae. Description.— Female-. (Figs. 62-63) Ventral setae, except adanal and postanal

Dorsal plate length 523 ,". width 374 n.. setae, rather long and slender, extending Gnathosomal and hypostomal setae seta- well beyond base of adjacent posterior ceous; medial hyj)ostomal setae of medium setae; holoventral plate very broad be- length, reaching slightly more than half tween coxae II and III, narrowing be- distance to base of gnathosomal setae. Pos- tween coxae IV, and expanded posterior terior margin of sternal plate moderately to coxae IV; expanded area between geni- invaginated. invagination reaching to or tal setae and anal orifice bearing 5 pairs slightly beyond level of 3rd sternal setae; of setaceous setae; adanal setae of medium

setae st. 1 relatively long, reaching to or length, extending to or slightly beyond almost to level of 3rd sternal setae. An- base of postanal seta; adanal setae set at terior flap of genital plate overlapping level near posterior of anal orifice; post- posterior margin of sternal plate only anal seta somewhat longer and consider- slightly; distance between 1st genital setae ably more robust than adanal setae. Meta- considerably greater than distance be- podal plates inapparent, apparently fused tween 4th genital setae, and distance be- to lateral extensions of holoventral plate; tween 2nd genital setae subequal to dis- unarmed venter bearing approximately tance between 3rd; greatest width of 8 to 10 pairs of setaceous setae adjacent to genital plate at or betw^een 2nd and 3rd holoventral plate, marginal seta longer. genital setae. Anal plate triangular in Peritreme extending to middle of coxa I. general shape, about as wide as long, Dorsal plate bearing 39 pairs of setaceous with anterior margin straight to slightly setae; length and position approximately 156 GREAT BASIN NATURALIST Vol. 36, No. 2

- 62 I vw r-ri//'' 63

Figs. 62-63. Laelaps roubaudi Taufflieb, female. (62) venter; (63) dorsum, scale = 100^.

scale 100^. Figs. 64-66. Laelaps rouhaiuli Taiiffiieb, male. (64) venter; (65) dorsum, = ;

June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 157 as in female. Both proximal and distal results. The 5 taxa of this subgroup differ setae of coxa I setaceous, proximal seta from subgroup A by the presence of one somewhat longer and more robust than or more blunt, peglike setae a})ically on distal seta; setae ad 1 and pd 1 of fenuu" tarsi II and III, but there is no set of key 1 suhecjual and medium in length; an- characters which easily distinguishes this terior seta of coxae II and III, posterior snbgrouj) from subgroup (]. As noted pre- seta of coxa II and seta of coxa IV seta- xiously, L. aethiopicus differs from the ceous; posterior seta of coxa III short, ro- oth(u- taxa of this subgroup in bearing bust, and spinelike; several preapical setae blunt, peglike setae both j)roximally and of tarsi II and III somewhat robust and distally on coxa I; however, it is phenetic- spinelike; most other leg setae seta((Mius ally similar to the taxa of this subgroup and normally developed; however, som(> in overall similarity. may be shorter and rather spinelike.

Collection records Laelaps (Laelaps) nuttalli Hirst Crocidura sp. Figs. 66-72 Nigeria; 1 coll. (1 female); AMP

Tatera kempi Laelaps nuttalli Hirst. 191 '5. Bull. Ent. Res. 6: Dahomey; 1 coll. (4 females); AMP 18S (Holotype: Colombo, Ceylon; British Mu- Ivorv Coast; 2 coll. (3 females, 5 males) seum Natural Historv], London); I Zumpt,

AMP 1950, S. Afr. .1. Med. Sci. 15:87; Keegan, Taterillus nigeriae 1956. J. Egvpt. Publ. Hlth. Assoc. 31:262; 1 Nigeria (northern); 1+ coll. ( -f female); Zumpt and Till, 1958. .1. Ent. Soc. So. Afr. AMP 21:266; Taufflieb, 1959. J. Ent. Soc. So. Afr. Dasymys foxi 22:406; Tipton. 1960. Univ. Calif. Publ. Ent. Nigeria; 2 coll. (4 females); AMP 16:278; Coffee. 1971. Zeitsc.h. Angew. Zool. Dasymys incomptus 58:43-52.

Congo (Brazzaville) ; 1 1 females (type specimens): Taufflieb. 1954 Description.-— Female: (Figs. 66-70) Ivorv Coast; 8 coll. (20 females. Dorsal plate length 621 /-., width 423 /x. 6 ny.); AMP Gnathosomal and hypostomal setae seta- Lophuromys sikapusi Congo (Brazzaville); Taufflieb. 1962 ceous; medial hypostomal setae relatively Pelomys sp. long, not reaching to base of gnathosomal Congo (Brazzaville); Taufflieb. 1962 setae. Posterior margin of sternal plate Praomrs tullbergi only slightly invaginated; setae st. 1 rel- Ghana; 1 coll. (1 female); AMP atively long, reaching almost to posterior Remarks.— As noted previously, L. margin of sternal plate. Anterior flap of roubaudi is most similar to L. peregrinus genital plate slightly overlapping posterior but differs in being a smaller species of sternal plate; distance between 1st geni- (length of dorsal plate less than 6()0 /x). tal setae and 4th genital setae approxi- Also, the peritreme extends distinctly mateh' equal; greatest width of genital further anteriorly (to level of middle of plate at or slightly anterior to 3rd pair coxa I), and the distance between 2nd of genital setae. Anal plate roundly tri- genital setae is no greater than that be- angular, anterior margin somewhat tween 3rd. It may be separated from L. rounded, width approximately equal to tillae by the more robust setae on coxa I length; adanal setae of medium length, and the deeper invagination of the pos- less than distance to postanal seta; adanal terior margin of the sternal plate. setae set at level of posterior end of anal L. roubaudi has been collected from a orifice. Unarmed venter bearing about variety of different hosts in northwest 12 pairs of setaceous setae, 5 pairs adja- Africa south of the vSahara. More speci- cent to genital and anal plates plus about mens have been collected from Dasymys 7 pairs near or on posterior lateral body incomptus than from any other host, but margins; metapodal plates elongate oval. the number of collections from any one Peritreme extending to or nearly to mid- host is species not sufficient to draw ac- dle of coxa I. Dorsal plate bearing 39 curate conclusions on host-parasitic re- pairs of setaceous setae; mo-^t dorsal setae lationships. of medium length, length almost equal to distance between adjacent setae; sub- Subgroup B terminal setae (J5) reaching almost to The formation of this subgroup is based posterior margin of dorsal plate. Ten to primarily on the numerical taxonomic 12 pairs of setae bordering dorsal opistho- Vol. 36, No. 2 158 GREAT BASIN NATURALIST

dorsum, scale lOO.u: (68) ventral Figs 66-70. Laelaps nullalli Hirst, fo.nalo. (66) vonter: (67) (70) vontral vm-u of tarsus IV. scale - 50,«. ,ew of tarsus IT; (69) vo.itral view of tarsus III: June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 159

Figs. 71-72. Laelaps nultalli Hirst, niiile. (71 j venter; (72) doisuni. scale =^ 100/i. soma on soft integiunent. Proximal seta setae of medium length, extending slightly of coxa I setaceous and of medium length, beyond base of postanal seta; adanal setae distal seta relatively short, blunt, and set at level near posterior end of anal peglike; setae ])d 1 and ad 1 of femur I orifice; |)osterior seta approximately twice subequaj in length, with ad 1 seta only as long as adanals antl usually slightly slightly longer; anterior seta of coxae T more robust. Metapodal plates inapparent, and II and seta of coxa IV setaceous. Pos- apparently fused to lateral extension of terior seta of coxae II and III robust and holoventral plate; unarmed venter bear- peglike; tarsi II and III each with two ing approximatelv 12 to 14 pairs of seta- blunt, preapical setae, and tarsus IV with ceous setae adjacent to holo\entral [)late, one blunt, prea[)i(al s(>tae; all other leg those closer to holoventral plate rather setae setaceous and normall}- develo})0(l. short, with those on margins quite long.

to of I. Male: (Figs. 71-72) Gnathosomal and Peritreme extends middle coxa Dorsal of hy{)ostomal setae setaceous; medial hy- plate bearing 39 pairs setaceous setae; length position of postomal setae of moderate length reach- and setae ap- proxiiTiately as in female. Both proximal ing to base of gnathosomal setae. Ventral and distal setae of coxa I setaceous, proxi- setae, except adanal setae, rather long and slender, each extending well beyond base mal seta slightly longer and more robust of posterior seta; holoventral plate rather than distal seta; setae pd 1 and ad 1 of broad between coxae II and III, greatly femur I subecjual in length, jxl 1 seta narrowing between coxae IV, and greatly slightly longer; anterior seta of coxae II expanded posterior to coxae IV; expanded and III, and posterior seta of coxa IV area between genital setae and anal orifice mostlv setaceous but somewhat robust ba- bears 5 pairs of setaceous setae; adanal sal ly: posterior seta of coxa III short, ;

160 GREAT BASIN NATURALIST Vol. 36, No. 2 quite robust, and spinelike to peglike; sev- setae; setae st. 1 long, reaching beyond eral preapical setae of tarsi II and III posterior margin of sternal plate. Anterior rather robust and spinelike; most other leg flap of genital j)late overlapping posterior setae setaceous and normally developed; margin of sternal plate only slightly; dis- however, some may be shorter and some- tance between 1st genital setae distinctly what spinelike. less than distance between 4th genital setae, and distance between 2nd genital Collection records setae distinctly less than distance between Rattus rattus 3rd genital setae; greatest width of genital Belgian Congo (Elizabethville) ; Znnipt. 1961 plate at level of 3rd pair of genital setae. Madagascar; 1 female; Zumpt, 1961, Anal j)late roundly triangular, almost as and Coffey, 1971 wide as long, with anterior margins Madagascar; 4 coll. (65 females, rounded; adanal setae of moderate length, 2 males) ; AMP Mauritius; 4 coll. (12 females); AMP extending to or slightly beyond base of Maslomys coucha postanal seta; adanal setae set at level of South Africa; Tipton, 1960 posterior end of anal orifice. Unarmed Remarks.— L. nuttalli and L. aethio- venter bearing approximately 12 pairs picus may be easily distinguished from the of setaceous setae, 4 pairs adjacent other taxa of subgroup B by the longer to genital and anal plate plus approx- peritreme which extends anteriorly to near imately 6 pairs near or on posterior lateral middle of coxa I. These two taxa also body margins; metapodal plates differ from the taxa of subgroup C, except elongate-oval. Peritreme extending to level for L. myomys, in the same character. L. of middle or at least posterior of coxa I. nuttalli and L. myomys may be separated Dorsal plate bearing 39 pairs of setaceous from L. aethiopicus in that the proximal setae; most dorsal setae relatively long, seta of coxa I is setaceous rather than length equal to or slightly greater than blunt and peglike. L. nuttalli bears smaller distance between adjacent setae; subter- setaceous gnathosomal setae, setaceous minal setae (J5) reaching to or slightly ventral leg setae, and moderate-length beyond posterior margin of dorsal plate. adanal setae rather than robust, spinelike, Eighteen to 20 pairs of setae bordering or peglike ventral leg setae and short dorsal opisthosoma on soft integument. adanal setae as in L. myomys. Both jiroximal and distal setae of coxa I robust, blunt, and peglike, with distal seta L. nuttalli is reported almost exclusively slightly shorter than proximal seta; setae from Rattus rattus in the Ethiopian region. 1 of femur I slightly longer than ad All collections of the African Mammal I)d 1 anterior seta of coxae II and III and seta Project were from this host in Madagascar of coxa IV setaceous; posterior seta of and Mauritius. L. nuttalli is a rather cos- coxae II and III robust, blunt, and peglike; mopolitan mite, occurring worldwide tarsus II with four or five robust, blunt wherever Rattus species are found. preapical setae, tarsus III with three or four blunt, robust preapical setae, and Laelaps (Laelaps) aethiopicus Hirst tarsus IV with two blunt preapical setae; Figs. 73-76 all other leg setae setaceous and normal-

Laelaps aethiopicus Hirst, 1925, Proc. Zool. Soc ly developed. Lond. 4:56 (Holotype: Ashundwa's Camp, Collection records Wanga, Kenya; British Museum | Naural His "Bats" tory], London); 7umpt, 1950, So. Afr. ,1. Med. Soc. 15:78; Badford, 1950, Parasitology 40 Kenya (Ashundwa's Camp, Wanga); Hirst. (3-4):368; Keegan, 1956, Egypt. Publ. Hlth. 1925 Assoc. 31 (6): 258. Ri'.marks. — L. aethiopicus is easily dis- Description.— Female: (Figs. 73-76) tinguished from other taxa of subgroup B

Dorsal plate length 672 /-., width 501 /x. as well as all others of major group II (jnathosomaj and hypostomal setae seta- by the robust, peglike })ro\imal seta of ceous; medial hypostomal setae long, coxa I. Based u]ion this one character reaching almost to base of giialbosonial alone, this mite would be [ilaced in major setae. Posterior margin of sternal plate group III with L. ransomereni; however, slightly invaginated, invagination reach- in overall morphological characters it ing no further than Unol of -Jrd stern;d most closeK' resembles l.. tuittalli. L. June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 161

Figs. 73-76. Laelaps aethiopicus Hirst, female. (73) venter; (74) dorsum, scale = lOO/z; (75) ventral view of tarsus II; (76) ventral view of tarsus III. scale = 50//.

aethiopicus differs from taxa of major mens have yet been recovered from the group III by the following characters: African Mammal Project collections. gnathosomal setae setaceous, never robust and spinelike or peglike; 1st sternal setae Laelaps {Laelaps) liberiensis Hirst long, extending beyond posterior margin Figs. 77-83 of sternal plate; and adanal setae slender and setaceous, not robust and spinelike. Laelaps liberiensis Hirst. 1925, Proc. Zool. Soc. Lond. 4:68 (Holotype: Gonj'on CountpS Li- L. aethiopicus is known only from the beria; British Museum [Natural History], type collection is Hirst which reported by London); Keegan, 1956, J. Egypt. Publ. Hlth. (1925) from Kenya on "Rats." No speci- Assoc. 31:261; Zumpt and Till 1958. J. Ent. 162 GREAT BASIN NATURALIST Vol. 36, No. 2

Figs. 77-81. Lorlaps liberirnsis Hirst, foninlp. (77) voiitiM-; (78) dorsum, scale - lOO/i; (79) ventral view of tarsus II; (80) ventral view of tarsus III; (81) ventral view of tarsus IV, scale = 50/i. June 1976 HERRIN. TIPTON: AITIICAN LAELAPS 163

Soc. So. Afr. 21:266; Taufflich. l<)5y. J. Ent. one blunt preapical seta; all other leg Soc. So. Afr. 22:406; Tipton, 1960, Univ. setae setaceous and normally developed. Calif. Publ. Ent. 16:275; Coffee. 1971, Zeitsch. Angew. Zool. 18:43-46. Ma/e: (Figs. 82-83) Gnathosomal and Laelaps lamborni Hirst. 1925, Proc. Zool. Soc. hypostomal setae all setaceous, with me- Eond. 4:61 (Holotype: Karonga, Nyasaland; dial hypostomal setae moderately long British Mnesinii [Natural History |. London); Zumpt. 1950, So. Afr. J. Med. Soc. 15:78; but not reaching to base of gnathosomal Keegan. 1956, .1. Eg>pt. Puhl. HIth. Assoc. setae. Ventral setae, except adanal and ,31:260; Zumpt and Till. 1958, J. Ent. Soc. So. postanal setae, relatively long, each ex- Afr. 21:266; Taufflieh. 1959, J. Ent. Soc. So. tending in length of Afr. 22:460; Tipton, I960, Univ. Calif. Publ. beyond base seta im- Ent. 16:273; Coffee. 1971, Zeitsch. Angew. mediately })osterior by about one-third its Zool. 58:43-46. length; holoventral plate rather narrow between coxae IV but considerably ex-

