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inecticut
ibraries

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Part I, Argasidae]

[October, 1908

TICKS

A MONOGRAPH OF THE

IXODOIDEA

By
GEORGE H. F. NUTTALL

M.A., M.D., Ph.D., Sc.D, F.R.S.

Fellow of Magdalene College

Quick Professor of Biology in the University of Cambridge

CECIL WARBURTON

M.A., F.Z.S.
Christ's College, Zoologist to the Royal Agricultural Society

W. F. COOPER and L. E. ROBINSON

B.A., F.Z.S., F.L.S. A.R.C.Sc. (Lond.)

CAMBRIDGE
AT THE UNIVERSITY PRESS

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THE

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EDITED BY

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CONTENTS OF VOL. VIII, NO. 4 (October 1908)

{All rights reserved)

Boycott, A. E. and Damant, G. C. C. Experiments on the

Influence of Fatness on Susceptibility to Caisson Disease. 44.5

CuRRiE, J. K. Abnormal Reactions to Horse Serum in the

Serum Treatment of Cerebrospinal Fever. (Three Figures.) 457

DucKERiNG, G. Elmhirst. The Cause of Lead Poisoning in the

Tinning of Metals. (One Figure.) . . . . . 474

RuFFER, Marc Armand and Willmorb, J. Graham. The

Drinking Water of Steamships 504

Green, Alan B. Some Experiments on Immunity against

Vaccinia in Animals . . . . . . . .521

Green, Alan B. The Influence of Temperature, and some

other Physical Conditions, on Calf Vaccine. (One Figure.) 525

Watson, Herbert EdmEston. A Note on the Variation of the
Rate of Disinfection with Change in the Concentration of
the Disinfectant. (One Figure.) . . . . . .536

Wilson, W. James. Bacteriological Obseryations on Colon
Bacilli infecting the Urinary Tract, with special remarks
on certain Colon Bacilli of the " Anaerogenes " class . 543

Abstracts of Official Publications, etc. The Campaign

against Ankylostomiasis in Porto Rico .... 553

Publications received 556

TICKS

A MONOGRAPH OF THE

IXODOIDEA

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[All rights reserved]

QL
TICKS (^^

A MONOGRAPH OF THE lAfhC

IXODOIDEA

By
GEORGE H. F. NUTTALL

M.A., M.D., Ph.D., Sc.D., F.R.S.

Fellow of Magdalene College

Quick Professor of Biology in the University of Cambridge

CECIL WARBURTON

M.A., F.Z.S.
Christ's College, Zoologist to the Royal Agricultural Society

W. F. COOPER and L. E. ROBINSON

B.A., F.Z.S., F.L.S. A.R.C.Sc. (Lond.)

Cambridge

at the University Press

1908

im.

CambrtOgt:

PRINTED BY JOHN CLAY, M.A.
AT THE UNIVERSITY PRESS

\>

PREFATORY NOTE

THE discovery of the economic importance of ticks as carriers of
disease to man and domesticated animals has led to a vast increase
of our knowledge of this group. No existing work in any language
attempts to deal with the subject in a comprehensive manner, and the
student is confronted with a very extensive and widely scattered
literature from which he derives an impression of hopeless confusion.
There is therefore urgent need for a work of the nature here attempted.

The book will deal with the Classification, Structure and Biology of
Ticks, the study of the group having occupied the authors for several
years. Practically all that has been published on the subject has
received adequate consideration. The parts on Classification have
entailed much labour since it was found necessary to revise a large
amount of the work which has been done by others. The book will be
very fully illustrated by numerous text figures and plates, the majority
of which are original, the remainder reproduced from the best sources.

It was at first intended to publish a full account of the Ixodoidea
or Ticks as a complete vohime, but the increasing demand for a work
dealing with this group of parasites has caused us to decide to issue
without delay the part relating to the Argasidae. Other parts will
follow, and the whole, we hope, will be ready in about a year.

The parts will be complete in themselves but are designed to form
a volume of about 500 pages when all the parts have been published.
Each part will be issued in a stiff paper cover and will include a
bibliography printed on one side of thin paper so that the references
can be conveniently cut out and gummed on index cards. A complete
bibliography, including all the publications cited in each part, Avill
conclude the volume.

In the text the Harvard system of references is adopted, the year
and page of the authors' papers being added after their names. Unless
otherwise stated all the authors cited have been consulted in the
original. The completed volume will also contain an adequate intro-
duction, which we think it better to omit for the present.

Cambridge

August 1908

PAET I

THE ARGASIDAE

by

G. H. F. NUTTALL

and

C. WARBURTON

aided by
W. F. COOPER and L. E. ROBINSON

NOTE.

G. H. F. Nuttall and C. Warburton are responsible for the systematic
portion of this Part, while the biological section has been almost en-
tirely in the hands of the former. W. F. Cooper and L. E. Robinson
have assisted in collating the literature on the subject.

CONTENTS

OF

PART I
THE ARGASIDAE

SECTION I

DEALING WITH THE CLASSIFICATION OF THE ARGASIDAE

(Illustrated by 3 plates and 116 figures in the text.)

PAGE

The features which distinguish the Argasidae from the Ixodidae ... 1

Family characters, synonymy and literature relating to the Argasidae . . 2-3

The genus Argas : generic characters, synonymy and literature ... 4

List of species ............. 5

Terms and signs used in the descriptions 6

Key for the determination of species of Argas 8

Description, Synonymy, Iconography and Literature relating to the
following species :

Argas persicus ............. 8

reflexus ............. 22

hermanni ............ 27

cucwnerinus ............ 28

transgariepinus ........... 29

hrumpti 30

aequalis . ............ 33

vespertilionis ............ 34

transversa ............ 5

hrevipes ............. 5

The genus Ornithodoros : synonymy and literature, generic characters . 39

List of species 41

Key for the determination of species of Ornithodoros ..... 42

Vlll

Conteiits

Description, Synonymy, Iconography and Literature relating to the following
species :

Ornithodoros savignyi

„ mouhata

„ coriaceus

„ turicata

„ talaje .

„ pavimentosus

„ erraticus

„ tholozani

„ lahorensis

„ furcosus

„ megnini

„ canestrinii

•)■> papillipes

„ morbillosus

PAGE
42

46
55
57
59
62
63
65
67
70
71
78
79
80

SECTION II

DEALING WITH THE GENERAL BIOLOGY OF THE ARGASIDAE—

THE EFFECTS OF THEIR BITES,— THEIR RELATION TO THE
SPREAD OF DISEASE, ETC.

The genus Argas.

Argas persicus : Life history from the egg to the adult . . . . 81

Eflfects of bite on man and animals 85

Economic importance 87

In relation to disease 88

Methods of destroying 90

Argas reflexus : Life history 91

Eflfects of bite 92

In relation to disease .......... 94

Argas brumpti, eflfects of bite, habits, etc. ...... 95

The genus Ornithodoros.

Ornithodoros moubata : Life history ....... 96

Eflfects of bite, treatment, prevention 98

In relation to disease .......... 100

Ornithodoros coriaceus, biology, effects of bite, etc 102

„ turicata 102

„ talaje . 103

„ tholozani 103

„ megnini .......... 103

For what is known regarding other species of Argasidae in this respect see
Section I. The internal anatomy will be considered in the general introduction to
the completed volume.

LIST OF ILLUSTEATIONS

Plate I.

Fig.

1.

Fig-

2.

Fig.

3.

Fig.

4-5.

Plate II.

Fig.

1-2

Plate III.

Fig.

1-2

Argas reflexus ^, dorsum.
„ reflexus $ , venter.
„ persicus $, dorsum.
„ vespertilionis, last stage nymph, dorsum and

venter .to face p. 4

Ornithodoros savignyi 5 , dorsum and venter

42

talaje var. capensis, dorsum and venter

to face p. 60

IN THE TEXT

FIGURE

A and B. Schematic, to show differences between Argasidae and Ixodidae

1. Ornithodoros coriaceus $ with names of parts referred to in the

descriptions of Argasidae .......

2. Argas persicus ^ , the capitulum with names of parts referred

to in the descriptions of Argasidae

3-4. A. persicus 9 , dorsum and venter

5-8. A. persicus, details of integumental structure

9. A. persicus, the anus .....

10. ^. jjemcMs, the spiracle ....
11-12. A. persicus, the male and female genital orifices, etc. .
13-14. A. persicus, the capitulum and lateral view of the pali
15-18. A. persicus, the digits of the chelicerae, the hypostome, and

tarsus I seen in profile
19-23. A. persicus, representing stages of development from unfed larva

to second stage nymph ....

24-25. A. persicus, the larva, dorsum and venter .

26. A. persicus, tarsus and foot, and hypostome of larva

27-28. A. reflexus ^ , dorsum and venter
29-32. A. reflexus, a palpal hair, digits and hypostome .

33. A. reflexus, tarsi I and IV .

34-35. A. hermanni, digit and hypostome
36-37. A. transgariepinus 9 , dorsum and venter .
38-41. .4. /;;•;< ;»/;<t 5, dorsum and venter

PAGE

1

13

14
14
15
15

15

16

16

-17

17

-18

19

24

24

25

28

29

30

-31

List of Ilhistrations

FIGURE

42-43. A. hrumpti, capitulum of 5 , ventral and lateral aspects

44-47. A. hrumpti, digit, tarsi I and IV, details of integument

48. A. vespertilionis 5 j venter ......

49. A. vespertilionis, detail of peculiar structui'e on venter
50-51. A. vespertilionis, first and second stage nymphs .
52-56. A. vespertilionis, intestinal caeca, detail of integument, capitu

lum, hypostome, digit

57. A. vespertilio7ds, larval hypostome, digit, dorsal plate, anus

58. The tarsi I and IV of 9 species of Ornitkodoros compared
59-60. Ornithodo7-os savigiiyi, details of integument
61-63. 0. savignyi, palp, hypostome, digit ....
64-65. 0. saoignyi, tarsi I, III and IV, larva, dorsum and venter

66. 0. mouhata $ , dorsum and venter ....

67-68. 0. mo7ibata, part of ^ venter, cast skin of nymph

69. 0. moubata, capitulum of 9 , ventral aspect

70-71. 0. savignyi, legs I and IV .

72-73. 0. mouhata, legs I and IV .

74-75. 0. moubata, nymph, venter and capitulum

76-79. 0. moubata, egg, larva and details thereof

80. 0. moubata, capitulum and palp of larva

81-82. 0. coriaceus 9 , dorsum and venter

83-84, 0. coriaceus 5 , side view and venter .

85-87. 0. turicata ^ , dorsum and venter, digit

88. 0. talaje, hypostome and digit

89. 0. talaje var. capensis ^ , anterior part of venter
90-92. 0. pavimentosus, legs I and IV, and I and II
93-94. 0. tholozani, digits of $ and 5 • ...
95-97. 0. lahorensis 5 , dorsum and venter .