Description.^— Female: (Figs. 77-81 ) panded posterior to coxae IV; expanded Dorsal plate length 634 width 432 /(.. ^, area between genital setae and anal orifice Gnathosomal and hypostomal setae seta- bears five pairs of setaceous setae; adanal ceous; medial hypostomal setae moderately setae of moderate length, extending slight- long but not reaching base of gnathosomal ly beyond base of postanal seta; metapodal setae. Posterior margin of sternal plate [)lates not ap})arent; unarmed venter bear- slightly invaginated; setae st. 1 moderate- ing 15 to 20 pairs of setaceous setae vary- ly long, reaching to base of .setae st. 3. ing in length from rather short to quite Anterior flap of genital plate slightly long. Peritreme extending to middle of overlapping posterior margin of sternal coxa II. Dorsal plate bearing 39 pairs of plate; distance between 1st genital setae setaceous setae; length and position of subequal to that between 4th genital setae; setae as in female. Both proximal and distance between 2nd genital setae distinc- distal setae of coxa I setaceous; however, tly less than distance between 3rd genital proximal seta much larger and more ro- setae; greatest width of genital plate at bust with distal seta about half the length; level of 3rd genital setae. Anal plate rela- setae pd 1 and ad 1 of femur I subequal tively triangular, almost as wide as long, in length; anterior seta of coxae II and with anterior margin relatively straight; III and seta of coxa IV setaceous; pos- adanal setae rather long, extending dis- terior seta of coxa II setaceous but some- tinctly beyond base of postanal seta; what robust; posterior seta of coxa III adanal setae set at level near posterior rather short, robust, and spinelike; one end of anal orifice. Unarmed venter bear- or two preapical setae on tarsi II and III ing approximately 10 to 12 pairs of seta- spinelike but not blunt and peglike; all ceous setae, 4 or 5 pairs adjacent to geni- other leg setae setaceous and normally tal and anal plates plus 5 or 6 pairs near developed. or on posterior body margins; metapodal plates oval. Peritreme extending to level Collection records FAephanlulus mrurus of middle or anterior of coxa II. Dorsal South Africa (ORS); 1 coll. (1 female); plate bearing 59 pairs of setaceous setae; AMP most dorsal setae of medium length, ]\ lacroscelides proboscideus length approximately equal to distance South Africa (ORS); 1 coll. (1 female); between adjacent setae; subterminal setae AMP Crocidura sp. extend beyond posterior margins of dorsal Ghana; 2 coll. (3 females); AMP plate. Nine to 12 pairs of setae bordering Crocidura hirtn dorsal opisthosomal on soft integument. Rhodesia; 1 coll. (2 females); AMP Proximal seta of coxa I of moderate length Hvpsignatliae monslrnsus Ivory Coast; 1 coll. d female); AMP and setaceous, distal seta of coxa I rela- Nrcfrris hispi/la tively short, bkmt, and peglike; setae Mauritania; 1 coll. (3 females): AMP pd 1 and ad 1 of femur I subequal in Nrcteris macrotis Senegal; 1 coll. female); length; anterior seta of coxae II and III (1 AMP Rhinolophus simulator and seta of coxa IV setaceous; posterior Rhodesia; 1 coll. fl female) ; AMP seta of coxae II and III robust, blunt, and Eptesicus rapensis South Africa (ORS); 1 coll. (2 females); peglike; tarsus II \\'ith two robust, blunt AMP preapical setae, tarsus III with one blunt, Tadarida leoiiis robust preapical seta, and tarsus IV with Senegal; 2 coll. (3 females); AMP 164 GREAT BASIN NATURALIST Vol. 36, No. 2

Figs. 82-83. Laelaps liberiensis Hirst, male. (82) venter; (83) dorsum, scale 100^.

Tadarida major South Africa; 1 coll. (1 female,

Upper Volta; 1 roll. (1 female); AMP I uy.); AMP Tadarida pumila Tatera sp. Togo; 1 coll. (2 females); AMP Congo (Leopoldvillo) ; 6 females; Galago senegalensis Taufflieb, 1964 Upper Volta; 3 coll. f5 females. Tatera garnbianus 2ny.); AMP Senegal; 3 coll. (1 female, 1 male. Ceropithecus mitis 3ny.);AMP Rhodesia; 1 coll. (1 female); AMP Erythrocehus pata Tatera guineae Ghana; 1 coll. (1 female); AMP Upper Volta; 1 coll. (1 female,

1 male); AMP Tatera kempi

Lepus saxtilis Dahomc^'; I coll. (1 female,

Botswana; 1 coll. (4 females); AMP 1 male); AMP

Iryptomys hnltentotiis Ghana; 1 coll. (12 females); AMP

Botswana; 2 coll. (5 females. Ghana (south); 1 female; 1 male); AMP Paperna et al, 1970

South Africa (ORS); 1 coll. (1 female); Ivorv Coast; 6 coll. (8 fenuiles.

AMP 2 males. 1 nv.) ; AMP

South Africa; 2 coll. (2 females, Upper Volta; \ coll. (1 female); AMP

1 male); AMP Tatera Icucogaster Graphiurus murinus Botswana; 2 coll. (4 females); AMP Nigeria fAfon); 2 females; (Coffey. 1971 South Africa (ORS); 1 coll. Desmodillns auricularis ( I female) ; AMP

South Africa (ORS); 1 coll. (1 female); AMP South Africa; '5 coll. (4 femnl(>s. 1 male); AMP Desniodillus braiirri Upper Volta; 2 coll. (2 females, Tatrrilli/s gracilis

Volta; 1 coll. females. 3 ny.); AMP Upper (5 Gcrhillus paeba 3 ny.); AMP

.South Africa (ORS); 1 coll. (1 female); Taterillus nigeriae

AMP Nigeria (Dada): 1 male; Coffey. 1971 June 1976 HERRIN, TIPTON: AFRICAN LAEL.\PS 165

Dendromus melanotis Mastomys sp.

South Africa (ORS); 1 coll. (2 females); Angola (Dundo); 2 females; AMP Taufflieb. 1962 Malacolhrix typicus Mastomys albicaudatus

South Africa (ORS); 2 coll. (4 females); South' Africa (ORS); 1 coll. AMP (2 females); AMP Steatomys caurinus Mastomys coucha

Ivory Coast; 1 coll. (6 females); AMP Bas-Congo (Boma-Matadi) ; 178 females; AcoTTiYS cahirinus Taufflieb, 1964 Ghana; 1 coll. (1 female); AMP Congo (Brazzaville); Taufflieb, 1962 Aelhomys chrrsophilus Mastomys erythroleucus Botswana; 2 coll. (2 females. Ivorv Coast; 26 coll. (120 females.

1 niiile); AMP 39 males, 129 ny.); AMP

Rhodesia; 1 coll. (1 female); AMP Mastomys natalensis South Africa (ORS); 4 coll. Botswana; 1 coll. (3 females); AMP (6 females); AMP Dahomey; 1 coll. (6 females); AMP South Africa; 22 coll. (32 females, Ghana (Acra-Tema); Papema ct al., 1970 16 males, 4 ny.); AMP Ghana; 100 coll. (321 females. AethoTTiYS namaquensis 64 males, 101 ny.); AMP South' Africa (ORS); 4 coll. (4 females); Ivory Coast; 54 coll. (168 females, AMP 61 males. 39 ny.); AMP

South Africa (Cape) ; 1 female; Nigeria (Panisau); l+colL; AMP Zumpt Taufflieb. 1964 collection Aethomys stannarius Nigeria; 68 coll. (292 females. Nigeria (Ugar. Jabar); 6 females; 94 males. 246 ny.); AMP Coffey. 1971 Rhodesia; 59 coll. (258 females. Arvicanthis niloticus 9 males, 4 ny.); AMP Egypt (El Talbivo. Giza); Keegan, 1956 Senegal; 213 coll. (877 females.

Ghana; 1 coll. (1 female); AMP 112 males, 361 ny.); AMP Ivory Coast; 1 coll. (1 female); AMP South Africa (ORS); 27 coll.

Nigeria; 1 coll. (2 females) ; AMP (50 females); AMP

Nigeria (Panyam Fish Farm); 15 South Africa (Transvaal); 1 female; females. 3 males; Coffey. 1971 Taufflieb, 1964 Nigeria (Ugar, Jabar); 3 females. South Africa; 102 coll. (302 females. 1 male; Coffey. 1971 119 males, 69 ny.); AMP

Cricetomys emini Togo; 4 coll. (10 females, 1 male, Upper Volta; 1 coll. (1 female); 2ny.); AMP AMP Upper Volta; 63 coll. (79 females, Cricetomys gambianus 62males. 89ny.); AMP Nigeria; 1 coll. (1 female, Mus minutoides

1 male) ; AMP South Africa (ORS); 4 coll. Dasymys incomptis (5 females); AMP Rhodesia; 1 coll. (2 females); AMP South Africa; 1 coll. (1 male); AMP Dephomys defua Mus musculus Ghana; 1 coll. (2 females); AMP Egypt (Nahya. Imbaba. Giza); Keegan, 1956

Liberia (Gonvon Country) ; 1 female; Mus musculoides Hirst. 1925 Nigeria (Ibadan); ^ females; Grammomys dolichurus Coffey. 1971

Upper Volta; 1 coll. (3 females); Nigeria (Federal Dist.); 2 females; AMP Coffey. 1971 Hylomyscus alleni Senegal; 1 coll. (1 female); AMP Ghana; 1 coll. (3 females. Togo; 1 coll. (2 females. 5 males, 11 ny.); AMP 1 male); AMP

Togo; 3 coll. (3 females) ; AMP Myomys daltoni Lemniscomys barbarus Ghana; 3 coll. (4 females); AMP

Nigei-ia (Upper Ogum Ranch); 1 Ivory Coast; 1 coll. (1 female); AMP female. 1 male; Coffey, 1971 Senegal; 9 coll. (19 females, Lemniscomys griselda 7 males. 11 ny.); AMP South Africa; 1 coll. (\ female. Praomys daltoni 3 males); AMP Nigeria (Zaria); 2 females; Coffey, 1971

South Africa (Transvaal); 1 female; Praomys fumatus Taufflieb. 1964 Nigeria (lella); Coffey, 1971 Lemniscomys striatus Praomys jacksoni

Nigeria; 2 coll. (6 females, Angola (Dundo) ; 1 female; 4 males); AMP Taufflieb. 1962

Togo; 3 coll. (8 females); AMP Congo (Leopoldville) ; 1 female; Lophuromys sikapusi Taufflieb. 1964

Ghana; 1 coll. (4 females) ; AMP Kenya (Rift Valley Prov.); Nigeria (Ibadan); 1 female; Coffey, 1971 Keegan, 1956 Malacomys longipes Praomys tullbergi Ivory Coast; 1 coll. (2 females, Congo (Brazzaville); Taufflieb, 1962 3 males. 9 ny.); AMP Ghana; 4 coll. (4 females); AMP 166 GREAT BASIN NATURALIST Vol. 36, No. 2

Nigeria; 3 coll. (5 females. has not been reported from the Ethiopian 3 males, 3 ny.); AMP rep^ion, although it does occur in Africa Togo; 1 coll. (1 female, just north of the Sahara. L. liberiensis 1 ny) ; AMP "Rats" differs from L. henoiti by a distinctly Kenya (No. Kitosh & Wamia); shallower invagination of the posterior Hirst, 1925 margin of the sternal plate and the ab- Nyasaland (Karonga); Hirst. 1925 posterior Praomys morio sence of a pair of projections on Cameroon; Taufflieb and Mouchet, 1959 the sternal plate; also, the bodv setae, Congo (Brazzaville); Zumpt, 1961 especially dorsally, are somewhat less ro- Rattus frugivorus bust than in L. henoiti. L. liheriensis is Congo (Brazzaville): Taufflieb, 1962 easil}" separated from L. setzeri by the size Rattus norvegicus Cameroon (Yaounde); Zumpt. 1961 of the posterior central setae of the dorsal Rattus rattus plate. In the former all dorsal setae are Congo (Leopoldville) ; 1 female; medium size to long, whereas in the latter Taufflieb, 1964 the setae of the posterior central area Nigeria (Onitri); Keegan, 1956 Rhabdomys pumilio are much reduced in length. Another Kenya (Njoro. Rift Valley); Keegan, 1956 character which may be used to separate South Africa (ORS); 10 coll. (13 L. liheriensis from other taxa of major females); AMP group II is the presence of only two blunt, South Africa; 6 coll. (16 females, tarsus II, 7 males, 3ny.); AMP peglike preapical setae on and Saccostomus campestris two or three such setae on tarsus III but South Africa (ORS); 4 coll. with only one in the preapical position. (5 females); AMP L. liheriensis was synonymized with L. South Africa; 1 coll. (3 females); AMP Thamnomys rutilans lamhorni by Coffey (1971), a decision i

Togo; 1 coll. (1 female); AMP with which we fully agree. This species ' Uranomys ruddi is the most widely distributed of all Ivory Coast; 1 coll. (1 female); AMP Laelaps species in Africa, both in geo- Otomys irroratus graphic distribution and in host associ- | South Africa (ORS) ; 1 coll.

(1 female); AMP ation. It has been reported from a multi- ' Thryonomys swinderianus tude of hosts throughout the Ethiopian Rhodesia; 1 coll. (1 female); AMP region as in of the as well Africa north j Funisciurus pyrrhopus Sahara (Egypt and Morocco). The host i Ivory Coast; 1 coll. (1 female); AMP

Nigeria (Felele); 1 female; Coffey, 1971 with which it is most closely associated Ictonyx striatus is Mostoniys natalensis; in the collections

South Africa (ORS); 1 coll. (2 females); AMP of the African Mammal Project by far the majority of the collections of this mite Genetta servalina this species. Senegal; 1 coll. (1 female); AMP were from host Genetta villiersi During the examination of specimens Ivory Coast; 1 coll. (1 female); AMP of L. liheriensis from the different locali- Crossarchus obscurus ties and hosts, a certain amount of mor- Ivory Coast; 1 coll. (1 female. phological variability was observed, pri- 1 male) ; AMP Herpestes sanguineus marily in the size and general shape of Rhodesia; 1 coll. (2 females); AMP body structures. The large collection of Unknown host specimens in the African Mammal Project Botswana; 18 coll. (204 females, 12 males, 5 ny.); AMP woidd be ideal for further statistical anal- Ivory Coast; 5 coll. (28 females, yses of inlraspecific variability between

9 males, 3 ny.); AMP ]()( alities and host species. Rhodesia; 1 coll. (7 females); AMP Togo; 1 coll. (8 females); AMP South Afri(a; 35 coll. (73 females. J.dehips {Laelaps) setzeri ('offee 21 males, 29 ny.); AMP Figs. 84-90

Remarks.— L. liberiensis is j)heneti(al- f.aelaps setzeri Coffee. 1971. '/citsihr. Angmv. ly quite close to L. setzeri, L. henoiti, L. nl- Zool. 58:49-51 (Holofype: Tsaiuliagn, North- ern Nigeria; U. S. National Musuem. Wash- frericus, and L. nuttalU. It is easily sepa- ington. F"). C'.). rated from L. nuttaUi by the shorter peri- trerne, and from T.. al^cricus by the lack ni-.sciuPTioN.— Female: (Figs. 84-88) of a heavily seleroli/.cd anterolateral mar- Dorsal plate length 714 /(, witlth '512 /i.

gin of the dorsal plate. Also, /.. (il^crirus (rnathosomal atid hvpostomal setae seta- June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 167

Figs 84-88. Laelaps setzeri Coffee, female. (84) venter; (85) dorsum, scale = 200ju; (86) ventral viewr of tarsus II; (87) ventral viev\' of tarsus III; (88) ventral view of tarsus IV. scale = 50^. 168 GREAT BASIN NATURALIST Vol. 36, No. 2

Figs. 89-90. Laelaps setzeri Coffee, male. (89) venter; (90) dorsum, scale = 100/i. ceous; medial hypostomal setae of moder- terior, lateral, and all marginal dorsal ate length, not reaching to base of gnatho- setae relatively long, but about 7 pairs of somal setae. Posterior margin of sternal posterior central dorsal setae rather small; plate slightly invaginated medially; setae subterminal setae (J5) smallest, reaching St. 1 of moderate length, reaching almost no further than level of base of setae Z5. to level of base of setae st. 3. Anterior Ten to 12 pairs of setae border dorsal flap of genital plate slightly overlapping opisthosoma on soft integument. Proximal posterior margin of sternal plate; distance seta of coxa I setaceous and of moderate between 1st genital setae and 4th genital length, distal seta of coxa I short, robust, setae subequal; distance between 2nd geni- and peglike; setae pd 1 and ad 1 of femur tal setae distinctly less than distance be- I of moderate length and subequal; an- tween 3rd genital setae; greatest width of terior seta of coxae II and III and seta of genital plate at level near 3rd pair of coxa IV setaceous; posterior seta of coxa genital setae. Anal plate triangular in II of moderate length, blunt, and peglike; shape, almost as wide as long, and with posterior seta of coxa III rather short, anterior margin straight; adanal setae of robust, and peglike; tarsus II witli two ro- moderate length, extending to or slightly bust, blunt, peglike preaj)ical setae; tarsi beyond base of postanal seta; adanal setae III and IV each with one rather robust, set at level of posterior end of anal orifice. blunt preapical setae; all other leg setae Unarmed venter bearing 14 to 16 pairs setaceous and normally developed. of setaceous setae, 4 pairs adjacent to geni- Male: (Figs. 89-90) Gnathosomal and tal and anal plates, plus ai)proximately hypostomal setae setaceous; medial hy- 10 to 12 i)airs near or on f)osterior lateral postomal setae of moderate l(>iigth. reach- body margin; metapodal plates irregu- ing almost to base of gnathosomal setae; larly oval, width approximately equal to ventral setae, except adanal and postanal length. Peritreme extending to level of setae, very long and slender, each extend- middle or anterior of coxa II. Dorsal plate ing much beyond base of adjacent pos- bearing 39 pairs of setaceous setae; anterior setae; holoventral ])late broad be- June 1976 HERRIN, TIPTON: AFRICAN L.\EL/\PS 169 tween coxae II and III, extremely nar- Praomys jacksoni rowing between coxae IV, and greatly ex- Nigeria (Igb6-0ra); 2 females; Coffey, 1971 Nigeria (Kudo); I female; panded posterior to coxae IV; expanded Coffey, 1971 area between genital setae and anal ori- Praomys tullbergi fice bearing 5 pairs of setaceous setae; Ghana; 31 coll. (32 females, adanal setae of medium length, extending 21 males. 12 ny.); AMP Ivorv Coast; 39 coll. (64 females, to or slightly beyond base of postanal seta; 62 males, 140 ny.); AMP adanal setae set at level near posterior Nigeria (Sapaba); 1 female; end of anal orifice; postanal seta at least Coffey, 1971 twice as long as adanal setae and some- Nigeria (Federal Dist.); 3 females; Coffey. 1971 what more robust. Metapodal plates inap- Nigeria; 1 coll. (1 female. parent, apparently fused to lateral exten- 1 male. > ny.); AMP sion of holoventral plate; unarmed venter Senegal; 2 coll. (7 females. bearing approximately 12 to 14 pairs of 1 male,); AMP Togo; 63 coll. (149 females, setaceous setae adjacent to holoventral 36 males. 8 ny.); AMP plate, more marginal setae much longer. Rattus rattus

Feritreme extending to middle of coxae Ivory Coast; I coll. (1 female); AMP II. Dorsal plate bearing 39 pairs of seta- Unknown host Ivorv Coast; 1 coll. (1 female); AMP ceous setae; length and position of setae Togo; 1 coll. (2 females); AMP approximately as in female. Both proximal and distal setae of coxa I setaceous; Rem.\rks.— L. setzeri may be easily however, proximal seta much longer and distinguished from other closely related much more robust than short, slender taxa by the following characters: distinct- distal seta; setae pd 1 and ad 1 of femur ly shorter setae on the dorsal plate pos-

I subequal in length, pd 1 seta slightly terior and central in position; tarsus II longer; anterior seta of coxae II, III, and with only two blunt, peglike preapical IV setaceous and somewhat enlarged setae; and the posterior margin of the basally; posterior seta of coxa II of me- sternal plate only slightly invaginated and dium length, rather robust, and somewhat if pair of posterior projections present, spinelike; and posterior seta of coxa III rather small. short, robust, and spinelike to peglike; This taxon is reported primarily from

1 seta of each tarsus II and III robust, Paromys species, most frequently P. tull- blunt, and peglike; several other pairs of bergi, in northwestern Africa south of the setae of tarsi II and III rather robust and Sahara. spinelike; most other leg setae setaceous and normally developed; however, some Laelaps (Laelaps) benoiti Taufflieb may be shorter and rather spinelike. Figs. 91-97

COLLKCTION RECORDS Laelaps benoiti Taufflieb, 1964. Rev. Zool. Bot. Afr. 69(3-4) :377-380 (Holotype: Kibombo. Hipposideros caffer Lwiro, Kivu. Congo-Leopoldville; Musee Royal Ivory Coast; 1 coll. (2 females, de I'Afrique Centrale, Turvuren, Belgium). 1 male) : AMP Scotophilus nigrita Description. — Female-. (Figs. 91-95) Ivory Coast; 1 coll. (6 females); AMP Dorsal plate length 770 /i, width 570 /x. AcoTUYS cahirinus Gnathosomal and hypostomal setae seta- Ghana; 1 coll. (1 female); AMP Malaconiys longipes ceous; medial hypostomal setae relatively Togo; 1 coll. (4 females. long, but not reaching to base of gnatho- 3 males. 1 ny.); AMP somal setae. Posterior margin of sternal Mastomrs natalensis invaginated medially, Togo;' 2 coll. (2 females); AMP plate moderately Mus musculoides with two posterior projections between Nigeria (Ilashe); 3 females; 3rd sternal setae and medial invagi- Coffey, 1971 nation; setae st. 1 of medium length, Togo; 2 coll. (4 females); AMP Praomys alleni reaching to level halfway between 2nd Nigeria (Tsanchaga); 13 females; and 3rd sternal setae. Anterior flap of Coffey. 1971 genital plate overlapping posterior margin Nigeria (Federal Dist.); 2 females; of sternal plate only slightly; distance Coffey. 1971 less Nigeria (Igbo-Ora); 2 females; between 1st genital setae somewhat Coffey, 1971 than distance between 4th genital setae, 170 GREAT BASIN NATURALIST Vol. 36, No. 2

•Figs. 91-95. Laelaps benoiti 'I'auffliob, foniiilc. (91) venter; (92) dorsum, scale =- 200/i; (93) ventral view of tiirsus II; (94) voiiti;il view of tnrsiis III; (9'i) ventral viow of tarsus IV. scalp ^ 50/i. June 1976 HERRIN. TIPTON: AFRICAN LAELAPS 171

distance between 2n(l genital setae less as subterminal setae. Six to 8 pairs of than distance between 3rd genital setae; setae border dorsal opisthosoma on soft greatest width of genital plate at level of integument. Proximal seta of coxa I seta- 3rd genital setae. Anal plate triangular ceous and of medium length; distal seta in general shape, almost as wide as long, of coxa I short, robust, and peglike; setae

with anterior margins straight to slightly [)d 1 and ad 1 of femur I unequal in convex or invaginated; adanal setae of l(Migth, seta j)d 1 somewhat longer than

medimn length, extending slightly be- ad 1 ; anterior seta of coxae II and III and yond base of postanal seta; adanal setae seta of coxa IV setaceous, with coxa IV set at level posterior to anal orifice; post- seta rather small; posterior seta of coxae anal seta over twice as long as adanal seta II and III robust, blunt, and peglike; tarsi and more robust. Unarmed venter bearing II and III each with two rather robust, 12 to 14 pairs of setaceous setae, 4 pairs blunt preapical setae, and tarsus IV with adjacent to genital and anal plates plus one blmit preapical seta; most other leg apjiroximately 8 to 10 pairs near or on setae setaceous and normally developed. posterior lateral body margins; metapodal Male: (Figs. 96-97) Gnathosomal and plates oval, only slightly longer than wide. hypostomal setae setaceous; medial hy- Peritreme extending to level of middle or postomal setae quite long, extending to or anterior of coxa II. Dorsal plate bearing slightly beyond base of gnathosomal setae. 39 pairs of setaceous setae; most dorsal Ventral setae, except adanal setae, rather setae relatively long and robust, length long and somewhat robust, each extending slightly greater than distance between ad- in length well beyond base of seta im- jacent setae; subterminal setae (J5) of mediately posterior or adjacent; holoven- moderate length, reaching distinctly be- tral plate rather broad between coxae II yond posterior margin of dorsal plate; and III, much narrowing between coxae terminal setae (Z5) about twice as long IV, and greatly expanded posterior to

Figs. 96-97. Laelaps benoiti Taufflieb, male. (96) venter; (97) dorsum, scale = lOO/x. ;

172 GREAT BASIN NATURALIST Vol. 36, No. 2 coxae IV; expanded area between genital mately as long as wide. setae and anal orifice bearing 5 pairs of This mite has been collected primarily setae; adanal setae of medium length, ex- from Mus species in northwestern Africa tending well beyond base of postanal seta, south of the Sahara; however, one col- adanal setae set at level slightly posterior lection of four females and one male from to middle of anal orifice; postanal seta Rhodesia has been tentatively identified twice as long as adanals and rather robust. as L. benoiti. Metapodal plate inapparent, apparently fused with lateral extensions of holoven- Subgroup C tral plate; soft integument of opisthosoma bearing 14 to 16 pairs of setae. Proximal As noted in the discussion of subgroup seta of coxa I setaceous and of moderate B, there is no set of distinguishing charac- length; distal seta of coxa I short, robust, ters which can be used to separate this and spinelike; seta pd 1 of femur I slightly group of five taxa from those of subgroup longer than seta ad 1; anterior seta of B; however, in the several numerical taxo- coxae I and III and seta of coxa IV seta- nomic analyses four of these five taxa ceous; posterior seta of coxa II robust and clustered together. spinelike, but posterior seta of coxa III robust and peglike and somewhat shorter; Laelaps (Laelaps) hrandbergensis two setae of each tarsi II and II robust and Taufflieb spinelike rather than blunt; most other leg Figs. 98-104 setae setaceous and normally developed. Laelaps hrandbergensis Taufflieb. 1959. J. Ent. Collection records Soc. S. Afr. 22(2) :400. (Holotype: Brandberg, Southwest Africa; Soutli African Institute for Hipposideros caffer Medical Research, Johannesburg). Ivory Coast; 1 coll. (1 female); AMP Mus bella Description.— Female-. (Figs. 98-102) Congo-Leopoldville (Kwiro, Kivu, Kibombo) Dorsal /i, 2 females; Taufflieb, 1964 plate length 575 width 410 ^. Mus minutoides Gnathosomal and hypostomal setae seta- Ghana; 1 coll. (1 female); AMP ceous; medial hypostomal setae long, Rhodesia; 1 coll. (4 females. reaching to or slightly beyond base of 1 male); AMP of Mus musculoides gnathosomal setae. Posterior margin Ghana (Odomi, Jongo); 1+ coll.; AMP sternal plate slightly invaginated, invagi- Zumpt Collection nation reaching no further than level of Ghana; 3 coll. (3 females); AMP 3rd sternal setae; setae st. 1 long, reach- Ivory Coast; 3 coll. (4 females); AMP ing almost to posterior margin of sternal Nigeria (Ilashe, Igbo-Ora); 3-]- coll.; AMP Zumpt Collection plate. Anterior flap of genital plate Mus setulosus slightly overlapping posterior margin of Ghana; 6 coll. (15 females, sternal plate; distance between 1st genital 2 males); AMP setae slightly greater than distance be- Ivory Coast; 8 coll. (14 females, 4th genital setae; distance between 1 male) ; AMP tween Praomrs tullbergi 2nd genital setae distinctly greater than Ghana; t coll. (1 female); AMP distance between 3rd genital setae; Togo; 1 coll. (1 female); AMP greatest width of genital plate at level of Unknown host of genital setae. Anal plate Togo; 1 coll. (1 female); AMP 2nd pair roiuully triangidar, as wide as long, with Remarks.— L. henoiti may be separated anterior margins slightly rounded; adanal from the other taxa of subgroup B by the setae of moderate length, extending presence of a pair of prominent projec- slightly beyond base of postanal seta; tions on the sternal ])late, with a moderate adanal setae set at level of posterior invagination medially between the projec- end of anal orifice. Unarmed venter bear- tions. In this character this mite resembles ing aproximately 12 pairs of setaceous T.. hrazzai but differs from it in several setae, 6 pairs adjaccMit to genital and anal

other notable characters, i. e., all dorsal plates plus a])proximately () pairs near setae distinctly longer, setae J4 extending or oti posterior lateral body margins; ven- almost to level of .setae .15, setae .15 extend- tral setae all relatively long and somewhat ing beyond posterior margin of dorsal robust; metapodal plates irregularly oval, plate, and sternal ])lnto longer, approxi- \vi(Ith ecpial to length. Peritreme extending June 1976 HEREIN, TIPTON: AFRICAN LAELAPS 173

Figs. 98-102. Laelaps brandbergemis Tauffliob. female. (98) venter; (99) dorsum, scale = lOO/i; (100) ventral view of tarsus II; (101) ventral view of tarsus III; (102) ventral view of tarsus IV, scale = 50/i. 174 GREAT BASIN NATURALIST Vol. 36, No. 2

Figs. 103-104. Laelaps brandbergensis Taufflieb, male. (103) venter; (104) dorsum, scale = 100/z.

to middle of coxa II. Dorsal plate bearing adanal setae, rather long and slender, each 39 pairs of setaceous setae; most dorsal extending well beyond base of adjacent setae relatively long, length usually great- posterior seta; holoventral plate rather

er than distance between adjacent setae; broad between coxae II and III, greatly i subterminal setae (J5) quite short, reach- narrowing between coxae IV, and rather ing no further than terminal setae (Z5). narrow posterior to coxae IV; holoventral Six to 8 pairs of setae bordering dorsal plate posterior to coxae IV with irregular opisthosoma on soft integument. Proximal lateral margins; expanded area between seta of coxa I setaceous and of moderate genital setae and anal orifice bearing length; distal seta of coxa I quite robust, only 3 pairs of setaceous setae, other 2 short, and peglike; seta pd 1 of femur I pairs which are usually on the holoven- rather long, almost twice the length of tral plate set off on soft integument;

seta ad 1 ; anterior seta of coxae II and III adanal setae rather short, not extending and seta of coxa IV slender and setaceous; to base of postanal seta; adanal setae set posterior seta of coxae II and III rather at level of posterior third of anal orifice; robust, blunt, and peglike; tarsi II and III ])ostanal seta 2 or 3 times as long as each with two rather short, robust, blunt, adanal setae and (juite robust; unarmed prea})ical setae, tarsus IV with one mod- venter bearing approximately 14 to 15 erately long, blunt preapical seta; other pairs of setaceous setae adjacent to holo- leg setae mostly setaceous and normally ventral plate, those more anterior and me- developed. dial in position shorter with the more Male: (Figs. 10^-104) (inathosomal and marginal setae rather long. Metapodal hypostomal setae setaceous; medial hy- plates somewhat oval in shape. Peritreme postomal setae of moderate length, reach- extending to middle or anterior of coxa ing at least half distance to base of II. Dorsal plate bearing 39 pairs of seta- gnathosomal setae. Ventral setae, except ceous setae; length and position of setae ;