98-100. 0. lahorensis, digit, hypostome, tarsi I to IV

101. 0. ftircosus 5 , tarsi I, II and IV . . .

102-103. 0. me^'nMu cJ , dorsum and venter

104. 0. megnini, digits of nymph and 5 . . .

105-107. 0. megni7ii, young nymph, dorsum and venter .

108. 0. megnini, capitulum of nymph in ventral aspect

109-110. 0. megnini, larva, dorsum and venter .

111-112. 0. «?e^mm, leg of larva, gorged larva .

113. 0. canestrinii 5 , anterior part of venter, tarsus I

114. 0. papillipes $ , anterior part of venter, tarsus IV

PAGE

31
32
35
36
36

37
38
40
43
44
45
49
49
49
50
50
51
51
52
55
56

58-59
60
62
63
66

67-68
68
70
72
72

73-74
74
76
77
78
79

SECTION I

DEALING WITH THE CLASSIFICATION OF THE ARGASIDAE

Superfamily: IXODOIDEA Banks.

Family I : ARGASIDAE Canestrini, 1890.
Family II : IXODIDAE Murray, 1877.

The most striking difference between the Argasidae and the
Ixodidae is the possession by the latter of a shield or scutum, which
covers the whole body of the male, and prevents any large degree of
distention, while it forms a small patch on the anterior part of the
dorsum of the female, the rest of the body being greatly distensible.
The Argasidae possess no such scutum, but are covered by a more or
less uniform leathery integument. Many other differences, however,
exist, and the principal may be tabulated as follows :

Figures illustrating the main points wherein the Argasidcie (A, an Argas)
diiier from the Ixodidae (B, an Amblyomma ? ).

Argasidae

Ixodidae

Sexual dimorphism

Slight

Marked.

Capituhim :

Base

Ventral, in camerostome, no
porose areas in ?

Anterior, porose areas in ? .

Palps

Leg-like, with sub-equal ar-

Eelatively rigid, of very varied

ticles

form, 4th article rudimen-
tary.

Body :

Scutum

Absent

Present.

Festoons

Absent

Generally present.

Eyes (when present)

Lateral, on supra-coxal folds

Dorsal, on the sides of the
scutum.

Spiracles

Very small, more anterior

Generally large, well behind
coxa IV.

Legs : Coxae

Unarmed

Generally armed with spurs.

Tarsi

Without ventral spurs

Generally armed with 1 or 2
ventral spurs.

Pad (pulvillus)

Absent, or rudimentary

Always present.

N. I.

1

2 Family Argasidae

The differences extend to their habits as well as to their structure.
The Argasidae feed moderately, and the change of shape in both sexes
is chiefly a dorso-ventral thickening, while the females of the Ixodidae,
when gorged, are greatly increased in size, and their coxae widely
separated. The Argasidae live for several years, and as a rule lay
comparatively few eggs in small batches, have two or more nymphal
stages, and may moult after attaining maturity. The Ixodidae probably
do not live more than two years, lay a single huge batch of eggs, and
moult twice only, on leaving the larval and nymphal stages.

Family I. ARGASIDAE Canestrini, 1890.

Family Characters. Non-scutate ticks luith leathery integument
and slight sexual dimorphism, with capitulum inferior in the nymphs
and adidts, the palps being fy^ee and leg-like, with their articles sub-equal
in length. The spiracles are small and usually anterior to coxae iv.
Pulvilla absent or rudimentary, porose areas absent on the female
capitulum. The capitulum lies partly (larvae) or wholly (adults and
nymphs) in a hollow of the overhanging anteriorly protruding body.
When gorged the hollow is shallower.

The Argasidae, even when gorged, never increase much in size, and
in their flattened appearance when fasting they bear some general
resemblance to bed-bugs. Their principal hosts are man, birds and
bats. They are mostly inhabitants of warm climates or choose a
habitat in colder climates which assures them protection : thus Argas
reflexus lives in pigeon coops and fowl houses, Argas vespertilionis in
the retreats of bats. Ornithodoros moubata and savignyi inhabit native
dwellings, retreating into chinks in the walls or dusty floors. The
Argasidae appear to be chiefly nocturnal in their habits, like Cimex
lectidarius, and to prefer dryness.

The family Argasidae includes two genera, Argas and Ornithodoros.

N.B. There is some difference of opinion as to the number of
genera to be admitted under the family Argasidae. Pocock (1907,
p. 189) has advocated the revival of Latreille's genus Catns (or Carios)
for the aberrant form vespertilionis, which he considers clearly marked
off from other members of the family by the possession of a "conspicuous
transverse, slightly curved groove just behind the anus," and he would
also remove 0. talaje to another genus, for which he proposes the name
Alectorobius, on account of the movable lateral wings of the camero-
stome. Now the post-anal structure so conspicuous in vespertilionis,

Genus Argas 3

and which on examination appears to be a curious paired organ rather
than a simple groove, occurs also in other forms, notably in 0. talaje,
and is not peculiar to vespertilionis. With regard to 0. talaje, we
hesitate to remove it from Or-nithodoros, as in most respects it more
closely resembles 0. erraticus than that species resembles, for example,
0. savignyi.

In fact we are by no means sure that the family Argasidae contains
more than one genus, Argas^. At the time Ornithodoros was separated
off from it by Audouin (1827), the peculiar characteristics of 0. savignyi
seemed to warrant that proceeding, but the gap has largely been
bridged over by subsequently discovered species. For the present,
however, we prefer to retain the genus Ornithodoros, though some of
the distinctions which were supposed to exist between it and Argas
have to be discarded.

Synonymy and Literature :

Family ARGASIDAE Canestrini, 1890^.

Argasiden Koch, 1844, p. 219 ; 1847, p. 11.— Fam. Argantidae Agassiz, 1846, p. 32.
— Fam. Argasides Fiirstenberg, 1861, p. 208. — Subfiim. Argasidae Murray, 1877,
p. 180.— Berlese, 1885, p. 131 (iiicl. one genus : ^/-gros).— Riley, 1887, p. 744.—
Tribe Argasides Megnin, 1880, p. 132 (comprising but one genus : Argas). —
Laboulbfene and Megnin, 1882, p. 320 (both papers contain the same). — Megnin,
1892, p. 64 (brief mention only).— Tribe Argasin^S Railliet, 1886, p. 499.—
Neumann, 1888, p. 89 ; 1892a, p. 93.— Subfam. Argasinae Trouessart, 1892,
p. 38.— Railliet, 1893, p. 704.— Neumann, 1896, p. 2.— Dubreuilh and Beille,
1896, p. 68.— Ward, 1900 a, p. 196 ; 1900b, p. 436, and numerous recent
authors. — Trouessart, 1891, j). 308; misquotation for Argasidae. — Fam.
Argasini Canestrini and Fanzago, 1877, p. 192. — Canestrini, 1890, p. 530. —
Fam. Argasidae Canestrini, 1890, p. 530 ; 1892, p. 582.— Marx, 1892, p. 232.—
Trouessart, 1892, p. 22.— Salmon and Stiles, 1901, p. 400.— Pavesi, P., 1884,

1 In support of this view we may cite the case of Argas aequalis (Nn.) Neumann.
This species was first named Ornithodoros aequalis by Neumann (1901, p. 259), being
referred to the genus Ornithodoros because of its general form, parallel sides, and especially
the anterior portion of the body which narrows to a short rounded point. In Argus the
anterior portion of the body is usually broadly rounded and this had appeared to Neumann
to constitute a generic distinction until he came to describe A. brumpti which possesses the
physiognomy of Ornithodoros, together with essential characters of Argas. Neumann
(1908, p. 26) now refers aequalis to the genus Argas, because, on closer examination, it
has been found to possess the typical margin radially striated or with quadrangular areas,
visible ventrally and dorsally, which he now regards as a generic distinctiou. Neumann
appears to be almost in favour of degrading Ornithodoros to a subgenus, differing from
Argas by the ventral folds and the absence of the margin above mentioned.

- The above Synonymy of the Family is taken from Salmon and Stiles, 1901.
Additions have been made to date.

1—2

4 Family Argasidae

p. 485, used as a heading only to brief mention of A. erraticus = Ornithodoros
q.v. — Banks, 1904, p. 45, brief mention only. — Donitz, 1907, pp. 17, 24. —
Pocock, 1907, p. 189.— Nuttall, 1908, p. 389.— Banks, 1908, p. 14.— Tribe
Argasinae Neumann, 1892 b, p. 96. — Group Catastomata Marx, 1892, p. 233.
—Morgan, 1899, p. 136.— Subfam. Argasin^S Railliet, 1893, p. 715.

Genus 1. ARGAS^ Latreille, 1796.

Generic Characters. Body flattened, oval or rounded, with a
distinct flattened margin differing in structure from the general integu-
ment; this margin gives the body a sharp edge which is not entirely
obliterated even when the tick is fully fed. Capitulum (in adults and
nymphs) entirely invisible dorsally, distant in adults by about its mvn
length from the anterior border. On both dorsum and venter there are
numerous symmetrically arranged discs^, generally round or oval, more
or less disposed in radial lines. Elseiuhere the integument is minutely
wrinkled into irregular zig-zag folds. Eyes absent.

Type species : Argas persicus^.
Synonymy and Literature: Genus ARGAS Latreille, 1796*.

Argas Latreille, 1796, p. 178, obs. specimen 8 mm. long, had seen specimens from
Tuscany {A. rejlexus, doubtless) ; 1804, p. 53 : type, Acarus reflexiis ; 1806, p. 155 ;
1829, p. 288 ; 1835, p. 512.— Leach, 1815, p. 397.— Risso, 1826, p. 182.— Heyden,
1826, p. 612.— Audouin, 1832, p. 413.— Sundvall, 1833.— Dugfes, 1834 a, p. 15
(included in the family Oamasei by this author) ; 1834 c, p. 31. — Gervais, 1844,
p. 229 ; 1847, p. 351.— Koch, 1844, p. 219 ; 1847, p. 12.— Gervais and van
Beneden, 1859, p. 455.— Gerstaecker, 1860, p. 464 ; 1863, p. 344.— Furstenberg,
1861, p. 208.— Moquin-Tandon, 1861, p. 304.— Verrill, 1870, p. 116.— Spicer,
W. W., 1874 a, p. 185.— Taschenberg, E. L., 1874.— Megnin, 1876, p. 288; 1880,
pp. 133, 320.— Conil, 1877, p. 27 ; 1878, p. 101.— Murray, 1877, p. 180.—
Canestrini and Fanzago, 1877, p. 193. — Kiichenmeister and Zui5i,,1879, p. 538
et se^.— Berlese, 1885, p. 131.— Ludwig, 1886, p. 612.— Railliet, 1886, p. 499 ;

1 Often misspelt Argus, see Ant. Dug^s and Milne Edwards, pi. 27 ; FuUager, 1874 a ;
Francis, 1894, p. 452 ; Fuller, 1897, p. 590, etc. The derivation of ''Argas" attributed to
the Greek ap-y-qs ( = white, glistening) by Spicer (1874a, p. 185), who adds that being
eyeless, it can have nothing to do with him "of the hundred eyes." The numerous discs
on A. reflexus may well have been considered to represent "many eyes" by earlier
observers. We have never seen any specimens which could be described as white or
glistening. The white incrustations dotting the backs of A. reflexus found at Canterbury
in 1874 (noted by Gulliver) appears to have been derived from the plaster in the walls, or
it may have been excrement.