June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 175 approximately as in female. Proximal distance between 2nd genital setae sub- seta of coxa I rather lon^ and setaceous, equal to distance between 3rd genital with distal seta of coxa I short, robust, setae; greatest width of genital plate at and spinelike; seta pd 1 of femur I ap- level between 2nd and 3rd pairs of genital proximately twice as long as seta ad 1 setae. Anal plate triangular in general anterior seta of coxae II and III, i)osteri{)r shape, anterior margin slightly convex to seta of coxa II, and seta of coxa IV seta- slightly concave; adanal setae of moderate ceous; posterior seta of coxa III shorter, length, extending to or almost to base of robust, and spinelike; 2 or 3 pairs of postanal seta; adanal setae set at level of mostly preapical setae of tarsi II and TIT posterior eiifl of anal orifice. Unarmed short, robust, and j)eglike, with some oth(M- venter bearing a])proximately 12 to 14 setae of tarsi II. III. and IV somewhat [lairs of setaceous setae, 4 pairs adjacent spinelike; most otlier leg setae setaceous to genital and anal plate plus approximate- and normally developed; however, some ly H to 10 pairs near or on posterior may be shorter and somewhat spinelike. lateral body margins; metapodal plate almost circular, \^^dth almost equal to Collection records length. Peritreme extending to level of Prtrnmyscus collinus Soutluvest Africa; 7 females (type middle of coxa II. Dorsal plate bearing specimens); Taufflieb, 1959 39 pairs of setaceous setae; most dorsal South Africa (ORS); 16 coll. setae relatively long, length slightly females); (31 AMP greater than distance between adjacent Aethomrs namaquensis setae; subterminal setae South Africa (ORS); 1 coll. (J5) reaching (1 female); AMP slighth' beyond posterior margin of dor- Re.marks.— The most diagnostic char- sal plate. Approximately 9 pairs of setae bordering dorsal opisthosoma on soft in- acter of L. brandbergensis is the unusually tegimient. Proximal seta of coxa I seta- long pd 1 seta of femur I: the pd 1 seta ceous, distal seta short, blunt, and peglike, is nearl}' two times as long as the ad 1 and approximately half the length seta. All other phenetically similar taxa of proximal seta; setae pd 1 and ad 1 of bear a much shorter pd 1 seta on femur femur I subequal in length, with ad 1 I, only slightly longer than the ad 1 seta. etae slightly longer; anterior seta of rhis mite is known only from southern coxae II III Africa primarily parasitic on Petroniyscus and and seta of coxa IV setaceous; posterior seta of coxae II and III collinus. A single collection is reported from Aethomys namaquensis. robust, blunt, and peglike; tarsi II and III each wdth two rather robust, blunt preapical setae; tarsus IV may have one Laelaps (Laclaps) zumpti Keegan blunt preapical setae or all setaceous; all Figs. 105-111 other leg setae setaceous and normally Laelaps zumpti Keegan. 1956, J. Egypt. Publ developed. Hlth. Assn. 31(6):263 (Holotype: Mjoro, Rift Valley Province. Kenya; U.S. National Mu- Male: (Figs. 110-111) Gnathosomal and seum. Washington, D.C.); Tipton. 1960. Univ. hypostomal setae setaceous; medial hy- Calif. Publ. Ent. 16(6):285. I)ostomal setae long, reaching almost to Description.— Female: (Figs 105-109) base of gnathosomal setae. Ventral setae,

Dorsal plate length 614 /-, \^'idth 401 jx. except adanal setae and postanal seta, of Gnathosomal and hypostomal setae seta- moderate length, each extending in length ceous; medial hypostomal setae long, well beyond base of seta immediately pos- reaching to or almost to base of gnath- terior; holoventral plate filling area be- osomal setae. Posterior margin of sternal tween coxae II and III, narrowing con- plate moderately invaginated, invagination siderably between coxae IV, and greatly reaching to level of Srd sternal setae; setae expanded i)osterior to coxae IV; expanded St. 1 relatively long, reaching beyond level area between genital setae and anal orifice of 2nd pair of sternal pores but not to pos- bearing 5 pairs of setaceous setae; adanal terior margin of sternal plate. Anterior setae relatively short, extending no further flap of genital plate only slightly over- than to base of postanal seta; postanal seta lapping posterior margin of sternal plate; somewhat more robust and about twice as distance between 1st genital setae subequal long as adanal setae. Metapodal plates in- to distance between 4th genital setae, and ay)f)arent or joining holoventral plate lat- 176 GREAT BASIN NATURALIST Vol. 36, No. 2

iFigs. 105-109. Laelaps zumpli Keegan, female. (105) venter; (106) dorsum, scale = lOQ/i; (107) ventral view of tarsus II; (108) ventral view of tarsus III; (109) ventral view of tarsus IV ' scale =-= 50//. ;

Tune 1976 HERRIN, TIPTON: AFRICAN LAELAPS 177

[Figs. 110-111. Laelaps zumpti Keegan, male. (110) venter; (111) dorsum, scale 100/1. erally; unarmed venter bearing 6 to 8 Mus triton pairs of setae adjacent to holoventral plate. Kenya (Rift Valley) ; 1 female, 2 males, 2 ny. (type specimens) Peritreme extending to level of anterior Keegan, 1956 of coxa II. Dorsal plate bearing 39 pairs Remarks.— L. zumpti is quite similar of setaceous setae; length and position as in to L. hrazzai and L. brandbergensis in female. Soft integument of opisthosoma overall characteristics; however, it may be bearing about 8 to 12 pairs of setae. Both easily separated by the unusually short proximal and distal setae of coxae I seta- adanal setae and by the shape of the ster- ceous, with proximal seta considerably nal plate which has a distinctly broader longer than distal seta; setae pd 1 and ad and deeper invaginated posterior margin 1 of femur I subequal in length; anterior and prominent extensions posterior and seta of coxae II and III and seta of coxa lateral to the 3rd sternal setae. In L. IV setaceous; posterior seta of coxa II seta- brandbergensis the posterior margin of the ceous, posterior seta of coxa III spinelike; sternal plate is only slightly invaginated tarsi II and III each with about two pairs with no posterior lateral projections, and of spinelike preapical setae; all other leg in L. brazzai the posterior invagination setae setaceous and normally developed. is only slight and is between two small, Collection records more medial posterior projections. Aethomys chrysophilus This taxon is parasitic primarily on Rhodesia; 1 coll. (1 female); AMP Mus species in the southern half of Africa Lemniscomys striatus from Congo and Kenya to Rhodesia.

Kenya (Rift Valley) ; 1 female; Keegan. 1966 Laelaps {Laelaps) brazzai Taufflieb Mus bella Figs. 112-118 Congo (Leopoldville) ; 10 females.

1 male; Keegan, 1956 Laelaps brazzai Taufflieb, 1962, Acarologia t. IV. Mus minutoides fasc. 4:499 501 (Holotype: Brazzaville, Congo; Rhodesia; 1 coll. (1 female); AMP {jers. coll. of R. Taufflieb). 178 GREAT BASIN NATURALIST Vol. 36, No. 2

Description.— Female: (Figs. 112- reaching almost to base of gnathosomal

116). Dorsal plate length 408 //,, width setae. Ventral setae, except adanal setae,

397 fx. Gnathosomal and hypostomal setae rather long and slender, each extending setaceous; medial hypostomal setae long, well beyond base of adjacent posterior reaching beyond base of gnathosomal seta; holoventral plate rather broad be- setae. Posterior margin of sternal plate tween coxae II and III, greatly narrowing irregular, slightly invaginated mediall}' between coxae IV, and greatly expanded between two small posterior projections; posterior to coxae IV; expanded area be- setae st. 1 of moderate length, reaching tween genital setae and anal orifice bear- halfway between 2nd and 3rd sternal ing 5 pairs of setaceous setae; adanal setae; sternal setae as well as 4 pairs of setae of medium length, extending to or genital setae rather robust. Anterior flap slightly beyond base of postanal seta; of genital plate overlapping posterior mar- adanal setae set at level of posterior third gin of sternal plate slightly; distance be- of anal orifice; postanal seta at least twdce tween 1st genital setae slightly less than as long as adanals and somewhat more distance between 4th genital setae, and robust. Metapodal plates inapparent, ap- distance between 2nd genital setae less parently fused to lateral extensions of than distance between 3rd genital setae; holoventral plate; unarmed venter bear- greatest width of genital plate at level of ing approximately 10 pairs of setaceous 3rd pair of genital setae. Anal plate setae adjacent to holoventral plate, 2 or roundly triangular, almost as wide as 3 posteriorly located pairs rather long, long, with anterior margins irregularly with other more anteriorly located pairs rounded; adanal setae of moderate length, about half this length. Peritreme extend- extending somewhat beyond base of post- ing to middle or anterior of coxa II. anal seta; adanal setae set at level of Dorsal plate bearing 39 pairs of setaceous posterior end of anal orifice; postanal seta setae; length and position of setae ap- rather robust. Unarmed venter bearing proximately as in female. Soft integument approximately 12 pairs of setaceous setae, of opisthosoma bearing approximately 8 5 or 6 pairs adjacent to genital and anal to 10 pairs of setaceous setae. Both prox- plates plus 4 to 6 pairs near or jon pos- imal and distal setae of coxa I setaceous, terior lateral body margins; metapodal ]iroximal seta somewhat longer than distal plates irregularly oval, slightly longer seta; setae pd 1 and ad 1 of femur I than wide. Peritreme extending to level of subequal in length, pd 1 slightly longer; middle or anterior of coxa II. Dorsal plate anterior seta of coxae II and III, posterior bearing 39 pairs of setaceous setae; most seta of coxa II, and seta of coxa IV all seta- dorsal setae of medium length, length ceous; posterior seta of coxa III shorter, usually no greater than distance between robust, and spinelike; no blunt, preapical adjacent setae; subterminal setae (J5) setae on tarsi II, III, or IV; however, some reaching no further than posterior margin preapical setae robust and spinelike; most of dorsal plate. Eight to 10 pairs of setae other leg setae setaceous and normally bordering dorsal opisthosoma on soft in- developed; however, some often shorter tegument. Proximal seta of coxa I of mod- and spinelike. erate length and spinelike, distal seta of coxa I quite robust and peglike; seta pd 1 C>olli:(;tion records CnIom}s fxoslingi of femur I somewhat longer than seta ad Congo-Loopoldville (Lwiro, Biikavu. Kivu): 1; anterior seta of coxae II and III and 2 fonialos; Taufflieb, 1964 seta of coxa IV setaceous, coxa IV seta Dasymys inconiptus rather small; posterior seta of coxae II Congo (Brazzaville); Taufflieh. 1962 and III rather robust, blunt, and peglike; T.rniniscomys striatus tarsi II and III each with 3 blunt, ])re- Congo (Brazzaville); Taufflieb. 1%2 T.nplniromys apuilus rita apical setae, and tarsus IV with 1 or 2 Angola (Dundo); 1 female; blunt |)reapical setae; most other leg setae Taufflieh. 1962 setaceous and normally developed; how- Lophurornys sikapusi ever, .some may be shorter and spinelike. Congo-Leopoldville (Lwiro, Biikavu. Kivu); 2 females; Taufflieb, 1964 Male: (Figs. 117-118) (matho.somal Mastornys natalensis and hypostomal setae setaceous; medial Congo-Leopoldville (Lwiro, Bukavu, Kiwi); hypostomal setae of moderate length. 2 f(>males; Taufflieb. 1964 June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 179

116

Figs. 112-116. Laelaps brazzai Taufflieb, femalo. (112) venter; (113) dorsum, scale = 100/t; (114) ventral view of tarsus II; (115) ventral view of tarsus III; (116) ventral view of tarsus iv'' scale = 50/i. ;

180 GREAT BASIN NATURALIST Vol. 36, No. 2

Figs. 117-118. 'Laelaps brazzai TauHlieh, male. (117) venter; (118) dorsum, scale lOOyu'.

Mus Bella rather long but J5 quite short, not reach- Congo-Leopoldville (Lwiro, Bukavu, Kivu); ing to posterior margin of dorsal plate. 1 female; Taufflieb, 1964 Pelomys foxi L. hrazzai is recorded from a variety Congo-Leopoldville (Lwiro, Bukavu, Kivu) of different hosts by Taufflieb (1962, 1 female; Taufflieb, 1964 1964). No collections of this taxon have Praomys jacksoni yet been identified from the African Angola (Dundo); 3 females; Taufflieb, 1962 Congo-Leopoldville (Lwiro, Bukavu, Kivu); Mammal Project material. 12 females; Taufflieb, 1964 Congo (Musoshi, Elizabethville, Laelaps {Laelaps) myomys, n. sp. Haut-Katanga); 1 female; Taufflieb, 1964 Figs. 119-125 Praomys tullbergi Congo (Brazzaville); Taufflieb, 1962 Holotype, female; type locality: Sedhiou, Cas- Rattus frugivorus manee Region, Senegal; in U.S. National Mu- Congo (Brazzaville); Taufflieb, 1962 seum, Washington, D.C. Rattus verreauxi (Figs. 119- South Africa (Citrusdal, Cape Prov.); Description. — Female: 28 females, I'l males, 3 ny.; Taufflieb, 123) Dorsal plate length 531' /i, width

1964 378 /I. Gnathosomal and hypostomal setae Remarks.— L. hrazzai may be distin- setaceous; medial hypostomal setae long, guished from all other phenetically simi- reaching to or almost to ba.se of gnath- lar taxa by the following characters: pos- osomal setae. Posterior margin of sternal terior margin of sternal plate sHghtly in- plate slightly invaginated medially; setae vaginated medially between pair of St. 1 relatively long reaching almost to rather prominent posterior projections level of 3rd sternal setae; sternal setae posterior and mecHal to setae St. 3; genital as well as 4 pairs of genital setae rather plate somewhat narrower, greatest width long and slender, although somewhat ro- at level of 3rd pair of setae rather than bust basallv. Anterior flap of gcMiital plate at level of 2nd pair; and dorsal setae Z5 overla[)])iiig posterior margin of sternal June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 181

Figs. 119-125. Laelaps myomys n. sp.. female. (119) venter: (120) dorsum, scale = IOOac (121) ventral view of tarsus II; (122) ventral view of tarsus III; (123) ventral view of tarsus IV ' scale = 50/i. 182 GREAT BASIN NATURALIST Vol. 36, No. 2 plate to level slightly anterior to 3rd (J5) reaching almost to posterior margin sternal setae; distance between 1st genital of dorsal plate. Eight to 10 j)airs of setae setae subequal to or slightly less than dis- bordering dorsal opisthosoma on soft in- tance between 4th genital setae; distance tegument. Proximal seta of coxa I of mod- between 2nd genital setae slightly less erate length and setaceous, yet somewhat than distance between 3rd genital setae; robust, distal seta of coxa I quite robust greatest width of genital plate at level of and peglike; seta pd 1 of femur I distinct- 3rd genital setae. Anal plate roundly tri- ly longer than seta ad 1; anterior seta angular to heartshaped, as wide as long, of coxae II and III of moderate length with rounded anterior lateral margins and setaceous but somewhat robust ba- and slightly concave posterior margin; sally; seta of coxa IV shorter and seta- adanal setae of moderate length and slen- ceous; posterior seta of coxae II and III der, extending slightly beyond base of rather large, robust, and peglike; tarsus II postanal seta; adanal setae set at level of with three robust, peglike preapical seta, posterior end of anal orifice; postanal seta tarsus III with two robust, peglike pre- rather long and robust. Unarmed venter apical setae with 2 or 3 pairs of peglike bearing approximately 6 to 8 pairs of setae more proximal in position, and tar- setaceous setae, some rather long with sus IV with 1 blunt peglike preapical seta; others rather short; metapodal plates of most other leg setae setaceous and normal- moderate size, irregularly elongate in ly developed. shape. Peritreme extending to level of an- Male: (Figs. 124-125) Gnathosomal terior of coxa II. Dorsal plate bearing 39 and hypostomal setae slender and seta- pairs of setaceous setae; most dorsal setae ceous. Ventral setae, except adanal setae, slender and rather elongate, length equal rather long and slender, each extending to or slightly greater than distance be- well beyond base of adjacent posterior tween adjacent setae; subterminal setae seta; holoventral plate broad between