2 Scutella, foveolae, patellae, pits of other authors.

" A. reflexus (Fabr.) has been the hitherto accepted type of the genus (vide Nn. , 1896,
p. 2). We prefer to i&ke pcrsicus for our type because it is world-wide in its distribution
and commonly found, besides having been the better studied.

* Literature given by Salmon and Stiles, 1901, p. 401, revised and with additions.

TICKS

PLATE I

\^

M

Fig. 1.

Fig. 2.

■■■■

2JHH!5SS||

^^=

B

^E

FiR. 3.

Fig. 4.
Fig. 1. J. reflexus d , x 10.

Fig. 5.
Fig. 3. J . persicus <f , x 9.

Fig. 2. .-1. rejlexus ? , x 10.

(Wheler, I'JOG, figs. 6, 7. 39.)

Fig. 4. A. ves2)ertiUoiiis. Last uympbal stage, dorsal aspect, x 16. (Specimen from Cape Colony

Lounsbury Coll.)
Fig. 5. The same, ventral aspect. (Original, C. and K. phot.)

Grenus Ai^gas

i>

1893, p. 715.— Neumann, 1888, p. 89 ; 1892 a, p. 92 ; 1892 b, p. 96; 1896, p. 3,
etc., and 1908, p. 26.— Blanchard, 1890, p. 329.— Canestrini, 1890, p. 530; 1892,
p. 582.— Trouessart, 1891, p. 290; 1892, p. 35.— Efisio, 1892, p. 257.— Marx,
1892, p. 233.— Du Buysson, 1896.— Moniez, 1896, pp. 499-509.— Osborn, 1896,
p. 255.— Morgan, 1899, p. 137.— Ward, 1900a, p. 196; 1900b, p. 437.— Salmon
and Stiles, 1901, p. 401.— Banks, 1904, p. 45.— Aclogue, 1905, p. 536 (in-
accessible).—Dbnitz, 1907, p. 24.— Pocock, 1907, p. 189.— Nuttall, 1908, p. 390.
— Banks, 1908, p. 14, and many recent authors making casual mention.
(Not to be confused with Argas Seoul., 1835, crustacean ; Arges Hann., 1835,
crustacean ; Arges Goldf., 1839, crustacean ; Arges Val., 1840, fish ; Argus
Boh., 1761, mollusk ; Argus Scop., 1777, lepidopteron ; Argus Poli, 1795,
mollusk ; Argus Temm., 1815, bird; Argus Walk., 1837, arachnoid.)
Carios Latreille, 1796, p. 176 ; 1806, p. 161. — Kolenati, 1857, p. 16. —Canestrini,

1890, p. 480.— Neumann, 1896, p. 19.
Rhynchoprion Hermann, 1804, p. 69. — Heyden, 1826, p. 612. — Treviranus, 1831,
p. 188.— Latreille, 1829, p. 288.— Canestrini, 1890, p. 531.— Trouessart, 1891,
p. 290.— Railliet, 1893, p. 715.— Neumann, 1896, p. 3.

(Not to be confused with Rhynchoprion or Rhyncoprion Oken, 1815, dipteron ;
Rhynchoprium Marx, 1895, arachnoid.)
Caris Latreille, 1806, p. 161 ; type, vespertilionis { = Carios renamed for no obvious
reason); 1829, p. 290.— Risso, 1826, p. 182.— Heyden, 1826, p. 613.— Latreille,
1835, p. 511.— Gerstaecker, 1863, p. 343. (See also Audouin, 1832, p. 413.)—
Dugfes, 1834 c, p. 14 (regards it as Argas larva). — Kolenati, 1857, p. 15 ; 1858,
p. 4. — Murray, 1877, p. 181. — Canestrini and Fanzago, 1877, p. 184. — Haller,
1882, p. 312.— Canestrini, 1890, p. 480.— Trouessart, 1891, p. 290.— Neumann,
1896, p. 19.— Pocock, 1907, p. 189.
(Not to be confused with Caris Fisch., 1821, col.)
" Rhyachoprion {sic) Hermann," of Duges, 1834 c, p. 14.

In our opinion the genus Argas comprises six well established
species :

1.

2.

persicus
reflexus

3.

4.
5.
6.

transgariepinus
brumpti
aequalis
vespertilionis

There are four doubtful species :

7. cucumerinus (possibly = reflexus var. magnus) .

8. hermanni (near to reflexus var. magtius)

9. transversa Banks, 1902, p. 70, PI. II, Fig. 9. (Species

founded on one specimen ; description insufficient.)
10. brevipes Banks, 1908, p. 15, PI. I, Figs. 3, 4. (Descrip-
tion insufficient. Legs shorter than in A. persicus;
may be but a variety.)

8
22
29
30
33
34

28

27

6 Family Argasidae

The following species have been suppressed :

americanus, chinche, mauritianus, miniatus, radiatus, sanchezi =
A. persicus ; columbae, magnus, marginatus — A. reflexus (including
var. magnus) ; kochi = A. transgariepinus ; decussata, elUptica, fischeri,
inermis, longimcma, pipistrellae, pidchella = A. vespertilionis ; strogu-
loides Gervais, 1844, p. 231, is a purely nominal species (Nn., 1896,
p. 25). A.forskali Audouin, 1827, p. 430, figured as a hexapod larva in
Savigny, 1826, PI. IX, Fig. 13 (1 and 2 g.); mentioned in Duges, 1834,
p. 32 ; called Hyalomma Forskaelii (Aud.) by C. L. Koch, 1844, p. 222 ;
possibly if. aegyptium according to Nn., 1896, p. 24, and 1901, p. 255.

Explanation of terms and signs used in tlie following diagnoses.

L. = the extreme length of the tick.

W. = its width at the widest part.

{I. and 10. refer to length and width of various structures.)
Dorsum = the whole dorsal surface of the body.

Mammillae = imnu.te hemispherical or conical protuberances into which the
integument is generally raised.

Discs =i\ie foveolae, patellae, scutella, pits of other authors, see Figs. 6, 7, 23.
Venter = the whole ventral surface of the body.

spiracle = the " peritreme " or " stigmal plate " of some authors.

vulva = t\xQ genital orifice of the female.

grooves = \inei^r depressions. In Fig. 1 note pre-anal groove, post-anal groove
and post-anal median groove.

/o^o?s = prominent integumental ridges. In Fig. 1 note supra-coxal and coxal
folds.

earner ostome = the cavity in which the capitulum lies.

hood=t\\Q projection of the integument forming the walls of the camerostome.
Capitulum = the "rostrum," " head" or " false head " of various authors (Fig. 2).

basis capituli or shortly &ase = the basal ring, etc., of most authors.

h7/post077ie = th.e "maxiUa," "radula," "labium," or " Unterkiefer " of various
authors. The dentition is indicated by figures on either side of a vertical
line. Thus 3 | 3 means three longitudinal files of teeth on each half of the
hypostome.

chelicerae =t'he "mandibles" or " pseudo-chelicerae " of some authors. The
terminology usually applied to the portions of the digit of the chelicerae
appears to us wrong and confusing, and we prefer to recognise (1) an
" internal article," the latter bearing a " dorsal process " which is a portion
of it (not articulated), and (2) an "external article" which articulates with
the internal article upon its outer side.

1. internal article — " median apophysis " or " immovable finger " of some

authors.
dorsal jtwoce5s=" internal apophysis" of some authors.

2. external arft'c^e = " external apophysis" or "free" or "movable finger" of

some authors.

Genus Argas 7

Palps, the pedipalpi, 4-jointed, leg-like appendages on either side of the

hypostome.
(For details of the capitulum consult Fig. 2.)
Legs, six articles, coxa, trochanter, femur, tibia, protarsus and tarsus are always
recognisable, and there may be additional " false articulations." Articles
1 and 6 are generally referred to as the coxa and the tarsus respectively, the
others being indicated by their numbers. Certain dorsal prominences often
present on articles 5 and 6 are referred to as humps or dorsal protuberances.

---eye

Supracoocal fold
— genital orifice
eye

coxal fold
spiracle

median
postanal

groove

preanal grooue-

transuerse
postanal grooue

Fig. 1. Ventral aspect of Ornithodoros coriaceus i with names of parts
referred to in the descriptions (Nuttall, 1908, Fig. 2).

Palp article 4
Art. 3

-Internal \^f>^^f
-External \ofChelicera

. hypostome

7^ I ^^^-'- \-—\-St^afl or
A Ha TK, J\\ 11 .Chelicera

Fig. 2. Capitulum of Argas persiciis s with names of parts referred to
in the descriptions (Nuttall, 1908, Fig. 4).

8

(a)
(h)
(c)
(d)

(e)

Genus Argas

l^QY for the determination of species of Argas.
(Exclusive of doubtful species q.v. p. 5)

[Body circular ....

I „ oblong ....

[Margin striate ....
\ „ with quadrangular " cells "

[Body sub-conical in front
I „ rounded in front

[Dorsum with polygonal depressed areas

I „ without „ „ „

'Body scarcely narrower in front, capitu-
lum very small and posterior

Body narrower anteriorly, capitulum
large and more anterior

vespertilionis
b

persicus

d
e

hrumpti
aequalis

transgariepinus
reflexus

PAGE

34

30
33

29
22

ARGAS PERSICUS (Oken) 1818\

Synonymy. Owing to its wide geographical distribution (q.v.) due,
it appears to us, to the distribution of its chief host, the domesticated
fowl, this species has been given a great many specific names :

Rhynchoprion persicum Oken, 1818, p. 1567.

Argas persicus Fischer de Waldheim, 1823, p. 269.

Argas mauritianus Guerin-Meueville, 1829-43. — Megnin, 1880, p. 134 (only men-
tioned). Neumann, 1901, p. 256, examined a dried specimen collected by
Desjardins in Mauritius (Paris Mus.) and identified it as A. persicus.

Argas miniatus C. L. Koch, 1844, p. 219 ; 1847, p. 32 (PI. I, Fig. 4, legend states it
is a (J but text states ^ is unknown. Description insufiicient, specimen from
Demerara, Brit. Guiana. Original description translated in Neumann, 1896,
p. 24). Neumann, 1901, p. 255, examined the type. Name commonly used by
American writers and others, see var. miniatus.