Figs. 124-125. Laelaps myomys n. sp., male. (124) venter; (125) dorsum, scale -- 100/i. June 1976 HEREIN, TIPTON: AFRICAN LAELAPS 183

coxae II and III, greatly narrowing be- female (TJM 1173); 1 female (TJM 1188); 1 female (TJM 1213); 1 tween coxae IV, and greatly expanded pos- female and 1 dnv. (TJM 1220); 1 terior to coxae IV; expanded area be- female (WPM 56); 1 female (WPM tween genital setae and anal orifice bear- 69); 1 female (WPM 103); AMP ing 5 pairs of slender setaceous setae; Ivorv Coast (Tyenka); 4 females and 1

1 1 female adanal setae of medium length, extending dnv. (LWR 876); male. add 1 (hiv. (LWR 878); (Bouna); 1 slightly beyond base of j)ostanal seta; male and 1 female (LWR 1611); adanal setae set at level of posterior third (Fetekro); 1 female (LWR 1679); of anal orifice; postanal seta considerably 2 females (LWR 1680-81); 3 females longer than adanal setae. Metapodal plates (LWR 1682-84); 1 females (LWR 1691); 1 male and 3 females (LWR 1693); inapparent, apparently fused to lateral AMP extensions of holoventral plate; unarmed Nigeria (Panyam Fish Farm, Northern venter bearing 6 to 8 pairs of setaceous Region); 1 male and 3 females setae adjacent to holoventral plate. Peri- (HWS 4508); 2 males and 1 female (HJH 1588); (1 mi S Kabvvira, extending to anterior of coxa II. treme Northern Region); 4 females (HWS Dorsal })]ate bearing ^9 j)airs of slender 4588) ; 6 males and 7 females setaceous setae; anterior and lateral dorsal (HWS 4610); AMP setae distinctly longer than posterior cen- Senegal (6 km E Kaolack. Sine-Saloum Region); 1 female (RMD 1547); tral setae. Soft integument of opisthosoma (Koussanar. Oriental Region); 1 female bearing approximately 6 to 8 pairs of seta- (RMD 1804); 2 females (RMD 1847); ceous setae. Roth proximal and distal setae AMP of coxa I setaceous, proximal seta con- Upper Voha (Dio); 1 male and 4 females (REV 1710); 1 female (REV siderably longer and larger than small, 1714); 1 female (REV 1843); (9 km NE slender distal seta; seta pd 1 of f(>mur I Barga); 1 female (REV 1917); (6 km SE somewhat longer than seta ad 1 ; anterior Sequenega); 5 females (REV 2330); seta of coxae II and III and seta of coxa (Ougarou); 5 females (REV 3049); 3 females (REV 3050); 1 female IV setaceous, with coxa IV seta somewhat (REV 3057); 1 female (REV 3066); smaller; posterior seta of coxa II long and 1 female (REV 3067); (5 km SW setaceous, but j)osterior seta of coxa III Koutoura); 5 females (REV 4147); 1 shorter and more spinelike; no blunt, pre- female (REV 4148); 3 females (REV 4169); 2 females (REV 4170); apical setae on tarsi II, III, or IV; how- 2 females (REV 4173); 4 females (REV ever, some tarsal setae may be rather ro- 4190); 1 female (REV 4193); bust and spinelike; most other leg setae (Sideradougou); 2 females (REV 4271); setaceous and normally developed; how- (Djipologo): 1 female (REV 44-14); AMP ever, some often shorter and spinelike. Remarks.-— L. myomys n. sp. differs Type material from most other closely related taxa by Myomys daltoni the three blunt, peglike preapical setae on Senegal: Sedhiou. Casamanee Region; female tarsus II rather than two such setae; how- holotype. allotype, and 8 para- male female ever, it is similar to L. hrazzoi in this one types (RMD 2385); 2 female parat\-pes character but differs in several others. (RMD 2386). This taxon is considerably smaller than Additional collection records other taxa of this subgroup, and it is Nycteris macrotis easily separated from L. hrazzai by the Gambia f Kudang) ; 1 female shape of the sternal plate, i.e., posterior (RMD 2519-27); AMP Tatera Kempii margin only slightly in"\aginated without Upper Volta (Fo); 1 female [)rominent posterior projections. (REV 3755); AMP L. myomys has been collected only from Cricetomys gambianus northwest Africa south of the Sahara pri- Upper Volta (Konankira); 1 female (REV 3279); AMP marily on Myomys daltoni. Only a very Mastomys natalensis few single collections are reported from Upper Volta (5 km. N. Boussouma); other host species. 1 female (REV 1545-46); AMP Myomys daltoni

Gambia (Kudang) ; 4 females Laelnps (Laelaps) malacumys n. sp. (RMD 2517): 1 female (RMD 2518); Figs. 126-132

1 female (RMD 2568) : 1 female (RMD 2570); 1 female (RMD 2573); Holotype. female; type locality: Belekoum, Ivory I female (RMD 2585); AMP Coast; in U.S. National Museum, Washington,

Ghana ( Damongo. Northern Region); 1 D.C. 184 GREAT BASIN NATURALIST Vol. 36, No. 2

Figs. 126-130. Laelaps malacomys n. sp., fcinalo. (126) venter; (127) dorsum, scale = lOO/t; (128) ventral view of tarsus II; (129) ventral view of tarsus III; (130) ventral view of tarsus IV, scale = 50yu. June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 185

Description.— Female: (Figs. 12()- terior third of anal orifice; postanal seta 130) Dorsal plate length 509 /i, width much larger, robust, and spinelike to al- 339 jx. Gnathosonial setae rather long and most jieglike. Unarmed venter bearing 10 quite robust basally; medial hvpostonud to 12 pairs of mostly short, setaceous setae quite long, slender, and setaceous, setae, some almost spinelike; metapodal reaching distinctly beyond base of gnatli- plates irregularly oval, slightly longer osomal setae; other hypostomal setae than wide. Peritreme extending anteriorly smaller and setaceous. Posterior margin to posterior of coxa I. Dorsal plate bearing of sternal plate slightly invaginated, in- 39 pairs of setaceous setae; most dorsal vagination extending no further than lev- setae of medium length; length usually el of 5rd sternal setae; setae st. 1 of mod- no greater than distance between adjacent erate length, reaching halfway between setae; subtorminal setae (J5) of moderate 2nd and '5rd sternal setae; sternal setae length, extending to or slightly beyond as well as 4 pairs of genital setae all of posterior margin of dorsal plate. Approxi- moderate length and setaceous. Anterior mately 10 ])airs of setaceous setae border- flap of genital plate slightly overlapping ing dorsal opisthosoma on soft integument. [posterior margin of sternal plate; dis- Proximal seta of coxa I rather long, ro- tance between 1st genital setae equal to bust, and spinelike; distal seta of coxa I distance between 4th genital setae; dis- (juite short, robust, and peglike; a number tance between 2nd genital setae subequal of ventral setae of leg I short, robust, and

to distance between 3rd genital setae; peglike or spinelike; seta pd 1 and ad 1 greatest width of genital plate at both of femur I subequal in length and rather level of 2nd and 3rd genital setae. Anal robust; anterior seta of coxae II and III plate roundly triangular, approximately of moderate length, somewhat robust and as wide as long with anterior margin spinelike; seta of coxa IV slender and seta- straight to slightly rounded; adanal setae ceous; j)osterior seta of coxae II and III very small, almost minute, yet somewhat rather robust and peglike, with seta of robust; adanal setae set at level of pos- coxa II considerably larger than that of

131

Figs. 131-132. Laelaps malacomys n. sp., male. (131) venter; (132) dorsum, scale = 100/i. . ; ;

186 GREAT BASIN NATURALIST Vol. 36, No. 2 coxa III; tarsi II, III, and IV each with Additional collection records 2 or 3 moderately robust, blunt, peglike Hipposideros commersoni setae; most other leg setae setaceous and Ivory Coast (Yabrasso) : 1 female (I.WR 1528): AMP normally developed; howeAer, ven- some Aethomys chrysophilus tral leg setae often short, robust, and spine- Rhodesia (20 mi N Salisbury, Mashona- like or peglike as those on venter of leg I. land); 1 female (SWG 1747); AMP Lemniscomys slriatus

Male: (Figs. 131-132) Gnathosomal and Ivory Coast (Fetekro); 1 female hypostomal setae setaceous; medial hy- (LWR 1739); AMP Mas setulosus postomal setae long and slender reaching Ivorv Coast (Kahin); 1 female almost to base of gnathosomal setae; gnath- (LWR 763); AMP osomal setae rather short and setaceous. Malaconiys edwardsi Ventral setae, except adanal setae, rather Ghana (Adamso. Ashant Region); 2 females (TJM 1136); AMP long and slender, each extending well be- Ivory Coast (10 mi WNW Soubre) yond base of adjacent posterior seta; holo- ,3 females (LWR 1443); 2 females ventral plate rather broad between coxae (LWR 1449); 4 females (LWR 1451); II and III, greatly narrowing between 1 female (LWR 1475); 23 females and 2 males (LWR (Niebe); 17 females coxae IV, and greatly expanded posterior 1477); (JWL 3061); AMP to coxae IV; expanded area between geni- Malacorny^s longipes tal setae and anal orifice bearing 5 pairs Ghana (Adamso, Ashant Region) of slender, setaceous setae; adanal setae 1 female (WPM 12); AMP Ivory Coast (10 mi Soubre); 4 relatively short, extending to or slightly WNW females (LWR 1450); 1 female beyond base of postanal seta; adanal setae (LWR 1452); 19 females (LWR 1460); set at level of posterior third of anal ori- 13 females (LWR 1462); 2 females fice; postanal seta at least twice as long as (LWR 1476); 4 females (LWR 1478); 5 females (Niebe); adanals and slender. Metapodal plate in- (LWR 1479); 10 females (JWL 3049); 4 females apparent, apparently fused to lateral ex- (JWL 3060); 7 females (JWL 3070); tensions of holoventral plate; unarmed AMP venter bearing approximately 8 pairs of Remarks.— L. nialaconiys is tentatively setaceous setae adjacent to holoventral placed in subgroup C of major group II; plate. Peritreme extending to level of an- however, it differs in several major char- terior of coxa II. Dorsal plate bearing 39 acters: gnathosomal setae robust and pairs of setaceous setae; dorsal setae all longer than medial hypostomal setae; prox- of moderate length, each extending to or imal seta of coxa I robust and long; some slightly beyond base of adjacent posterior ventral leg setae short, blunt, and peglike, seta; subterminal setae (J5) of medium especially on leg I; dorsal setae more ro- length, extending well beyond posterior bust than normal; and peritreme longer, margin of dorsal plate. Both proximal and extending anteriorly to posterior of coxa distal setae of coxa I setaceous, proximal I. In the numerical taxonomic analysis seta somewhat longer than distal seta; L. malacornys clustered with subgroup seta pd 1 of femur I slightly longer than B of major group III because of the simi- seta ad 1 ; anterior seta of coxae II and III lar robust body S(4ae; howe\ er, it is most and seta of coxa IV of medium length and similar to taxa of major group II in other setaceous; posterior seta of coxa II of prominent characters, such as the form medium length and setaceous, but en- of the setae of coxa I (blunt, peglike dis- larged basally; posterior seta of coxa IV tal seta and elongate, setaceous proximal short, relatively robust, and spinelike; pre- seta ) apical setae of tarsi II and III may be somewhat enlarged and spinelike; most This mite j^arasitized Malaconiys spe- other leg setae setaceous and normally cies, with but few exce])tions, in north- developed; however, some may be shorter west Africa south of the Sahara. Single and somewhat s|)inelik('. collections have been made from several other host species. Type material Malaconiys edwardsi Ivory Coast (Belekoum); fonialo liolotvpo Major (h'oup III (LWR 'S7^) male allotype, and 9 female The six taxa of this major group are paratyi)es (I.WR "xSS); 1 female paratvj)e (LWR 584); AMP characterized by the presence of two June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 187

blunt, peglike setae (both proximnlly nnd anal orifice and postanal seta; postanal (Ustally) on coxa I. Even though these seta distinctly longer than adanal seta and species share this one character in com- r(>lati\ely robust. Unarmed venter bear- mon, they form a rather chverse grou}), ing ajjproximately 18 pairs of mostly se- differing from each other in many mor- taceous setae, 6 pairs adjacent to genital phological characters. and anal plates plus approximately 10 to 12 pairs near or on posteriolateral body Subgroup A margins; metapodal plates generally oval in shape. Pcritreme extending to level of The two taxa of this subgroup (L. von- middle or anterior of coxa II. Dorsal somereni and L. acomys) differ from plate bearing 39 pairs of setaceous setae; subgroup B in having simple, setaceous most dorsal setae relatively long, length gnathosomal setae rather than robust, equal to or slightly greater than dis- si)inelike or peglike gnathosomal setae. taiue between adjacent setae; subtermi- L. vnnsotnercni and L. acomys differ iial setae (J5) reaching to or slightly be- from each other in several significant yond posterior margin of dorsal plate.

( haracters: the shape of the sternal plate About 12 pairs of setae bordering dorsal differs greatly, as well as the shape of the opisthosoma on soft integument. Both anal plate. proximal and distal setae of coxa I enlarged, robust, and peglike, with j)roximal Laelaps (Laelaps) vansonicreni Hirst seta considerably more robust than distal 1 of I Figs. 133-139 seta; setae pd 1 and ad femur usually stibequal in length; anterior seta Laelaps vansomereni Hirst. 1923, Ann. Nat. Hist. of coxae II and III and seta of coxa IV 12(67) :690. (Holotype: Busui. S. Bugishu. 3'et robust basally; Uganda; Britisli Museum [Natural History], setaceous, somewhat London); Hirst. 1925, Proc. Zool. Soc. Lond. posterior seta of coxae II and III greatly 4:55; Zumpt, 1950, S. Afr. J. Med. Sci. 15: enlarged, robust, and peglike; tarsi II, 78; Radford. 1950. Parasitology 40(304) :369; III, and IV each with 3 or 4 blunt, robust, Keegan. 1956, J. Egypt. Publ. Hlth. Assoc. setae; most other leg setae se- 31(6):256; Tipton. 1960. Univ. Calif. Publ. preapical Ent. 16(6):284-285. taceous and normally developed, some often rather spinelike. Description.— Female: (Figs. 133-

137) Dorsal plate length 656 /', width Male: (Figs. 138-139) Gnathosomal

466 /'.. Gnathosomal and hypostomal setae and hypostomal setae setaceous; medial setaceous; medial hypostomal setae of hypostomal setae of moderate length, moderate length, reaching only about reaching almost to base of gnathosomal half distance to gnathosomal setae. Pos- setae. Ventral setae, except adanal and terior margin of sternal plate slightly in- postanal setae, rather long and slender, vaginated, invagination reaching no fur- each extending well beyond base of adja- ther than level of 3rd sternal setae; setae cent posterior seta; holoventral plate St. 1 of moderate length, reaching about rather broad between coxae II and III, halfway between setae st. 2 and st. 3. An- f]uite narrow between coxae IV, and terior flap of genital plate overlapping moderately expanded posterior to coxae posterior margin of sternal plate only IV; expanded area between genital setae slightly if at all; distance between 1st gen- and anal orifice bearing 5 pairs of se- ital setae distinctly less than distance be- taceous setae; adanal setae of medium tween 4th genital setae, and distance be- length, extending almost to base of post- tween 2nd genital setae slightly less than anal seta; adanal setae set at level near distance between 3rd genital setae; great- middle of anal orifice; postanal seta est width of genital i)late at level of 3rfl rather slender and setaceous, and only pair of genital setae. Anal })late somewhat slighth- longer than adanal setae; un- broadly oval in general shape, width great armed venter bearing approximately 12 er than length, with margins broadly to 15 })airs of setaceous setae adjacent to rounded; adanal setae rather robust and holoventral plate, all rather slender with spinelike, length extending well beyond more posterior and marginal setae longer. base of postanal seta; anal orifice located Metapodal plates rather elongate. Peri- near anterior margin of anal plate, with treme extending to middle of coxa I. adanal setae set about halfwav between Dorsal plate bearing 39 pairs of setaceous Vol. No. 2 188 GREAT BASIN NATURALIST 36,