Argas americanus Packard, 1872, p. 740, identified with A. miniatus by Neumann,
1901, p. 255. Synonymy explained by Salmon and Stiles (1901, p. 406) : —
Ercolani (1859) gave the name A. americanus Latreille to a form which is
parasitic in the ears of horses in San Domingo, and Gamgee (1871) did like-
wise, consequently the specific name atnericanus has to lapse. Again, Amhlyomma
americanum was placed under Rhynchoprion by Hermann (1804), but Rhyncho-
prion = Argas. For this reason Railliet (1893, p. 718) gave the tick the name
of Argas radiatus (see below). Acarus americanus L. {Acarus nigvM de Geer)

1 The priority for this species has hitherto been given to Fischer de Waldheim.

Argas 2)ei^sicus 9

placed under Rhynchoprion by Hermann was probably A mblyomma americanum
according to Neumann, 1901, p. 255.— Ward, 1900 b, p. 196.

Argas sanchezi Alf. Dug^s, 1891, p. 20 and Neumann, 1896, p. 16. Referred to A.
miniatus by Neumann, 1901, p. 255, after seeing the type. This name recurs
occasionally in the literature, vide Ward, 1900 b, p. 198. — Banks, 1901, p. 590 ;
1902, p. 568, etc.

Argas chinche Goudet, referred to A. miniatus by Neumann, 1901, p. 344 (no
reference).

Argas radiatus Railliet, 1893, p. 718. Neumann examined Koch's type of A.
miniatus and degraded radiatus.

Argas miniatus firmatus Neumann, 1896, p. 12, a variety based on 1 $ from Algiers.

Argas persicus var. miniatus Neumann, 1905, p. 240. (Neumann finds no constant
difference between A. miniatus and the type, and largely because of its distribu-
tion retains it as a variety. We see no reason for recognising the variety.)

Popular names : "Fowl tick" (America, Australia and Africa), "Adobe tick"' in
Arizona and New Mexico. In S. Africa "Wandluis" or "Tampan" (Lounsbury,
1900). In Persia " Miana bug," " Miane bug," " Mialleh de Mianeh," " Garib-
guez " or " Guerib-guez," the last name meaning " stranger bug " according to
Churchill, 1880, p. xxi (see also Tholozan, 1881, Megnin, 1880, p. 134) ; " M^ll^h "
(Dupre, 1819, p. 324), "mall^h" (Fischer de Waldheim, 1823, p. 272) = native
name; "punaise de Miana" (Kotzebue, 1819, p. 180), Schlimmer (1874)
states it is called "bhebguez" ("Shebgaz," according to Churchill, 1880, p. xxi,
meaning " biting at night ") at Chahroude and Bestham.

Iconography : Oken, 1818, PI. XIX, Figs. 1-4, notes resemblance to A. rejlexus.
He gives sketchy figures of A. persicus ; dorsal and ventral aspects. — Fischer
DE Waldheim, 1823, PI. I, Figs. 8-10, shows dorsal aspect with correct distri-
bution of the discs ; ventral aspect and capitulum not so good. — Savigny, 1826,
PI. IX, Fig. 8, dorsal aspect, very good considering the date when it was drawn.
— Gu^RiN-M^NEViLLE, 1829-43, PL VI, Fig. 3, of J. mauritianus dorsal aspect.
Fig. 3a part of capitulum. Both figures inaccurate. — Koch, 1847, PI. I, Fig. 4.
— KoLLAR- (date?) is stated by Heller, 1858, to have figured the capitulum. —
Heller, 1858, Pis. I-IV, deals chiefly with the internal anatomy. — Packard^,
1872, Fig. 68, referred to by Neumann, 1901, p. 255 {A. americanus). — Megnin,
1880, PI. VI, Fig. 3 (poor, A. mauritianus). — Laboulb^ne and Megnin, 1882,
Pl. XXI, Figs. 1-10 ; none of them accurate, except that of the capitulum,
which is fair. PI. XXIII, Figs. 1, 2, represent the larva and egg. — DuGfes^,
1891, Plate referred to by Neumann, 1901, p. 255 {A. sanchezi). — Marx in
Osborne, 1896, PI. Ill, Figs. 3-3 k. Drawings by Marx of A. americanus 9
and larva. — Neumann, 1896, p. 8, Figs. 4, 5, of digit (2 views) and tarsus I ;
Figs. 6, 7 {A. americanus), details of integument, 8* anus, 9* digit, 10* hypo-
storae, 11 tarsus IV; Figs. 16-21 {A. sanchezi), details of integument, digit,
hypostome, tarsus I, larval hypostome and ambulacrum. (Here reproduced,

' "Adobes" are sun-dried bricks used in building houses.

^ Inaccessible. Pohl and Kollar (1823) mention Argasidae.

^ These figures could not be studied, the originals being inaccessible.

* Figures marked by a star are reproduced in this work.

10 Genus Argas

except Figs. 5, 6, 9, 11, 16, 18 aud 20, which duplicate the others since the species
americanns and sanchezi no longer hold.) — Woods \ 1898, Figs. 20a, b, referred
to by Salmon and Stiles under bibliography (1901, p. 402) A. americanus. —
Hassall, 1899, Coloiu-ed Plate XVI {Argas americanus) of $ , dorsal and ventral
aspects : reproduced in Salmon and Stiles (1901). Text Fig. 16, larva (dorsum) ;
Fig. 17, ventral view of the larva (otherwise good), omits the coxae ; Figs. 18,
19, larval capitulum ; Figs. 20-22, larval ambulacrum (leg I), drawn like Figs.
16 and 17 from living specimens.— Ward, 1900a, Fig. 2; 1900b, Fig. 260, re-
ferred to inSalmon andStiles, 1901, p. 402.— Salmon and Stiles, 1901, A. miniatus,
Coloured PI. LXXVIII, 9 , reproduction of Hassall, 1899, PI. XVI. They give
a number of excellent text figures, of which those marked by a star are repro-
duced by us. Figs. 56, 57, views of chelicera complete. Figs. 71, 72* $ ; 73
portion of the marginal skin ; 74* tarsus I of adult ; 75, anus ; 77, 78*, larva
(dorsum and venter) ; 79, larval capitulum (quite as well shown in Fig. 78, but
both omit the two hairs on the basis capituli figured by Hassall) ; 80, larval
mandibles ; 81, larval Haller's organ.— Lounsbury, 1903, gives a photograph
illustrating A. persicus upon a sliver of wood torn from a fowl house ( x 2), and
a plate showing the 9 (ventral aspect), the larva (dorsal aspect) and a part of
the larval capitulum. He gives Wheler's photograph of the male (dorsum). —
Blanchard, reproduced in Neveu-Lemaire, 1904, p. 154, Figs. C and D, are very
inaccurate, and appear to be only poor copies from Savigny (1827). The same
figure appears in Perroncito (1882 and 1901) and is copied from Perroncito by
V. Linstow (1894).— Banks, 1904, Fig. 71, ? venter, poor.— Froggatt, 1906,
Figs. 1 and 2, showing Argas in crevices of splintered wood, and mouth-parts
of tick.— Wheler, 1906, Fig. 39* the S, dorsum.— Donitz, 1907, PL II, Figs.
9, 10$, 13, 14 larval leg. — Reaney, 1907, p. 401, two very bad figures of larva
and female.^MANSON, 1908, PI. IV, coloured figure of adult (0 dorsum. —
Nuttall, 1908, Fig. 4 ^ capitulum*.— Banks, 1908, PI. I, Figs. 1, 2, legs 1 and 4.

Literature. Papers marked " 0 " contain no original matter or
only make casual mention of ^. persicus.

1818. Oken, pp. 1567-1570.-1819. DuPRifi, p. 323, refers to efiects of bite.
Passage quoted in Fischer de Waldheim, 1823, p. 272.— Kotzebue, pp. 180,
194, rough description, fully quoted in Fischer de Waldheim, 1823, pp. 272-
274. Dwells on effect of bites on man.— 1821. Porter^. — 1823. Fischer de
Waldheim, pp. 269-273, gives brief description, dwells especially on effect of
bite.— 1828. Szovits^.- 1829. Latreille, b, p. 289. 0.-1829-43. Gu^rin-
Mi^NEViLLE, p. 17, only refers to plate and habitat {A. mauritianus). —
1833. CuviER, cited by George, 1876, p. 224, as referring to A. persiciis. —
1844. DuGfes, cited by Laboulbfene and Megnin, 1882. — Koch, C. L., p. 219.
A. miniatus, described in 18 words. — Walkenaer, p. 232. Also cited by
Heller, 1858, p. 297. Passage translated in Churchill, 1880a.— 1847. Koch,
C. L., p. 12. A. miniatus. 0.-1849. Lucas, H., p. 317. 0.-1855. Kuchen-

^ These figures could not be studied, the original being inaccessible.
* Figures marked by a star are reproduced in this work.
2 Cited in bibliography by Huber, 1899.