135

dorsum, scale 100^1; Figs nS-137. Laelaps vansomereni Hirst, female. (133) venter; (134) = ventral view of tarsus IV, (135) ventral view of tarsus II; (136) ventral view of tarsus III; (137) scale = 50/1. /

June 1976 IIERRIN. TIPTON: AFRICAN LAELAPS 189

138 -^'—

Figs. 138-139. Laelaps vansomereni Hirst, male. (138) venter; (139) dorsum, scale = lOOyu.

setae; length and postion of setae approx- Suncus varilla imately as in female. Soft integument of Southern Africa; Zumpt, 1950 Rkinolophus elivosus opisthosoma bearing approximately 6 to South Africa; 1 coll. (6 females); AMP 8 pairs of setaceous setae. Both proximal Cryptomys hotlentotus

and distal setae of coxa I setaceous, South Africa; 1 coll. (2 females) ; AMP proximal seta considerably longer and Gerbillus paeba Botswana; 1 coll. (1 female); AMP robust slender, shorter much more than Tatera leucogaster distal seta; setae pd 1 and ad 1 of femur South Africa; 2 coll. (11 females,

I rather short and robust, with ad 1 seta 1 male, 2 ny.); AMP Tatera afra somewhat longer than pd 1 seta; anterior Southern Africa; Zumpt, 1961 seta of coxae II and III of medium length, Aethomys chrysophilus rather robust, and spinelike; posterior seta Rhodesia (Bulawayo); Zumpt, 1950 of coxa II of medium length and somewhat Rhodesia; 22 coll. (127 females. setaceous, slightly robust; posterior seta 1 ny.); AMP South Africa (Pretoria, Transvaal); Zumpt, of coxae III short, robust, and spinelike; 1950 and seta of coxae IV shorter and much South Africa (Mfongos, Zululand); more setaceous; 3 preapical setae of tarsus Hirst, 1925 II short, robust, and peglike; some other South Africa (Vaalwater, Nvlstroom Transvaal); Taufflieb, 1964 seta of tarsi II and III shorter, somewhat South Africa (Naboomspruit, Transvaal); robust, and spinelike; most other leg Taufflieb. 1964

setae setaceous and normally developed; South Africa (ORS); 1 coll. however, some may be shorter and rather (18 females); AMP females, spinelike. South Africa; 50 coll. (308 4 males) ; AMP Collection records Aethomys namaquensis Southern Africa; Zumpt, 1950 Elephantulus intufi South Africa; 2 coll. Aethomys selindensis Rhodesia; 4 coll. (38 females); AMP (8 females) ; AMP Suncus etruscus Dasymys helukus iSouthern Africa; Zumpt, 1961 Uganda (Kampala); Tipton, 1960 190 GREAT BASIN NATURALIST Vol. 36, No. 2

Lemniscomys griselda relatively long and somewhat robust. An- South Africa; 1 coll. females); (3 AMP terior flap of genital plate not reaching Mastomys coucha Southern Africa; Zumpt, 1950 to posterior margin of sternal plate; dis-

Sudan (Torit, Equatoria); 1 female; tance between 1st genital setae distinctly Keegan, 1956 less than distance between 4th genital Mastomys natalensis setae; distance between 2nd genital setae Rhodesia; 1 coll. (1 female); AMP distinctly less South Africa; 17 coll. (40 females, than distance between 3rd 2 males, 11 ny.); AMP genital setae; greatest width of genital Southern Africa; Zumpt, 1961 plate at level slightly anterior to 3rd pair Rhabdomys pumilio of genital setae. Anal plate elongate, dis- South Africa; 5 coll. (5 females); AMP Saccostomus campestris tinctly longer than wide; adanal setae of Southern Africa; Zumpt, 1950 moderate length but very robust and South Africa (ORS); 1 coll. spinelike; postanal seta somewhat longer (1 female); AMP and equally robust and spinelike; adanal South Africa; 1 coll. (1 female); AMP "Rodent" setae set at level somewhat posterior to Uganda (Busiu, So. Bugishu); Hirst. 1923 anal orifice. Unarmed venter bearing 6 Uganda (Bumungi, Bugwe); Hirst, 1925 pairs of setaceous setae, anteriormost 5 "Rats" pairs of moderate length, and single pos- Kenya (Okwara's Camp); Hirst, 1925 terior Unknown host pair quite long and slender; meta- Rhodesia; 1 coll. (1 female); AMP podal plates small, oblong-oval. Peri- South Africa; 14 coll. (25 females. treme extending to level of middle or an- 1 male, 2 ny.); AMP terior of coxa II. Dorsal plate bearing 39 Remarks.— L. vansomereni may be pairs of setaceous, rather robust setae; separated from all other taxa of major most dorsal setae of medium length, group III by the following characters: length slightly less than distance be- gnathosomal and hypostomal setae se- tween adjacent setae; subterminal setae taceous; genital plate quite broad through- (J5) quite small, not reaching to pos- out with 1st genital setae considerably terior margin of dorsal plate. Nine closer together than 4th pair; anal plate pairs of medium length setaceous setae wider than long; and adanal setae robust border dorsal opisthosoma on soft integ- and spinelike. ument. Proximal and distal setae of coxa I rather large, robust, and peglike; one This taxon has been recorded from a seta on venter of femur I rather robust variety of different hosts in southern and spinelike; seta pd 1 of femur I some- Africa, with more collections from Aetho- what longer than seta ad ; anterior seta mys species and Mastomys species than 1 of coxae II and III and seta of coxa IV from all others. setaceous; posterior .seta of coxae II and III rather large, robust, and peglike; Laelaps (Laelaps) acomys n. sp. coxae II and III each with one slightly Figs. 140-145 robust, spinelike seta; most other leg setae setaceous and normally develo})ed; how- Holotype, female; type locality: Dunblaine. Mani- caland, Rhodesia; in U.S. National Museum, ever, some, particularly fenun- of each Washington, D.C. leg. often shorter and somewhat spinelike.

Description.— Female: (Figs. 140- Male: (Figs. 144-145) Gnathosomal

143) Dorsal plate length 574 /x, width and hypostomal setae setaceous; medial 421 /x. Gnathosomal and hypostomal setae h3^postomal setae of moderate length, setaceous; medial hypostomal setae me- reaching slightly more than half distance dium length, extending slightly half dis- to base of gnatho.somal setae; gnathosomal tance tf) gnathosomal sotae; gnathosomal setae short, setaceous. Ventral setae, ex- setae of medium length and rather ro- cept adanal and postanal s(^tae. rather bust. Posterior margin of sternal plate long and somewhat robust, each extending considerably invaginated, distinctly be- well beyond base of adjacent posterior yond level of ^rd sternal .setae; anterior seta; holoventral |)late broad between margin of sternal plate arched consider- coxa(» II and III, narrowing considerably ably; setae st. 1 extending to invaginated between coxae IV. and considerably ex- posterior margin of sternal plate; sternal panded posterior to coxae IV; expanded setae as well as 4 pairs of genital setae area between genital setae and anal orifice June 1976 HEREIN, TIPTON: AFRICAN LAELAPS 191

143

Figs. 140-143. Laelaps acomys n. sp., female. (140) venter; (141) dorsum, scale = lOO/i; (142) ventral view of tarsus II; (143) ventral view of tarsus III, scale = 50^. bearing 4 pairs of setaceous setae; adanal Proximal and distal setae of coxa I rather setae of niedium length and quite robust robust and peglike; seta pd 1 of femur I and spinelike; postanal seta some^vhat slightly longer than seta ad 1 ; anterior longer but equally as robust and spinelike; seta of coxae II and III and seta of coxa adanal setae set at level somewhat pos- IV slender and setaceous; posterior seta terior to anal orifice. Metapodal plates of coxae II and III rather robust and peg- small and oval. Unarmed venter bearing like; one preapical seta of tarsi II and III 5 or 6 pairs of setaceous setae adjacent to somewhat robust and spinelike; most other holoventral plate, 2 pairs considerably leg setae setaceous and normally devel- longer than others. Peritreme extending to oped; however, some, particularly on fe- anterior of coxa II and rather broad mora, often short and spinelike. throughout. Dorsal plate with 31 pairs of Type material setaceous setae; length and position of Acomys spinosissineus setae approximately as in female. Soft Rhodesia (Dunblaine. Manicaland) ; female holotvpe. male allotype. 8 female paratopes integument of opisthosoma bearing ap- (SWG 2120-22); 8 female paratypes (SWG proximately 6 jiairs of setaceous setae. 2129); AMP 192 GREAT BASIN NATURALIST Vol. 36, No. 2

144

Figs. 144-145. Laelaps acomys n. sp., male. (144) venter; (145) dorsum, scale = 100/u.

Additional collection records Subgroup B Acomys spinosissineus Rhodesia (3 mi NE Mt. Selinda, Farfell The four species of this subgroup (L. Farm. Manicaland) ; 2 females (HWS paraspinosus. L. hocquieri, L. spinifer, and 5406-58); 1 female 55486-87); HWS L. hreviperitremus) all bear robust, spine- 1 female (HWS 5470); (Chirinda Forest, like Manicaland) 3 females (HWS 5293); or peglike gnathosomal setae, but 2 females (SWG 1560); (Nyamkarara share few other characters in common. River. Manicaland) 7 females (SWG This is a rather diverse group in many 1899-1900); females 5 (SWG 1901-03); niori)hological characters; however, most 3 females (SWG 1946-49); 7 females (SWG 1968-72); (Ngorima Reserve clustered together in the numerical taxo- [East]; Manicaland) 11 females (SWG nomic analvsis. 2147-49); 9 females (SWG 2158-61); 7 females (SWG 2174-76); 5 females rSWG 2176-78); AMP Laelaps {Laelaps) spinijer Taufflieb Unknown liost and Mouchet Rhodesia; 1 female (SWG 2181); AMP Figs. 146-149 Remarks. L. differs — acomys from all r.arlaps spinijer Taufflieb and Mouchet. 1956. other Laelaps species in several unique Aim. Parasit. 31(3):302 (Holotype: Yaounde.

( haracters: anterior margin of sternal plate French Cameroon; Institut Pasteur, Paris); Tipton. 1960. Univ. Calif. Publ. Ent. strongly arched; posterior margin of ster- 16(6): 282. nal plate deeply invaginated; anal plate distinctly longer than wide; and adanal Description. — Female: (Figs. 14(i- setae atul f)ostanal setae robust and spine- 149) Dorsal plate length 438 /i, width like or j)eglike. 285 /I. (juathosomal setae short, robust, This species is known only from Acomys and jieglike; medial hypostomal .setae long, spinosissineus in Rhodesia. It no doubt oc- extending at least to base of gnathosomal curs on this host throughout southern setae; other two pairs of hypostomal setae Africa. short, setaceous. Posterior marijiu of ster- June 1976 HERRINj TIPTON: AFRICAN LAEL\PS 193

h^ '/^" 194 GREAT BASIN NATURALIST Vol. 36, No. 2 extending to level of middle or anterior Description.— Female-. (Figs. 150- of coxa II. Dorsal plate bearing 39 pairs 154). Dorsal plate length 543 /i, width of setae, more anterior and lateral setae 530 p.. Gnathosomal setae stout, robust, somewhat robust, setae il and r4 rather and spinelike to peglike; hypostomal setae small and spinelike, and subterminal setae setaceous, with medial hypostomal setae (J5) very small and setaceous; terminal long, reaching to beyond base of gnatho- setae (Z5) longer than any other dorsal somal setae. Posterior margin of sternal setae. Eight to 10 pairs of setae border dor- plate somewhat invaginated, invagination sal opisthosoma on soft integument, an- reaching no further than level of 3rd terior-most setae short and spinelike, with sternal setae; setae st. 1 rather long, posteriormost setae longer and more seta- reaching almost to posterior margin of ceous. Both proximal and distal setae of sternal plate. Anterior flap of genital coxa I very robust, blunt, and peglike; plate overlai)ping posterior margin of seta pd 1 of femur I somewhat longer sternal plate only slightly if at all; dis- than ad 1; anterior seta of coxae II and tance between 1st genital setae distinctly III somewhat enlarged and spinelike; pos- greater than distance between 4th genital terior seta of coxae II and III quite ro- setae, and distance between 2nd genital bust, blunt, and peglike; seta of coxa IV setae distinctly greater than distance be- small and setaceous; tarsus II with two tween 3rd genital setae; greatest wddth blunt preapical setae, tarsus III wdth one of genital plate at or slightly posterior to blunt preapical seta, and tarsus IV with level of 2nd pair of genital setae. Anal several somewhat spinelike preapical setae; plate roundly triangular, as wide as long, most other leg setae setaceous and normal- with anterior margins rounded; adanal ly developed. setae robust and spinelike and of moderate length, extending almost to base of Collection records postanal seta; adanal setae set at level near Arvicanthis rufinus posterior end of anal orifice; postanal seta French Cameroon (Yaounde); 6 females; Taufflieb and Mouchet, 1956 somewhat longer than adanal setae and Lophuromys aquilus rather robust. Unarmed venter bearing Congo-Leopoldville (Lwiro. Kivu); 2 approximately 50 pairs of mostly seta- females; Taufflieb, 1964 reous setae, some more anterior setae Lophuromys sikapusi rather short and stout wdth more pos- French Cameroon (Yaounde); Zumpt, 1961 terior setae much longer. Metapodal plates Remarks.— L. spinifer possesses a num- oval. Peritreme extending to level of mid- ber of short, spinelike setae ventrally and dle of coxa II. Dorsal plate bearing 38 ^ laterally posterior to coxae IV. It may pairs of setaceous setae, setae px3 absent; be distinguished from other taxa by the most dorsal setae of medium length, length robust, spinelike gnathosomal setae; pos- slightly less than distance between ad- terior margin of the sternal plate irreg- jacent setae; central dorsal setae shorter ularly straight; anal plate unusually wide, than lateral and posterior marginal setae; broadly triangular in general shape; subterminal setae (J5) reaching to or rather long Z5 setae but with tiny J5 setae; slightly beyond posterior margin of dor- and rather short posterior central dorsal sal plate, with terminal setae (Z5) quite setae. long. Approximately 18 pairs of setae L. spinifer has been collected from two bordering dorsal opisthosoma on soft in- Lophuromys species and Arvicanthis tegument. Both proximal and distal setae rufinus in French Cameroon and Congo- of coxa T greatly enlarged, robust, and Leopoldville. peglike or spinelike; ventral setae of trochanter I enlarged and spinelike with proximal posteriolateral seta nnich more

Laelaps (Laelaps) paraspinosus Tipton robust than others; setae pd 1 and ad 1 Figs. 150-156 of femur I subequal in length, with pd 1 seta longer; anterior seta of Laelaps paruulus Hirst (not Berlese, 1904 or Ber- somewhat lese, 1910), 1923, Ann. Nat. Hist., 12(67): coxae IT and III and seta of ( oxa IV rela- 691 (Holotype: South Africa; British Museum tively short, stout, and sjiiiielike to peg- [Natural History], London). like; po.sterior seta of coxae II and III Laelaps paraspinosus Tipton, 1960, Univ. Calif. Publ. Ent. 16(6): 278-280. greatly enlarged, robust, and peglike; ven- June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 195

153

dorsum, scale 100/t; Fiss 150-154 Laelaps paraspinosus Tipton, female. (150) venter; (151) = ventral view of tarsus IV, (152) ventral view of tarsus II; (153) ventral view of tarsus III; (154) scale = 50/i. 196 GREAT BASIN NATURALIST Vol. 36, No. 2

Figs. 155-156. Laelaps paraspinosus Tipton, male. (155) venter; (156) dorsum, scale = lOOiU.

tral anterolateral margin of coxa IV with Peritreme extending to level of middle of serrated, acute, spurlike process; tarsi II, coxa II. Dorsal plate bearing 37 pairs of III, and IV each with 4 to 6 rather robust, setaceous setae, setae px2 and px3 absent; blunt to pointed preapical setae; most length and position of setae as in female. other leg setae setaceous and normally Soft integument of opisthosoma bearing developed; however, some often rather ro- 15 to 20 pairs of setaceous setae. Both bust and spineHke. proximal and distal setae of coxa I en- Male: (Figs. 155-156) Gnathosomal larged, robust, and peglike or spinelike, setae short, robust, and spinelike; hypo- subequal in length; proximal posteriolat- stomal setae setaceous with medial hy- eral setae of trochanter I enlarged, robust, postomal setae longer than others, yet and peglike; setae pd 1 and ad 1 of femur reaching about half distance to gnathoso- I subequal in length and somewhat en- mal setae. Ventral setae, except adanal and larged; anterior seta of coxae II and III postanal setae, rather long, each extending and seta of coxa IV rather short, robust, in length well beyond base of seta im- and jieglike; posterior seta of coxae II mediately j)osterior; holoventral j)lato and Til quite enlarged, robust, and peglike; rather narrow between coxae IV and ventral anterolateral margin of coxa IV throughout entire length, although some- serrated with slender spur; tarsi II, III. what ex})anded ])Osterior to coxae IV; ex- and IV each with 4 to 6 blunt, peglike to panded area between genital setae and spinelike preapical setae; most other leg anal orifice bearing 4 pairs of setaceous setae setaceous; however, some may be setae; adanal setae relatively short, ro- spinelike. bust, and spinelike, length less than distance to postanal seta; postanal seta robust ('OIJ.I'.CTION RECORDS and spinelike, and somewhat longer than Myosntcx variiis Soutli Africa (Caxtoii. Transvaal); adanal setae. Unarmed venter bearing 35 Tipton. 1060 to 40 setaceous setae adjacent to holoven- Aethoniys namnqucnsis tral plate, with an additional 15 to 20 setae Soutii Africa (ORS); 1 coll. d female); on posteriolateral margin, setae closest to AMP bolf)v<'Mfral plate and coxae IV shorter A ri 'icdntliis dnrsalia South Africa; Hirst. 1923 with most posterior and lateral setae (piite f.rnifiiarnnn s' prisridd long. Metapodal i)lat('s rather ('longate. Soutli .'\fii(a; /unipt. 1'161 197 I line 1976 HERRIN. TIPTON: AFRICAN LAELAPS

Rhabdomys pumilio dle of anal orifice. Unarmed a enter bear- South Africa; Zunipt. 1961 ing 5 pairs of setaceous setae adjacent Otomys sp. to genital and anal jilates, no setae on South Africa (Pilgiinis Rest. Transvaal); Tipton. 1960 posterior and lateral margins of body; Otoniys illorat us metapodal plate small, oval. Peritreme ex- South Africa (Grahanistown) ; Hirst. 1925 t(>nding to level of middle or anterior of South Africa (Van Richeeck Nat. Res., coxa I. Dorsal plate bearing 30 to 32 pairs Pretoria); 1 -f coll; AMP Zumpt Collection of mostly setaceous setae; all dorsal setae

South Africa; 1 ( oil. ( 7 females. except setae rl, r2, si, and Z5, extremely 2ny.); AMP minute; setae r2 short and spinelike, setae setae si and Z5 Remarks.— L. paraspinosus bears sev- rl rather robust, and setaceous; position of setae, eral rather unique characters which distin short and absent, not determined be- guisli it from all other Laelaps species: a particular setae

( small size of setae pres- ij;reat many setae ventrally posterior to ause of extremely co.xae IV. lateral to (genital and atial plates, ent. Only one pair of setae apparently and posterolateral to dorsal plate; all co.xal bordering dorsal opisthosoma on soft in- setae and some ventral leg setae short, tegument. Both ])roximal and distal setae robust, and spinelike or peglike; dorsal of coxa I extremely robust and peglike; setae px3 absent with some dorsal setae seta ad 1 of femur I rather short and positioned differently from other Loelaps spikelike, seta pd 1 of femur I about twice species. as long and more setaceous; anterior seta This taxon is known only from South of coxae II and III quite robust and spine- Africa and has been collected from several like; posterior seta of coxa II more seta- different hosts, |)rimarily 0/ornys species. ceous; posterior seta of coxa III quite robust and peglike; seta of coxa IV slender Laelaps (Laelaps) hocquieri Taufflieb and setaceous; all preapical setae of tarsus II setaceous, most with slightly enlarged Figs. 157-163 bases; tarsi III and IV each with one or Laelaps bocquieri Taufflieb. 1962, Acarolngia t. two blunt preapical setae and several IV. Fasc. 4:497-499 (Holotype: Brazzaville. other pairs on tarsi blunt or spinelike; Congo; Pers. coll. of R. Taufflieb, Dakar. Senegal). many other leg setae short and spinelike to setaceous. Description. — Female: (Figs. 157-

KSl) Dorsal plate length 574 //, width Male: (Figs. 162-163) Gnathosomal

^f)5 /I. Gnathosomal setae very robust setae short, extremely robust, and peglike; and peglike; lateral hypostomal setae ro- lateral hypostomal setae somewhat robust and peglike; medial hypostomal and bust, recurved, and peglike; medial and distal hypostomal setae slender, short, distal hypostomal setae slender, setaceous, and setaceous. Posterior margin of sternal and of medium length. Ventral setae, ex- plate slightly invaginated, invagination ( opt adanal and postanal setae, of mod- reaching no further than level of 3rd erate length, extending in length slightly sternal setae; setae st. 1 of moderate beyond base of setae immediately pos- length, reaching slightly more than half- terior; holoventral plate rather broad be- way between setae st. 2 and st. 3. Anterior tween coxae II and III, somewhat narrow- flap of genital })late overlapping posterior ing between coxae IV, and slightly ex- margin of sternal plate to level of 2nd pair panded ])osterior to coxae IV; expanded of sternal pores; distance between 1st area between genital setae and anal orifice genital setae and 4th genital setae sub- bearing only 4 pairs of setaceous setae; equal; distance between 2nd genital setae adanal setae slender and of moderate distinctly greater than distance between IfMigth, extending distinctly beyond base oi 3rd genital setae; greatest width of genital postanal seta; adanal setae .set near level plate at level of 2nd genital setae. Anal of middle of anal orifice; postanal seta plate somewhat oval in general shape, considerably more robust and longer than longer than wide, with anterior and lat- adanal setae. Metapodal plates rather eral margins rounded; adanal setae slen- small, elongate-oval; unarmed venter der and of moderate length but not ex- bearing 4 pairs of setaceous setae. Peri- tending to base of postanal seta; adanal treme extending to middle or anterior of setae set at level slightly posterior to mid- coxa I. Dorsal plate setae as in female. 198 GREAT BASIN NATURALIST Vol. 36, No. 2

159 160

Figs. 1*57-161. Laelaps bocquieri Tnuffliob, female. (157) venter; (158) dorsum, scale = lOO/;: IV. (159) ventral view of tarsus II; (160) ventral view of tarsus III; (161) ventral view of tarsus scale =- 50/i. June 1976 HEREIN, TIPTON: AFRICAN LAELAPS 199

Figs. 162-163. Laelaps bocquicri Taufflieb, male. (162) venter; (163) dorsum, scale = lOOju.

Both proximal and distal setae of coxa I II, and both setae of coxa III robust and greatly enlarged, robust, and peglike; peglike or spinelike; posterior seta of coxa seta ad 1 of femur I short, robust, and II long and setaceous; seta ad 1 of femur spinelike, seta pd 1 at least twice as long I short and spinelike with seta pd 1 twice and rather robust; anterior seta of coxae as long; almost all dorsal setae minute, II and III somewhat enlarged and spine- setae Z5, rl, and si short and setaceous hke; posterior seta of coxa II of moderate and setae r2 short and spinelike. length and setaceous; posterior seta of L. bocquieri has been reported only from coxa III rather short, robust, and peglike; Chryssochloris leucorrhina in the Congo. seta of coxa IV slender and setaceous; tarsus I with 1 blunt preapical seta, tarsus Laelaps (Laelaps) breviperitremus II with 3 moderately long, blunt setae, 1 (Garrett and Strandtmann) being preapical, and tarsus IV with 4 Figs. 164-167 blunt setae, 2 being preapical; other leg Tur breviperitremus Garrett and Strandtmann, setae mostly setaceous; however, some 1967, J. Med. Ent. 4(2):240-246 (Holotype: short and spinelike. Clanwilliam. South Africa; U. S. National Museum, Washington, D.C.) Collection records Laelaps breviperitremus: Furman. 1972, BYU Sci. Chryssochloris leucorrhina Bull, Biol. Ser. 17(3):l-58. Congo (Brazzaville); 24 females, 18 males; Taufflieb, 1962 Description.— Female: (Figs. 164-

Remarks.— L. bocquieri differs from 165) Idiosoma length 890 p.. Gnathosomal all other Laelaps species in several unique setae short, robust, and spinelike; hyposto- characters: gnathosomal and lateral hy- mal setae mostly setaceous, wdth medial postomal setae short, robust, and peglike; hypostomal setae shorter, reaching ap- both setae of coxa I, anterior seta of coxa proximately halfway to base of gnathoso- 200 GREAT BASIN NATURALIST Vol. 36, No. 2 mal setae. Posterior margin of sternal mal setae somewhat more robust; medial plate moderately invaginated, at least to h3'postomal setae of moderate length, level of 3rd sternal setae; all 4 pairs of reaching almost to base of gnathosomal sternal setae short, robust, and spinelike; setae. Ventral setae, except adanal and anterior flap of genital plate overlapping postanal setae, rather long and slender, posterior margin of sternal plate very extending well beyond base of adjacent slightly if at all; genital plate expanded posterior setae; holoventral plate rather considerably posterior to coxae IV and set broad between coxae II and III, greatly very close to anal plate with posterior mar- narrowing between coxae IV, and greatly gin invaginated to accommodate anal expanded posterior to coxae IV; expanded plate; 3 pairs of genital setae rather short, area between genital setae and anal orifice robust, and spinelike, and only first 3 bearing 5 pairs of setaceous setae; adanal pairs set on genital plate; distance be- setae of moderate length, extending well tween 1 St genital setae much less than dis- beyond base of postanal seta; adanal setae tance between 4th genital setae, and dis- set near middle of anal orifice; postanal tance between 2nd genital setae distinct- setaceous; posterior seta of coxae II and ly less than distance between 3rd genital seta somewhat longer than adanals but setae; greatest width of genital plate at much more robust and spinelike. Meta- level of 3rd genital setae. Anal plate podal plates inapparent, apparently fused roundly triangular, almost as wide as to lateral extension of holoventral plate; long; adanal setae of moderate length, ex- unarmed venter bearing approximately tending somewhat beyond base of post- 10 to 12 pairs of slender setaceous setae anal seta; adanal setae at level slightly adjacent to holoventral plate. Peritreme posterior to middle of anal orifice; post- short, extending no further than posterior anal seta very large, rather long and ro- of coxa II. Dorsal plate bearing 40 pairs bust. Unarmed venter bearing approxi- of setaceous setae; most dorsal setae of mately 12 to 14 pairs of setaceous setae, moderate length, usually greater than dis- all rather long and most barbed; meta- tance between adjacent setae; subterminal podal plates irregularly oval, slightly setae (J5) of medium length, extending longer than wide. Peritreme very short, well beyond posterior margin of dorsal extending no further than posterior plate, but considerably shorter than of coxa II. Forty-one pairs of setae associ- terminal setae. Both proximal and distal ated with dorsal plate; more anterior setae setae of coxa I setaceous, proximal seta short, robust, and spinelike, with posterior 'Somewhat longer than distal seta; setae and posterior marginal setae longer and pd 1 and ad 1 of femur I subequal in more setaceous; subterminal setae (J5) length; anterior seta of coxae II and III, long and slender with terminal setae some- posterior seta of coxae II and III, and what longer and more robust. Approxi- sota of coxa IV all setaceous, but some mately 12 pairs of slender, setaceous setae may be robust basally; some preapical border dorsal opisthosoma of soft in- 9otae of tarsi TI, III, and IV robust and tegument. Both proximal and distal setae "^pinelike; most other leg setae setaceous of coxa I robust, blunt, and peglike, with and normally developed; however, some proximal seta somewhat larger; setae ad may be shorter and rather spinelike. 1 and pd 1 of femur I subequal in length; proximal posterior seta of trochanters I Coi,r,r.cTioN rkcords and II short, robust, and peglike; an- Arntnvs siihspinosus Soutli Africa (Pakliuis Pass, Clanwilliam. terior seta of coxae II and III of mod- Traiisviinl); 28 females (type specimens); erate length and setaceous, yet somewhat Garrett and Strandtmann. 1967 robust basally; seta of coxa IV short and South Africa (Goudveld, Cape Prov.);

III rather short, robust, and peglike; 29 females, 1 male, ^ ny.; tarsi II, III, and IV each with 3 to 5 Garrett and Strandtmann, 1967 blunt, peglike preapical setae; most other Rkmarks.— Garret and Strandtmann leg setae setaceous and normally devel- (1067) originally placed L. brcvipcritrem- oped; however, some may be shorter and i/s in the genus because of many rather spinelike. 7ur morphologic al characters possessed in com- Male: (Figs. 166-167) Gnathosomal and mon with 7'i/r which differ from any other hypostomal setae setaceous, with gnathoso- Ldi'hips species. Some of these distinguish- June 1976 IIKRRIN, TIPTON: AFRICAN LAEL.\PS 201

164 165 I

Figs. 164-165. Laelaps breviperitremus (Garrett and Strandtmann). female. (164) venter; (165) dorsum; redrawn from Garrett and Strandtmann (1967).