A. persiciis 11

MEISTER, p. 423. O.— 1858. Heller, C, pp. 297-326. Distinguished the sexes
and describes and figures the $ and $ genital orifice. Spiracle between legs 3,
4. Heller gives the first detailed study of the internal anatomy. — Kollar,
cited by Heller, 1858, p. 297, as giving first description and figure of capitulum.
—1862. HusEMANN, p. 248.— 1863. Gerstaecker, C. E. A., p. 344. 0.—
1867. Laboulb^ne, A., p. 54, obviously compiled from Fischer de Waldheim,
1823 iq.v.). — 1872. Packard, pp. 740, 741 {A. americana), cited by Salmon
and Stiles, 1901, p. 402.-1873. Fomouze, inaccessible i. — 1874. Schlimmer,
cited in Nuttall, 1899, p. 46. Notes occurrence in different parts of Persia and
records effects of bite. — Spicer, p. 209, quotes Walkenaer, 1844. — Taschenberg,
p. 171.-1875. Fritsch, p. 61 1.— 1877. Packard in Murray, p. 182 {A. ameri-
canus). — 1880. Churchill, S., p. xx, cites older authors and adds a few
observations of his own. — Ernodf, p. 256 1. — M^gnin, P., p. 134. More fully
dealt with in Laboulbfene and Megnin, 1882. — Goudet, J., cited by Megnin
(1880) as having described and figured "^. americamcs de Geer" and that it
looked like an Argas (see synonymy) = " chinche " or "nigua" in Colombia.
Megnin (1892, p. 67) thinks it must be 0. talaje. — 1881. Tholozan. See
Laboulbfene, 1881.— Laboulb^ne, A., (27, vii). Report from Tholozan. Cited
in full in Laboulbfene and Megnin, 1882, p. 327.— 1882. Bordier, p. 131, cited
by Nuttall, 1899, p. 47.— Cobbold, p. 77. — LABouLBfeNE and Megnin, p. 327,
deal at length with A. persicus, very diffuse, omit many essentials but describe
development and sexes. — Perroncito, p. 450. 0. — Tholozan, p. 15 1. — 1885.
MEGNIN, p. 460 {A. americanus)'^. — 1890. Blanchard, ii, p. 883 et seq. — 1892.
Marx, p. 234 {A. mnericanus)'^. — 1893. Hoehr, p. 348 {A. americamisY. —
Railliet, p. 718 {A. radiatus). — Riley and Howard, p. 267 {A. americana,
also A. americanus)^. — Webster, p. 149 {A. americanusY, inaccessible. — 1894.
Francis, p. 452 {Argus americanusy.—w. Linstow, p. 120. 0. — 1895. Braun,
p. 258. 0. — Howard, p. 41 7 {A. americanus) 2. — Lounsbury, p. 657.— Marx, p. 199
{A. amet-icafms)^.— Riley and Howard. — Packard, p. 417. — 1896. Dubreuilh
and Beille, p. 79. O. — Fuller, p. 593. General description of fowl tick. —
Neumann, pp. 9-12, 24, 25 (A. americanus). — Osborn, H., p. 256, quotes Packard's
description verbatim (A. americanus). — 1898. Woods, p. 102 {A. amei-icanus)'^.
— 1899. Hart, p. 180 {A. amei-icamis) in Trinidad. — Hassall, pp. 496-500 {A.
americanus). — Lounsbury, repr. A fuller account in his publication of 1903. —
Marlatt, C. L. (11, viii). O. — Morgan, p. 137 {A. americanus)'^. — Nuttall,
pp. 46, 47, cites older literature regarding effects of bite. — Taschenberg, 1900,
p. 738. 0.— Utility, p. 185, inaccessible^.— 1900. Ward, H. B., p. 197, quotes
Packard's description; mentions "J. sanchezV as another species. — Ward,
H. B., p. 437 {A. americanus). — 1901. Froggatt, p. 1349, inaccessible, cited
in Hunter and Hooker's Bibliogr. — Neumann, pj). 255, 256. — Perroncito,
p. 561. 0. — Salmon and Stiles, pp. 402-406, give a very full description {A.
miniatus), which we have utilized in part. — 1902. Brown, p. 86, cited by
Lounsbury, 1903, p. 6, re oviposition. — 1903. Lounsbury, pp. 1-15, gives a
very good description of the different stages and the biology. (Cited in text.) —
MARCHOUXand Salimbeni, p. 569. Regarding spirillosis in fowls. — 1904. Banks,

' Cited in bibliography by Huber, 1899.

' Cited in bibliography by Salmon and Stiles, 1901, p. 402.

12 Genus Argas

p. 331. 0. Described as "^. miniata." — Lounsbury (November. Repr., brief
mention), see Lounsbury, 1903. — Mosler and Peiper, p. 346. 0. — Neveu-
Lemaire, p. 154. 0. — 1905. Borrel and Marchoux, p. 362. Regarding
spirillosis in fowl. — Neumann, p. 240.— Nuttall, p. 22. 0. — Robertson, p. 561,
quoted by Hunter and Hooker, 1907.— 1906. Braun, p. 371. 0.— Froggatt,
pp. 14-18 ; gives general account of life historj' and habits ; measures for tick
destruction in infested fowl houses {A. americanus). — 1907. Donitz, W., p. 27. —
Hunter and Hooker, p. 71. Reaney, p. 401.— 1908. Hooker, a, p. 39. —
Manson, p. 204. 0. — Nuttall, p. 394 et seq. — Banks, p. 15.

(Not " Ai-gas americanus Latreille " of Ercolani, 1859, p. 248;
Gamgee, 1871, p. 219. Not " Argas americanus De Geer" of Megnin,
1880, p, 134. (Megnin thought it an Argas from a figure published by
J. Goudet; no reference.) Acarus americanus see Amhlyomma ameri-
canum. Dolly, 1894, p. 980. From Salmon and Stiles, 1901, p. 402.)

With the exception of a few unimportant papers all of the above
have been consulted in the original.

Specific Description.
(Refer to Figs. 2—26, PI. I, Fig. 3.)

Adults : — Body ovate or quite oval, widest towards the posterior end.
Margin (150-200 fx wide), composed of quadrangular units or " cells," each
enclosing a circular pit. Discs numerous, oval or round, arranged as shown
in figures 3, 4. Venter : anus nearly central, with smooth anal ring ;
spiracles small (125 /a) crescentic, equal to about half the width of the
anal ring; genital opening between coxae I and II \ Capitulum: base
ventrally with four long hairs directed forwards, two post-hypostomal,
one near the articulation of each palp (post-palpal). Palps about twice
as long as hypostome, second article longest, the others equal in length.
Chelicerae (see Figs. 15 a and h). Hypostome indented, 6 or 7 fine
denticles on each half distally, followed by stout teeth 2 | 2, the numbers
of teeth increasing to 3 | 3, 4 | 4, 5 | 5 basally, the teeth decreasing in size,
not attaining the external border nor extending beyond half the length of
the hypostome. Legs : sub-equal and similar ; coxa I distinctly separated
from coxa II in $ ; coxae II, III and IV contiguous; article 3 the
longest ; articles 3 and 4 broader distally ; tarsi with very slight dorsal
protuberance.

The above description applies to both sexes, which differ chiefly in

the slightly smaller size of the ^ — which is slightly narrower in front

(ovate, PI. I, Fig. 3) — and in the appearance of the genital orifice

(Figs. 11, 12). The latter is slit-like and broader than the capitulum

1 See Figs. 11, 12. In gravid ? it may be more anterior, as in Fig. 4.

A. persicus

13

immediately behind which it lies in the % (it is situate between
coxae II according to Laboulbene and Megnin, 1882, p. 333). In the
</ it is half-moon shaped and surrounded by an oval ring narrower
than the capitulum and placed further back.

The cT measures from 4 x 2"5 to 5 x 3 mm. (Nn. 1896) but it may
attain 8 x 5 mm. (Salmon and Stiles, 1901, p. 404). Fecundated ?s
measure 7x5 tolOxG mm. (Nn. 1896, p. 8) ; we have seen specimens
from Merced, California, measuring 10'4 x 6"3 mm. Gorged females may
however attain 11 x 8*5 mm. ; Lounsbury (1903, p. 5) and Donitz (1907,
p. 27) have seen specimens from Cape Colony 125 and 12'7 mm. long^

Fig. 3. Fig. 4.

Figs. 3, 4. A. persicus. Female, dorsal and ventral aspect. Drawn from mounted
specimen. Salmon and Stiles, 1901 (figs. 71 and 72, 17th Ann. Rep., B.A.I., U.S.
Dep. Agr.)2.

1 There is a considerable discrepancy in the measurements given by different authors,
due, no doubt, to the variation in size of A. persicus s and ? . Most authors do not
distinguish the sexes.

2 Lounsbury (1903, p. 5) notes that asymmetry is common, a fact we have also observed ;
these figures show such asymmetry.

14

Genus Argas

The integument, in this and all other species of Argas, is of a uniform
pale yellow colour, and the darker tints generally observable are
entirely due to the ingested blood. Thus the partially fed living
specimen appears of a leaden colour, with yellow margin, pale legs and
capitulum. Further feeding increases the darkened area and deepens
its hue to a blackish tint which may appear more or less red in preserved
specimens. The radiating series of discs (seen in Figs. 3, 4, 22, 23)
correspond to intervals between the digestive caeca, and serve for the
attachment of muscles.

Fig. 5.

FiK. 6.

Fig. 7.

Fig. 5. A. 2)ersicits. Portion of integumeut from posterior dorsal border of female,

X 44. Original N.
Fig. 6. A. persicus. A disc ("fossette") of medium size, x 110 (Nn. 1896, fig. 17:

A. sanchezi).
Fig. 7. A. persicus. Detail of integument showing a disc and irregular smooth areas bearing

hairs. From dorsum of adult, highly magnified. Original N. and W.

FiK. 8.

Fig. 9.

Fig. 8. A. persicus. Integumental hair, x 500. N.

Fig. 9. A. persicus. Anus, x 95 (Nn. 1896, fig. 8, of ^. americanus).

A. persicus

15

Nymphs resemble ^/s, but have no sexual orifice. There are two
nymphal stages {a and b) separated by a moulting. Nymphs (a, Figs.
21, 22) measure when full-fed, 4 to 45 mm. In second-stage nymphs
(6, Fig. 23) the discs are more marked, though not as distinct as in
adults ; when full-fed they measure 5'5 to 6'7 mm.

Fig. 10.

Fig. 11.

FiR. r^

Fig. 10. A. pernicits (Indian). Left spiracle of last stage nymph, oriented to the long
axis of the tick's body (compare with Fig. 4 of ? ). The crescentic cribriform plate
170 fjL long. Drawn from balsam specimen and opaque object with camera lucida.
Original N.

Figs. 11, 12. A. persicus 3 and ? genital orifices and coxae, etc. Specimens from
India. Original N. and W.

Fig. 13.

Fig. 13.
Fig. 14.

A. persicus <j capitulum, ventral aspect, x 71

A. persicus ? right palp in profile, external aspect, x 75

Fig. 14.
Indian specimen. Original N.

American specimen.

Original N.

16

Genus Argas

Fig. 15.

Fig. 16.

Fig. 17.

Fig. 15. A. persieus. Left digit A, dorsal and B external aspects, x 220 (Nn. 1896, fig. 4,

modified).
Figs. 16, 17. A. persieus. Hypostomes, x 135 (Nn. 1896, fig. 17 of A. sanchezi and

fig. 10 of ^. americanus respectively).

Fig. 18. A. persieus. Adult, tarsus I (^. miniaiMs). Salmon and Stiles, 1901
(fig. 74, 17th Ann. Eep., B.A.I., U.S. Dep. Agr.).

Fig. 19.

Fig. 20.

Fig. 21.

Fig. 22.

Figs. 19 — 22. J. ^^rsjCMS showing successive stages of development. Drawn from living
specimens, excepting Fig. 21. Specimens received from India and raised in Cambridge.
Fig. 19. Unfed larva 3 weeks after hatching. 1 mm. I. Fig. 20. Gorged larva
11 weeks after feeding (in winter). 2 mm. I. Fig. 21. Unfed nymph (Ist stage,
dead). 2-5 mm. I. Showing upturned margin as seen in hungering specimens when
alive, the intestinal caeca (dotted lines in Figs.) being practically empty. Fig. 22.
Nymph (1st stage) one month after gorging (in winter), note appearance of discs
omitted in previous figure. 3-3 mm. I. Original N.

A. persicus

17

Fig. 23. A. persicus. Nymph (2nd stage) 8 months after feeding, the discs more
numerous than in 1st stage nymph. Living specimen from India. 5 mm. I.
Original, N.