Figs. 166-167. Laelaps breviperitremus (Garrett and Straiultmaiin;. male. (166) venter; (167) dorsum, redrawn from Garrett and Strandtmann (1967).

ll 202 GREAT BASIN NATURALIST Vol. 36, No. 2 ing characters are: very short peritreme, Elephantulus myurus L. jritzumpti - South Africa (ORS) extending to posterior of coxa II; many L. liberiensis - South Africa (ORS) short, robust, spinelike setae ventrally and Elephantulus rupestris dorsally; only three pairs of setae on geni- L. fritzurnpti - South Africa (ORS) tal plate; greatly expanded genital plate; Macroscelides proboscideus L. fritzurnpti - South Africa (ORS) plus other less obvious phenotypic differ- L. liberiensis - South Africa (ORS) ences. L. simillimus - South Africa (ORS) L. breviperitremus is known only from L. transvaalensis - South Africa Acomys subspinosus in South Africa. (ORS) Superfamily Soricoidea Family Soricidae Host-Parasite Relationships Subfamily Crocidurinae Crocidura hirta For the most part, species of Laelaps in L. liberiensis - Rhodesia Africa are associated with myomorph ro- Crocidura sp. dents and more particularly rodents of the L. keegani - Upper Volta subfamily Murinae. However, there are L. liberiensis - Ghana L. roubaudi - Nigeria some exceptions to this statement. For ex- Sylvisorex gemmeus ample, L. transvaalensis L. paraspino- and L. lavieri - Ghana sus were collected from Otornys sp. (sub- Order Chiroptera family Otomyinae) more frequently than Suborder Megachiroptera from other hosts, but they were collected Family Pteropidae Subfamily Pteropinae from murine rodents as well. L. brand- Hypsignathae monstrosus bergensis has been collected principally L. lavieri - Ivory Coast from Petromyscus sp. (subfamily Dendro- L. liberiensis - Ivory Coast murinae), but again some specimens were Suborder Microchiroptera collected from murine rodents. L. con- Family Nycteridae goicola, L. moucheti, L. aethiopicus, and Nycteris arge L. lavieri - Upper Volta L. bocquieri are known only from single Nycteris hispida type collections in which the hosts were L. liberiensis - Mauritania not identified beyond "rat" or "rodent" Nycteris macrotis or the specific identification of the host L. liberiensis - Senegal cannot be confirmed. In the northern part Family Rhinolophidae Subfamily Rhinolophinae of Africa gerbils (Gerbillinae) are fre- Rhinolophus clivosus c}uently associated with species of Laelaps^ L. vansomereni - South Africa but the Laelaps species involved are ubiq- Rhinolophus simulator uitous and are associated with such a va- L. liberiensis - Rhodesia Subfamily Hipposiderinae riety of hosts that the true host-parasite Hipposideros baetus is relationship obscure. Contaminations L. lavoipierrei - Ivory Coast which may have occurred in the field or Hipposideros coffer laboratory may account for other unusual L. benoiti - Ivory Coast L. lavieri - Ivory Coast associations recorded in the list given be- /-. setzeri - Ivory Coast low. vSpecimens which were actually Hipposideros comniersoni found on nonmurine hosts likely repre- L. malacomys - Ivorj' Coast sented spurious associations. Hipposideros cyclops L. lavieri - Ivory Coast New collection records of species of Family Vespertilionidae Laelaps from the African Mammal Proj- Subfamily- Vespei'filioninae capensis ect. (ORS = Orange River Survey) Eptisicus L. liberiensis - South Africa (ORS) Order Insectivora Scotophilus nigrita Superfamily Erinaceoidea L. setzeri - Ivory Coast Family Erinaceidae Family Molossidae Subfamily Erinaceinae Ta/larida Ironis

Atelerix albiventris L. liberiensis - Senegal

L. keeaani - Upper Volta Tadarida major

Superfamily M;i( rostelidoidea L. grenieri - Upper Volta

Family Mac losc clididae L. liberiensis - Upper Volta FJephantulus intufi Tadarida midas L. keegani - South Africa (ORS) L. simillirinis South Africa L. simillimus - South Africa Tadarida pumila

L. vansomercni - South Africa L. liberiensis Togo June 1976 HEREIN, TIPTON: AFRICAN LAEL.\PS 203

Order Primata Malacothrix typicus Family Lorisidae L. liberiensis - South Africa (ORS) Subfamily Galaginae Stealomys caurinus Galago senegalensis L. liberiensis - Ivory Coast

L. liberiensis - Upper Volta Petromyscus collinus Family Cercopitlieridae L. brandbergensis - South Africa Subfamily Cercopithecinae (ORS) Cercopithecus rnitis L. fritzumpti - South Africa (ORS) L. liberiensis - Rhodesia Subfamily Murinae Erythrocebus pata Acomys cahirinus L. liberiensis - Upper Volta L. liberiensis - Ghana Order Lagomorpha L. setzeri - Ghana Familj' Leporidae Acomys spinosissineus Lepus saxadlis L. acomys - Rhodesia chrysophilus L. liberiensis - Botswana Aethomys L. fritzumpti - South Africa Order Rodcntia (ORS), Rhodesia Suborder Hystricomorpha L. lavieri - South Africa (ORS), Superfamily Bathyergoidea Rhodesia Family Bathyergidae L. liberiensis - South Africa Cryptomys hottentotus (ORS), Rhodesia, Botswana L. liberiensis - Botswana, L. malacornys - Rhodesia South Africa L. simillimus - Botswana. South L. transvaalensis - South Africa Africa, Rhodesia L. vansomereni - South Africa tillae - South Africa Family Thryonomyidae L. L. transvaalensis - South Africa Thryonomys swinderianus L. vansomereni - Soutli Afnca L. liberiensis - Rhodesia (ORS). Rhodesia Superfamily Octodontoidea L. zumpti - Rhodesia Family Petromyidae Aethomys namaquensis Petromus typicus L. brandbergensis - South Africa L. transvaalensis - South Africa (ORS) (ORS) L. fritzumpti - Botswana. Suborder M>omorpha South Africa (ORS) Superfamily Muroidea L. liberiensis - Soutli Africa Family Cricetidae fORS) Subfamily Gerbillinae L. paraspinosus - South Africa Desmodillus auricularis (ORS) L. fritzumpti - South Africa (ORS) Aethomys selindensis L. liberiensis - South Africa (ORS) L. vansomereni - Rhodesia Desmodilliscus braueri Arvicanthis niloticus L. liberiensis - Volta Upper L. keegani - Ghana, Ivory Coast, Gerbillus paeba Nigeria, Senegal L. - South Africa fritzumpti (ORS) L. liberiensis - Ghana, Ivory L. liberiensis - South Africa (ORS) Coast, Nigeria L. vansomereni - Botswana Cricetomys emini Tatera brandsi L. liberiensis - Upper Volta L. fritzumpti - South Africa (ORS) Cricetomys gambianus Tatera gambiana L. liberiensis - Nigeria L. liberiensis - Senegal L. myomys - Upper Volta Tatera guineae Dasymys foxi L. liberiensis - Guinea L. roubaudi - Nigeria Tatera kempi Dasymys incomptis L. keegani - Dahomey L. liberiensis - Rhodesia L. liberiensis - Ghana, Dahomey. L. roubaudi - Ivory Coast Ivory Coast, Upper Volta Dephomys defua L. - Upper Volta myomys L. liberiensis - Ghana L. - Ivor>' roubaudi Coast L. parasimillimus - Ivory Coast Tatera leucogaster Grammomys dolichurus L. fritzumpti - South Africa (ORS) L. liberiensis - Upper Volta L. lavieri - South Africa (ORS) Hybomys trivirgatus L. liberiensis - South Africa (ORS). L. grenieri - Ivory Coast Botswana Hylomyscus alleni L. simillimus - South Africa L. liberiensis - Togo L. transvaalensis - South Africa Hylomyscus sp. L. - Soutli vansomereni Africa L. liberiensis - Ghana Taterillus gracilis Lemniscomys barbarus L. liberiensis - Volta Upper L. grenieri - Ghana. Upper Volta Family Muridae Lemniscomys griselda Subfamily Dendromurinae L. liberiensis - South Africa Dendromus melanotis L. simillimus - South Africa, L. liberiensis - South Africa (ORS) Rhodesia 204 GREAT BASIN NATURALIST Vol. 36, No. 2

L. tillae - South Africa. Rhodesia Coast, Senegal, Upper Volta L. vansoTTiereni - South Africa L. myomys - Ghana. Ivorj' Coast, Lemniscomys macculeus Nigeria. Senegal, Upper Volta L. grenieri - Ivory Coast L. transvaalensis - Senegal L. lavieri - Ivory Coast Praomys tullbergi Lemniscomys striatus L. benoiti - Ghana, Togo L. grenieri - Ghana, Ivory L. grenieri - Nigeria, Togo Coast, Nigeria, Togo L. lavieri - Ghana, Ivory Coast L. lavieri - Togo L. lavoipierrei - Ivory Coast L. liberiensis - Nigeria, Togo L. liberiensis - Ghana, Nigeria, Togo Lophuromys sikapusi L. roubaudi - Ghana

L. grenieri - Nigeria L. setzeri - Ghana. Ivory Coast. L. lavieri - Ghana Nigeria, Senegal, Togo L. lavoipierrei - Ghana. Ivory L. thamnomys - Togo Coast, Nigeria Rattus rattus L. liberiensis - Ghana L. nuttalli - Madagascar, Malacomys edwardsi Mauritius L. malacomys - Ghana. Ivory Coast L. setzeri - Ivory Coast Malacomys longipes Rhabdomys pumilio L. liberiensis - Ivory Coast L. fritzumpti - South Africa (ORS) L. malacomys - Ghana, Ivory Coast L. liberiensis - South Africa (ORS)

L. parasimillimus - Ivory Coast L. peregrinus - South Africa (ORS) L. setzeri - Togo L. simillimus - South Africa Mastomys ulbicaiulafus L. tillae - South Africa L. liberiensis - South Africa (ORS) L. transvaalensis - South Africa Mastomys erythroleucus L. vansomereni - South Africa L. liberiensis - Ivory Coast Sacrostomus campestris Mastomys natalensis L. fritzumpti - South Africa (ORS) L. fritzumpti - South Africa (ORS) L. lavieri - South Africa (ORS) L. lavieri - Ghana. South L. liberiensis - South Africa (ORS) Africa (ORS) L. tillae - Rhodesia L. lavoipierrei - Ivory Coast, L. transvaalensis - South Africa Upper Volta L. vansomereni - South Africa L. liberiensis - Botswana, Rhodesia, (ORS) South Africa (ORS), Dahomey, Thallomys paedulcus Ghana, Ivory Coast. Nigeria, L. fritzumpti - South Africa (ORS) Senegal. Togo, Upper Volta Thamnomys rutilans L. myomys - Upper Volta L. liberiensis - Togo L. setzeri - Togo L. thamnomys - Ivory Coast. Togo L. simillimus - South Africa Uranomys oweni (ORS) L. grenieri - Senegal L. tillae - South Africa Uranomys ruddi L. transvaalensis - South L. grenieri - Ivory Coast Africa (ORS) L. lavoipierrei - Ghana L. vansomereni - South Africa, L. liberiensis - Ivory Coast Rhodesia Subfamily Otomyinae Mas haussa Otomys angoniensis L. lavieri - Nigeria L. transvaalensis - Rhodesia, Mus minutoides South Africa (ORS) L. benoili - Ghana, Rhodesia Otomys irroratus

L. fritzumpti - South Africa (ORS) L. liberiensis - South Africa (ORS')

L. lavieri - Ghana, Ivory Coast. L. paraspinosus - vSoutli Africa

South Africa (ORS). Rhodesia L. transvaalensis - South Africa L. liberiensis - South Africa (ORS) Parotomys brantsi L. ziimpti - South Africa L. fritzumpti - South Africa (ORS) (ORS). Rhodesia Suborder Sciuroniorpha Mus musculoides Superfamily Sciuroidoa L. benoiti - Ghana. Ivory Coast Famil>' Sciuridae L. keegani - Ghana Subfamily Sciurinae T.. lavieri - Ghana. Ivory Coast, Funisciurus pyrrhopus

Upper Volta L. liberiensis - Ivor>' Coast L. lavoipierrei - Ghana Order Carnivora L. liberiensis Senegal, Togo I'^amil^' Mustelidae L. setzeri Togo Subfamily Mustelinae L. - thamnomys Togo Ictonvx striatus

Mus setulosus I., lavieri South Africa (ORS)

L. benoiti Ghana, Ivory Coast /., liberiensis - South Africa (ORS) A. lavieri Ghana. Ivory Coast Fiimil\ \'ivori(lae L. malacomys - Ivory Coast Suhfamily Viverinae Myomys daltoni Genetta servalina

L. liberiensis (ihaiia. Ivory L. liberiensis - Senegal June 1976 HERRIN, TIPTON: AFRICAN LAELAPS 205

Genetta villiersi MiCHENER, C. D., AND R. R. SoKAL. 1957. A L. lavieri - Ivory Coast quantitative approach to a pioblem in clas- L. liberiensis - Ivorj' Coast sification. Evol. 11:130-162. Subfamil\' Heipestinae PaPERNA, I.. D. P. FlIRMAN, AND N. RoTHSTEIN. Crossarchus obscurus 1970. The parasitic fauna of rodents from L. liberirnsis - Ivory Coast urban and suburban areas of Accra-Tema, Herpesles sanguineus South Ghana. Rev. Zool. Bot. Afr. 81(3-4): L. liberiensis - Rhodesia 330-336. Family Felidae Radford, C. D. 1950. The mites (Acarina) Subfamily Folinae parasitic on manunals, birds, and reptiles. lybicae Felis Parasitology 40(34) : 366-394. L. keegani - Upper Volta SoKAL, R. R. 1961. Distance as a measure of taxonomic similarity'. Svst. Zool. 10:70-79. References Cited SoKAL, R. R., AND P. H.' A. Sneatii. 1973. Principles of numerical taxonomy. W. H. and Co.. San Francisco and Bedford, G. A. H. 1932. A synoptic check-list Freeman Lon- don, and host-list of ectoparasites found on South 359 pp. Taufflieb, R. 1954. species of African mammalia, aves and reptilia (2n(l Some new (Acarina: Laelaptidae) the ed.) 18th Rep. Dir. Vet. Ser. and Animal Laelaps from middle Congo (French Ecjuatorial Africa). Ind.. Aug. 1932. pp. 223-520. Ann. de Parasitol. Humaine et Comparee. . 1936. A synoptic check-list and host- 436-446. list of the ectoparasites found on South Afri- 29(4): can mammalia, aves and reptilia (Supplement . 1959. Nouvelles especes de Laelaps de la region Ethiopienne (Acarina: Laelaptidae) no. 1). Onderstepoort J. Vet. Sci. 7:69-110. Coffee. G. M. 1971. Discussion on the Laelaps et cle des especes Africaines du genre. J. Ent. vSoc. 394-408. nuttalli Hirst comple.\ with description of a So. Afr. 22(2): new species from the Ethiopian region (Aca- . 1962. Deux nouveles especes de Laelaps rina: Mesostigmata. Demianyssidae). Zeit- (Acarina: Laelaptidae) de la region de Braz- t. 4:497- schr. angew. zool. 58:43-52. zaville (Congo). Acarologia, 4. fasc. Evans, G. O.. .^nd W. M. Till. 1965. Studies 502. on the British Dermanyssidae (Acari: Meso- . 1964. Notes sur les Laelaps du Congo- Leopoldville d'apres les collections du museede stigmata) Part I. External morphology. Bull. :377- British Mus. (Nat. Hist.) Zool. 13(8): Tervuren. Rev. Zool. Bot. Afr. 69(3-4) 249-293. 383. FuRM.AN, D. P. 1972. Laelapid mites (Laelapi- Tatjfflieb, R., and J. Mouchet. 1956. Deux dae: Laelapinae) of Venezuela. Brigham nouvelles especes de Laelaps (Acarina: Lae- Young ITniv. Sci. Bull.. Biol. Ser. 17(3): laptidae) du Canuneroun Francais. Ann. 1-58. Parasitol. Humaine et Comparee. 31(3):302- Garrett, L. E.. and R. W. Str,andtmann. 1967. 307. Tur breviperitremus n. sp., a new laelapine TnuRMAN. E. B. 1958. Laelaps keegani, nom. mite from South Africa. J. Med. Ent. 4(2): nov. for Laelaps berlesi Keegan, 1956. Ent. 240-246. Soc. Wash. 60 (2): 74. HiR.scHMANN. W. 1957. Gangsystomatik der TiPTON, V. J. 1960. The genus Laelaps with a Parasitiformes. Teil I. Rumpfbehaai-ung und review of the Laelaptinae and a new sub- RuckenflacJien. Schriftenreihe fur Vergleich- family Alphalaelaptinae (Acari: Laelaptidae). ende Milbenkunde. 20 pp. Univ.^ Calif. Publ. Ent. 16(6) :233-356. HiR.sT. S. 1915. On some new acarine parasites F. 1950. Descriptions of two Lae- of rats. Bull. Ent. Res. 6:183-190. ZuMPT. new laps species from South Africa with a key . 1923. On three new species of Laelaps this (Aca- occurring on African rodents. Ann. Mag. to the Ethiopian species of genus rina: Laelaptidae). So. Afr. J. Med. Sci. 15: Nat. Hist., Ser. 9. 12(67) :690-692. 77-82. . 1925. Description of new Acari, mainly

. 1961. arthropod parasites of verte- parasitic on rodents. Proc. Zool. Soc. London, The in of tlie Sahara (Ethio- pt. 1:49-69. brates Africa south I Publ. So. Kkegan, H. L. 1956. Ectoparasitic laelapid and pian region). Vol. (Chelicerata). 1-457. dermanyssid mites of Egypt. Kenya and the Afr. Inst. Med. Res. 9(1): Sudan, primarilv based on NAMRU-3 col- ZuMPT. F.. and W. M. Till. 1958. Notes on lection, 1948-1953. J. Egypt. Publ. Hlth. the classification and synonjmy of Gamasid Assoc. 31 (6): 199-272. mites parasitic on vertebrates (Acarina: Meso-

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