Fig. 24. A. jiersiciis. Larva, dorsal aspect. Salmon and Stiles, 1901
(figs. 77, 78, 17th Ann. Eep., B.A.I., U.S. Dep. Agr.).

18

Genus Argas

Larva : 680-770 fM in length (minus capitulum), hexapod and nearly
spherical (Figs. 19, 20, 24, 25). The larva becomes flattened and longer
as it matures, whilst unfed. The capitulum, though inserted ventrally,
projects well in front of the body, which is much wrinkled dorsally
and ventrally, but has no spiracles or discs. There is a circular or
oval unwrinkled area or plate in the middle of the dorsum. There
are numerous pennate hairs, the most prominent being a ventral row

Fig. 25. A. persicus. Larva, ventral aspect. Salmon and Stiles, 1901
(figs. 77, 78, 17th Ann. Kep., B.A.I. , U.S. Dep. Agr.).

persicus

19

near the posterior margin of the body (hypostome and palps see
Figs, 24-26). Legs (Figs. 24-26) long ; there is a distinct pad on the
tarsi.

Fig. 26.

A. persicus. A. Larval tarsus II ( x 300) ; B. larval hypostome ( x 270)
(Nn. 1896, fig. 21, A. sanchezi).

Egg almost spherical, yellowish brown,
apparently without external structure.

'6 to "8 mm. in diameter,

Hosts : In all parts of the world A. persicus appears to be pre-
eminently a fowl tick. It commonly attacks man in Persia, where, at
an early date, it had acquired a most evil reputation. In South Africa,
Lounsbury (1903, p. 11) states that it also seriously infests ducks, geese
and turkeys, and that he has on several occasions found a few adults in
pigeon-coops. But a few were attracted to feed on pigeons under
experimental conditions. It has been reported from Graaff-Reinet
as killing young ostriches and from Grahamstown as attacking canary
birds. A gentleman at Graaff-Reinet suffered once from a severe bite.
Neumann (1896) and Ward (1900 a) state that the tick has been found
on quail and Packard (1895) on turkeys in California, and according to
Salmon and Stiles (1901, p. 405), Packard found it once among
Boophilus annulatus which had been collected from cattle. The larvae
have been found on the wild dove (California) and on Zenaidura
macroura L. (Mexico). Riley and Howard (1893, p. 267) state that the
fowl tick attaches itself chiefly under the wing and shoulder of infested
hens.

2—2

20 Genus Ai^gas

Geographical distribution.

A. persicKS appears to be cosmopolitan, though its representatives
in different countries have been allowed specific rank by different
writers (see Synonymy, p. 8).

Europe: Russia. Neumann states that it was collected at Ssamjam
by A. Spoof (Ouderaans Coll.).

Asia: Persia. Oken (1818) and Dupre (1819) were the first to report
the presence of this tick in Persia. Kotzebue (1819, p. 180) states that
it has infested Miana from time immemorial, that it hibernates in the
cracks in the walls to become aggressive in summer. He reports passing
several villages which the natives told him had been abandoned owing to
A. persicus. At Avanlung it was only necessary to break away a part
of a house wall to discover hundreds of the ticks. The houses were
built of clay mixed with straw. It was first described by Oken (1818)
and by Fischer de Waldheim (1823, p. 269). Schlimmer (1874)
reported it from Chahroude and Bestham on the main road from Teheran
to Khoragan. Churchill (1880, p. xx) states that the natives of Persia
believe it lives on fowls in winter and that it occurs at Mazrah on the
Resht-Teheran road, at Chesna Ali, near Asterabad, at Shahrud in
Khorassan and at Kashan. According to Neumann it is distributed
throughout N.E. and N.W. Persia. Tholozan collected specimens in
chicken coops in different parts of Persia and sent them to Laboulbene
(1881). The Paris Museum contains specimens from Miana, of which
Neumann (1901, p. 253) gives particulars. The latter author also records
and gives particulars of specimens from Palestine, E. Turkestan, and
Pekin, China (all in Paris Mus.). Captain E. W. W. Greig, I.M.S., has
sent us speciniens from India (Kasauli, Punjab, 1906).

Africa: Egypt. Taschenberg (1874, p. 171) had specimens collected
in houses in Egypt. Neumann (1901, p. 253) records persicus from
Egypt (Klunzinger Coll.) and from Dongola, collected by Ehrenberg
(Berlin Mus.). Nuttall has received numerous live specimens from
Cairo collected in fowl houses by Prof H. Bitter, and others from the
Soudan collected similarly by Dr A. Balfour (Khartoum). A ^ was
collected at Fort National, Algeria, by P. Lesne (Nn., 1901, p. 253).
Lounsbury (ix. 1903, p. 1) states it occurs in most if not all the towns
of Cape Colony, also in the Transvaal, Orange River Colony and Natal
(at Pietermaritzburg). In Soutli Africa it appears to thrive as well on
the coast as inland, being as abundant about Cape Town, Port Elizabeth
and Uitenhage as at Grahamstown and Graaff-Reinet. Nuttall has

A. pei^sicus 21

received numerous live specimens from Mr Lounsbury and others have
been examined (from the same source) by Neumann (1901, p. 253).
Guerin-Meneville's (1829-43) specimens came from the Island of
Mauritius {A. mauritianus) where they infested fowls.

America : It (^4. miniatus) appears to be widely distributed, having
been found in N. and S. America and in the West Indies. In the United
States it has been frequently reported from Texas ' : F. G. Schanph (accord-
ing to Packard, 1895, p. 417) reported in November, 1884, that it caused
a large mortality in chickens in Dunant County. It is also reported by
Marx, 1892 a, Riley and Howard, 1893, p. 267, Hoehr, 1893, p. 348,
Osborn, 1896, p. 256, as destroying fowls in Kinney Co. and San Diego.
Hassall, 1900, p. 499, believed that it was extending northward in
Texas. Banks (1901, p. 590 and 1902 a, p. 568) reports "A. sanchezi"
from Catalina Springs, Arizo^ia (April), and from New Mexico (July).
Several authors report the ticks from California where they infested
a fowl house at Lakeside (Marx, 1892 a. Banks, 1904, p. 331). Packard,
1895, received specimens from Merced^ where they attacked chickens
and turkeys, and they have been found on the quail at Mariposa
according to Neumann (1896, p. 18). They injure fowls in Florida
according to Salmon and Stiles, 1901, p. 405. The tick occurs in
Mexico (A. sanchezi) at Guanajuato and in Southern California.
Numerous specimens have reached us from Brazil from Dr A. Lutz
(Sao Paulo) and through the courtesy of Drs Marchoux and Borrel,
of the Pasteur Institute, the ticks being infected with Spirochaetes
causing disease in fowls. "A. chinche" also occurs in temperate parts
of Colombia where it attacks man, and Koch's type of " miniatus "
came from Demerara {Brit. Guiana). In the West Indies the fowl tick
has been found at Trinidad where Hart- (1899, p. 180) supposes they
were imported from the United States, and Goodwin (Nn., 1901, p. 235)
also reports it as infesting chickens at Antigua.

Australia : Lounsbury (ix. 1903, p. 1) says the fowl tick occurs in
all five of the Australian Colonies, and no doubt he is correct for
W. Australia and Queensland. We find it recorded for the other colonies
as follows : Victoria : specimens found on fowls at Melbourne by
Desmond (Nn., 1901, p. 253; see also Brown, 1902). Neiu South

1 We have examined specimens collected in fowl houses in Texas in 1907 (sent by
"W. D. Hunter), and others off fowls in Merced, collected by Ehrhorn in 1894. Banks
(1908, p. 15) has seen specimens from Austin, Georgetown, El Paso, Patton, San Antonio,
Brackettville, and Colorado City, Texas ; also from Riverside, California.

^ Nuttall has received specimens in all stages collected by Mr Hart from Mr H. A.
Ballon (Barbados).

22 Genus Argas

Wales: specimens in the collection of the Department of Mines and
Agriculture were examined by Neumann (1901, p. 255 ; see also
Fuller, 1896, 1897, p. 590). S. Australia: Michael (1892, p. 202)
received live specimens from Adelaide, and Nuttall (1906) received live
specimens from the same place, sent by Dr E. A. Johnson. In all places
it infests fowls. According to Froggatt (1906, p. 14) it was introduced
from America into N. S. Wales and has since spread over all the south-
western towns.

Habitats : When A. persicus abandons its host, be it bird or man, it
retreats into the crevices in the walls of the chicken house or dwelling
to digest its food, undergo its metamorphosis or to oviposit, etc,
Schanph (published by Packard, 1895, p. 418) in Texas found them
retreating into the cracks and under the bark of trees upon which
chickens roosted,

N.B. For Seasonal Prevalence, General Biology, Effects of bite and
part played in Pathology, see Section II.

ARGAS REFLEXUS (Fabricius) Latreille, 1796.
Figs, 27—33, PI, I, Figs, 1, 2.

Synonymy: Acarus reflexus Fabricius, 1794, p. 426 ; 1805, ^. 353.
Acarus marginatus Fabricius, 1794, p. 427 ; 1805, p. 354.
Ixodes rejlexits Fabricius in Latreille, 1829 b, p. 288.
Argas reflexus (Fabricius) Latreille, 1796, p. 178 ; 1829b, p. 288.
Rhynchoprion columhae Hermann, 1804, p. 69, No. 1 ; also Fabricius, 1805,

p. 356.
Argas magnus Neumann, 1896, p. 14, degraded to A. reflexus var. magmcs

Neumann, 1905, p. 239.
Argus reflexus in Fullager, 1874 a, p. 86.
Common name: Pigeon tick, Taubenzecke (Ger.) ; Schellack (1908, p. 487) states

that the larva is known as the " Lederwanze " by pigeon fanciers in Germany.

Iconography: Latreille, 1796, PI. VI, Fig. 3.— Hermann, 1804, PI. IV (coloured),
Figs. 10, 11, iV and 0, dorsal and ventral aspects, capitulum and tarsus (very
crude figures). — Du6i:s, Ant., and Milne Edwards, PI. 27, Fig. 2 (poor). —
Gerstaecker, 1860, PI. XV, Figs. 1-6.— Pagenstecher, 1861, PL I.— Bianconi,
1867, figured the larva and its capitulum (part seen dorsally) and the capitulum of
the "adult" in ventral aspect. Figures sketchy. — Fullager, J., 1874a, p. 122,
Fig. 86 ; inaccurate figure of tarsi corrected in author's paper of later date 1874 b,
p. 234, Fig. 164. He also gives other figures. — M^gnin, 1880, Fig. 45, same as
following : 1892, p. 64, Figs. 5-7, ? dorsal and ventral aspects, capitulum (very
poor figures). — Taschenberg, 1880, p. 152, Fig. 39, small sketch, side view
(poor figure, reproduced by v. Linstow (?)). — Glaus, 1887, Fig. 444, sexual
organs, after Pagenstecher, 1861. — Berlese, 1888, fasc. XLVII, No. 5.

A. reflexus 23

cJ dorsum (coloured), venter, detail of dorsal and marginal integument. —
Canestrini, 1890, PI. XLI, Fig. 2, digit and hypostome (very inaccurate). —
Railliet, 1895, Figs. 487, 488, capitulum (inaccurate) ; gorged 9 dorsum and
venter (sketchy).— Dubreuilh and Beille, 1896, Fig. 6 (sketchy).— Neumanx,
1896, Figs. 1-3, of digit, hair ; tarsus 1 in two positions, tarsus 4, Bailer's organ.
Figs. 14, 15 {A. magnus), digit, hypostome ; here reproduced. — Taschenbero,
1900, p. 739, dorsum and venter of adult (?) (inaccurate).— Wheler, E. G., 1906,
Figs. 6, 7, $ dorsum, 5 venter, both good photographs ; here reproduced.

Literature : 1794. Fabricius, pp. 426, 427. Ac. reflexus, Ac. marginatus, Rhync.
cohimhae (few words).— 1804. Hermann, p. 69.-1805. Fabricius, pp. 353, 354,
356 (Latin). Same as 1784.— (No date) DuGfes, Ant., and Milne Edwards
(no date).— 1837. Walckenaer, hi, p. 231. Qi.— 1839. Raspail, p. 9.-1844.
Koch, C. L., p. 219, only mentioned.— 1859. Gervais and van Beneden, i,
p. 459. O.— 1860. Gerstaecker, pp. 457-470; efTects of bite.— Moquin-
Tandon, p. 281. — BoscHULTE, p. 554.-1862. Pagenstecher, p. 142.— 1863.
Gerstaecker, p. 344 ; brief mention only. — Coquebert, PI. 27, Fig. 10 (cited
thus by Bianconi, 1867).— Bocquillons (before 1867), p. 113 (cited thus by
Bianconi, 1867). 0.— 1867. Bianconi, C. A., pp. 107-112; the first to describe
larva.— 1871. Gulliver & Gulliver.— 1872. Gulliver, p. 205.— 1874. Ful-
LAGER, J., {a) p. 121 ; (6) p. 234.— " R. E.," p. 161 ; brief mention. -Spicer,
(a) p. 185. 0. (6) p. 209. 0.— Taschenberg, p. 171 et seq. (mostly citations
from other authors).— 1877. Canestrini, G., and Fanzago, p. 125. — 1879.
Boschulte, p. 562. — 1880. M^gnin, p. 135 ; repeated verbatim in Laboulbt;ne
and Megnin, 1882, p. 322.— Rivolta and Delprato, p. 314.— 1882. Laboulb^ne
and MjSgnin, pp. 332, 340. — Chatelin (see re effects of bite).— Contarini,
pp. 16, 161 (cited by Berlese, 1888).— Zurn, p. 78.-1885. Johannessen,
p. 347.-1888. Berlese, fasc. XLVII, No. 5. Short Latin description.— 1890.
Blanchard, II. p. 883 et seq. — Canestrini, p. 532, cites Bianconi's description
of larva. He describes $ and $ .—1892. Alt, p. 531.— Brandes, d. p. 10.—
Mi^GNiN, p. 64 ; brief mention.— 1893. Terrenzi, pp. 73, 79 ; reports reflexus
attacking persons at Narni. — 1894. v. Linstow, p. 121, cites Brehm, p. 688 (see
below). 0.— 1895. Braun, p. 258. 0.— Planchon, p. 29 (inaccessible).— 1896.
Dubreuilh and Beille, p. 76. O.— Neumann, p. 4.— Moniez, pp. 499-509. —
Gibert {thesis).— li^n. Brandes, p. 747.-1898. Ajutolo, p. 222; attacking
persons in Bologna.— Mingazzini, pp. 245-249.-1899. Wheler, p. 12.— 1900.
Brehm (see Taschenberg). — Taschenberg, p. 739 (in Brehm's Tierleben). —
Ward, H. B., (a). 0.— 1901. Neumann, p. 253.— Perroncito, p. 568.-1904.
MosLER and Peiper, p. 345. O.— 1906. Tonnel, p. 552.— Braun, p. 369. 0.—
1907. Donitz, p. 26.-1908. Schnee, p. 32.— Schellack, p. 486 ; transmis.sion
of spirochaetosis. — Banks, p. 16.

Adult: (Figs. 27, 28, PL I, Figs. 1, 2). Bodij oval, widest toward

the posterior end. Margin irregular lurinkled into radial striae forming

a border 200 fx wide, which is slightly turned up especially in unfed

specimens (whence reflexus) and is stated to always remain yellow

^ Papers marked 0 contain no original matter.

24

Genus Argas

(marginatus) ; discs arranged much as in A. persicus but the integument
much more finely wrinkled. Ventral surface : anus nearly central, with
smooth anal ring. Spiracles crescentic, elongated transversely, equal in
length to the width of the anal ring ; genital opening as in J., persicus.
Gapitulum: base with two long post-hypostomal hairs ventrally, directed
forwards. Palps with articles sub-equal, the third the shortest, denticu-
lated hairs (Fig. 29) dorsally. Chelicerae : digit, see Figs. 30, 31.
Hypostome (Fig. 32) rounded terminally, some small denticles at the
tip, followed by 2 I 2 stout teeth merging into 3 | 3 to 6 J 6 progressively

Fig. 27. Fig. 28.

Figs. 27, 28. Argas rejlexus, j' x 10 dorsum (with detail of margin) and venter.
From specimen lent by Prof. Neumann. Original, N. and W.

Fig. 29.

Fig. 29.
Fig. 30.
Fig. 31.
Fig. 32.

Fig. 30.

Fig. 31.

Fig. 32.

A. rejlexus. Palpal hair, x 250 (Nn. 1896, fig. 2).
A. rejlexus Y&r. magnus. Left digit, x 225 (Nn. 1896, fig. 14, modified).
A. rejlexus. Left digit, dorsal aspect, x 220 (Nn. 1896, fig. 1, modified).
A. rejlexus var. magnus. Hypostome, x 100 (Nn. 1896, fig. 15).

A. reflexus 25

smaller teeth. Legs as in ^. persicus, except that the coxae are
contiguous and the tarsi (Fig. 33) have a more distinct dorsal promi-
nence.

The sexes are distinguished by the same characters as in ^4. persicus.
The male (Figs. 27, 28) may attain 4 x 8 mm. </ pore between coxae
II, ? as in A. persicus (Megnin, 1880, p. 136; also Canestrini, 1890,
p. 532). The newly emerged female is about 5x3 mm., but when
gorged attains 8x4 mm. or even 9x5 mm.

V V \ V K-

Fig. 33. A. reflexus. Tarsi I and IV of large nymph. Sketch. W.

Nymphs measure 4x2 mm. (1st stage) or 5"5 x 4'2 to 7'3 x 4'8
(2nd stage), they resemble the male, but the larger ones are without a
true sexual aperture.

Larvae almost spherical, L. 0*5 mm., at most 2 mm. (Megnin,
1880) with terminal or subterminal capitulum. Hypostome with
dentition 2 | 2. Colour testaceous.

Eggs ovoid, reddish, '5 mm. I.

The foregoing description is taken in part from Megnin, 1880,
Laboulbene and Megnin, 1882, and Neumann, 1896, pp. 4, 5 (con-
densed) ; the measurements of the nymphs were made on specimens
received by us from Germany in 1908.

ARGAS REFLEXUS var. MAGNUS Nn. 1905.

Figs. 30, 32 (see Synonymy above).

This variety only differs from the type in its large size, its relative
narrowness, and the fact that the posterior border of the body is more
lanceolate. The details of the digit appear also to differ (compare
Figs. 30 and 31). Origin : Equador and E. Patagonia (Gulf of
St George).

A. REFLEXUS.

Hosts : Pigeons, the larvae have been found once on the horse
(Starcovici), and the nymphs and adults may attack fowls and man
occasionally.

26 Grenus Ai^gas

The Geographical Distribution of this species appears to be
limited to Europe and N. Africa. Osborn (cited by Ward, 1900) has
stated that it occurs in the United States as far north as St Louis but
there is no certainty that the determination was correct,

England: A. rejiexus was found at Canterbury Cathedral by
Gulliver (1872, p. 205) who states that its presence had "long been
known in the dark recesses of our time-honoured fane " and that it was
regarded there as an " insect peculiar to Canterbury Cathedral," The
ticks were found " rather plentifully crawling about the inside of the
base of the Cathedral " (see also Fullager, 1874 a and b ; " R. E.," 1874 ;
Spicer, 1874, Taschenberg, 1874, Wheler, 1899, p. 12), Two specimens
were secured alive in the Cathedral by W. F. Cooper in March, 1908.
We find no other records of its presence in Great Britain although
Dcinitz (1907, p. 26) states that it used to prevail in Ireland (authority
not stated).

France: the pigeon tick, according to Neumann, occurs especially
in the Ardennes. Latreille (cited by Megnin, 1880, p. 136) once found it
in a house, but judging from the statements both of Latreille and
Hermann it has grown much rarer in France. Laboulbene and Megnin
(1882, p. 323) sought in vain for years to procure specimens from pigeon
fanciers, and Neumann could only lend us a single </ specimen for
purposes of study.

Germany: Pagenstecher (1862) records its presence in large
numbers in the loft of a house at Frankfort where it greatly disturbed
the sleep of the servants at night. Gerstaecker (1863, p. 344) notes its
having been recently found to attack man in Germany, and states
it occurs more commonly in Southern Europe. Taschenberg (1880,
p. 152) states it has long been known in Alsace and in the Province
of Saxony at Eisleben (Taschenberg, 1873; 1900, p. 740) having been
recorded there and in Westphalia and at Friedeberg a. d. Saale before
1871. Donitz (1907, p. 26) writes that it is difficult to obtain. Schnee
(1908, p. 32) records that about 1896 an old church was torn down at
Magdeburg and that persons visiting the church at the time were much
attacked. As is usual in such cases, pigeons had been kept there and
the ticks wandered into the church. Mertens in 1907 sent specimens to
Hamburg from Magdeburg and some of these were kindly placed at our
disposal by Dr Fulleborn (Institute of Tropical Medicine, Hamburg).
Schellack (1908, p. 487) found it in two pigeon coops in Magdeburg.

Italy: Bianconi (1867) reports it from Modena. Canestrini (1890)
states it was found in 1876 in the interstices of the mosaic in the church

A. reflexiis 27

of San Marco, Venice. Berlese (1888) states that it is found about
Venice and Florence. Laboulbene and Megnin (1882) obtained
specimens from Pisa, and according to Perroncito (1878 and 1901) it
prevailed to such a degree at Turin as to make it impossible to raise
young pigeons. Birds 1 to 2 weeks old were attacked and died of
exhaustion. Terrenzi (1893) reports it as attacking persons at Narni
and as present at Tivoli, and Ajutolo (1898) as attacking persons at
Bologna. At Narni, according to Terrenzi, they wandered out of a
pigeon coop which had been disused for years and gained entrance to an
adjoining house where they inflicted troublesome bites upon the inhabi-
tants at night.

Russia and Roumania : R. Blanchard records it from Odessa and
the larvae have been found on the horse in Roumania by Starcovici
(cited by Neumann, 1896, p. 5).

Africa : according to Neumann (1901, p. 253) Simon found a large
specimen at Tlemen, Algeria.

It is obvious from the foregoing that this species is not common.
One of us (N.) tried in vain for years to obtain specimens from
England, France, Italy, and Germany but it was only recently that
some were received from Magdeburg. Owing to the growing interest
in relation to ticks as disease-carriers it is however quite possible that
a more careful search in the future will lead to A. reflexus being found
to be more prevalent in Europe than has hitherto been supposed.

Habitat: A. reflexus lives chiefly in pigeon-coops where it hides in
cracks in the walls and woodwork, the adults and nymphs attacking the
birds at night.

N.B. For General Biology, Effects of bite and part played in
Pathology, see Section II.

ARGAS HERMANNI Audouin, 1827.

Figs. 34, 35.

Literature, etc. : Savigny, 1826, PI. IX, Fig. 7 (resembles A. reflexus). — Audouin,
1827.— Neumann, 1896, p. 13, Figs. 12, 13, here produced ; 1901, p. 253.

Adults : Body oval, almost always markedly convex. Integument
of uniform appearance, very finely wrinkled. Numerous small discs
arranged as in A. reflexus. Margin finely striate, 75 — 80 /x. wide.
Venter: spiracles about two-thirds (150 /u.) the width of the anal ring.
Anal aperture bordered by a few short hairs. Capituluni relatively

28

Genus Argas

small (275-300 /x I.) ; digit (see Fig. 34) ; hypostome (Fig. 35) tapering
and slightly rounded at the tip : two long post-hypostomal hairs on the
basis capituli. Females attain 8x5 mm., and the males are only slightly
smaller.

Fig. 34. Fig. 35.

Argas hermanni.

Fig. 84. Left digit, x 225 (after Nn. 1896, fig. 12, modified).
Fig. 85. Hypostome, x 135 (Nn. 1896, fig. 13).

Nymphs show great variation in size, ranging from 1"5 x 1 mm. up
to the size of the male.

Neumann identified a tick collected by Rafifray (E. Simon's Coll.)
in N. Abyssinia as belonging to this species on collating it with the
numerous specimens, of unknown origin, in the Paris Museum.
Subsequently, three specimens were brought from Egypt by Klunzinger
(Neumann, 1901, p. 253). Neumann separates this species from reflexus
chiefly on account of " the fineness of the texture of its integument,
and the minute size and details of its capitulum," We regard it as
a doubtful species.

ARGAS CUCUMERINUS Neumann, 1901.

Lit. : Neumann, 1901, p. 254.

Male : Body flat, long oval, 10 x 5 mm., almost as broad in front as
behind. Integument rather finely wrinkled. Margin striate, rather
narrow. Venter: anus almost circular, central, well behind the spiracles,
which are less than the anal width. Capitulum small and anterior ;
hypostome 2 | 2. Legs long ; coxa I slightly separated from coxa II ;

A. cuciimerinus

29

the space between the coxae of the two sides equal to ^ the body-width.
Tarsi with a thick dorsal protuberance distally.

Description based on two specimens found beneath rocks in dry
places at Lima, Peru, by Gaudichaud (Paris Museum).

This species is doubtfully distinct from A. reflexus. Neumann
thought it might be the male of A. magnus, which he has since fused
with A. reflexus (see p. 22).

ARGAS TRANSGARIEPINUS White, 1846.

Figs. 36 and 37.

Lit., Icon, and Syn. : Argas transgariepinus White, 1846, p. 363, PI. II, Fig. 4,
dorsal and ventral aspects (Figs, and description useless).
Argas kochi Neumann, 1901, p. 254.

Kesembling A. reflexus, but differing from it as follows :

Adults : Body short oval, and almost as broad in front as behind.
Margin striate like that of reflexus, but relatively broad (3 mm.).
Spiracles scarcely as long as the width of the anus. Sexual orifice of
the male very small, facing the posterior extremity of coxa I. Capitulum
very small ("8 mm.) and very posterior, far from coxae I, lying in a
clearly marked camerostome. Hypostome narrow, palps short. Coxa I
far from coxa II, and the space between the coxae on the two sides
very broad— equal to one-third the width of the venter. Tarsi I much
humped distally, the protuberances on the other tarsi slight.

Fig. 36. Fig. 37.

Figs. 36, 37. Argas transgariepinus ? , x 4. Dorsum aud venter. Sketch from type
specimen in Brit. Mus. Original, N.

30

Genus Ai^gas

N.B. These points are taken from Neumann's description of
A. kochi, based on a single (^ from Basutoland, collected by Cristol
(Paris Museum). White's description is insufficient, but Neumann
found the types (2 females) in the British Museum and they agreed
with A. kochi, except in the genital orifice (a sexual character) and the
fact that the tarsal protuberance was almost as well marked on tarsi
2, 3 and 4 as on tarsus I. If the examination of other specimens of
A. transgariepinus should show similar tarsi in both sexes, Neumann's
species would become A. transgariepinus var. kochi. (There are
3 females (dry specimens) in the British Museum collection. The
accompanying figure was drawn from one of these measuring 10x7 mm.,
a smaller specimen measured 8"5 x 6 mm.)

Fig. 38. Fig. 39.

Figs. 38, 39. Argas hruvipti ? , x 3. Dorsum and venter. Original, N. and W.

ARGAS BRUMPTI Neumann, 1907.

Figs. 38 to 47.

Lit. and Icon.: Neumann, 1907, pp. 224-229, Figs. 9-14, here reproduced.

Adults and Nymphs : Body flat, only a few nymphs and males
showing the dorsum convex through repletion. The adult $ (Figs. 38-41)
attains 20 x 13 mm., average size 15 x 10 mm., nymph or young </
measure 7x5 mm. Sides of the body parallel, its anterior border
conical, recalling Ornithodoros ; posterior border broadly rounded; margin
(Fig. 47) finely striate. Dorsum pitted by symmetrically arranged
polygonal depressed areas, bounded by rugose ridges, and containing

A. brimipti

31

small discs, as shown in Figs. 40, 46. Numerous short hairs, especially
along anterior border. Venter : the anterior conical portion is hollowed
out into a deep triangular camerostome bounded by lips divergent
behind, and connected behind the capitulum by a transverse fold, im-
mediately behind which is the vulva in the ? , the (/ genital orifice

Fig. 41.
Argas brumpti ? .
Figs. 40, 41. Dorsal and ventral aspects. Nn. 1907, fig. 9.

Fig. 42. Fig- 43.

Argas brumpti ? .
Fig. 42. Capitulum and anterior part of venter, x 28. Nn. 1907, fig. 12.
Fig. 43. Capitulum in profile, x 28. Nn. 1907, fig. 12.

32

Genus Argas

being somewhat more posterior. Spiracles crescentic, less than the
width of the anal ring, which is oval. Hairs rather more abundant than
on the dorsum. Capitulum (Figs. 42, 43) : with very thick base ; palps
tapering, the first two articles (especially art. 2) long, the 3rd and
4th short ; four strong hairs on the outer surface of 1st article.
Chelicerae: see digit. Fig. 44; hypostome (Fig. 42) long, narrow,
lanceolate, bearing two long hairs at its base; dentition 2 | 2, 14 — 15 teeth
per file, the inner teeth widely separated. Legs yellowish, relatively

Fig. 44. Fig. 45.

Fig. 44. Digit, x 210. Nn. 1907, fig. 13,

Fig. 45. Tarsi I and IV, x 14. Nn. 1907, fig. 14.

Fig. 46. Fig. 47.

Argas brumpti.

Fig. 46. Detail of dorsal integument, x 11.
Fig. 47. Marginal integument, x 25.
(Neumann, 1907, figs. 11 and 10.)

A. hi'imipti 33

short and strong; coxa I (Figs. 39, 41) separated from coxa II by an
interval equal to their breadth ; coxae II, III and IV contiguous ; all
coxae have their anterior and posterior borders covered by striated
integument ; tarsi (Fig. 45) with a spiniform protuberance which gives
them a bifid appearance.

Origin : Description (condensed from Neumann) based on 35
specimens collected by Dr Brumpt at Ogaden, Somaliland.

Note: Dr Brumpt having kindly sent us a specimen of A. hrumpti %
(Dakhatto Valley, Somaliland), we have been able to compare it with
Neumann's description. We have figured our specimen (Figs. 38, 39).

Effects of Bite, etc. See Section II.

ARGAS AEQUALIS (Nn.) Neumann, 1908.

Syn. and Lit. ; Ornithodorus aeqxuxlis Neumann, 1901, p. 259.
Argas aequalis (Nn.) Neumann, 1908, p. 26.

Nymph: Body flat, 5 x 2-5 mm., with lateral borders parallel, rounded
posteriorly, tapering anteriorly to a short rounded point ; colour
reddish, capitulum and legs yellowish white. Integument folded, with
very fine granulations, covered with whitish hairs, excepting on the
venter between coxae. A distinct striated margin visible dorsally and
ventrally ; a dorsal protuberance corresponding to the capitulum ;
symmetrical depressions occupied by discs. Venter very prominent
centrally. Sexual orifice punctiform, between coxae I. No eyes.
Capitulum twice as long (1 mm., including base) as broad, the base twice
as broad as long, resting on a broad transverse fold equal to twice its
breadth ; hypostome long, narrow, pointed, lanceolate, with numerous
teeth anteriorly, followed by dentition 2 j 2, not extending to the middle
of its length. Legs fairly long ; coxa I stout, thick, somewhat separated
from coxa II ; tarsi without protuberance.

The description (modified from Neumann) is based on a specimen
(late-stage nymph) collected at Utengala, German East Africa, by
Fiilleborn (Berlin Mus.). This species is unfortunately not figured.
(See p. 3, footnote 1.)

34 Genus Argas

ARGAS VESPERTILIONIS (Latreille), 1796.

Figs. 48-57, PL I, Figs. 4 and 5.

Synonymy and Literature : Carios vespertilionis Latreille, 1796, p. 176, larvae.

Caris vespertilionis Latreille, 1804, p. 161, larvae. Generic name changed for
no apparent reason. Latreille, 1829 b, p. 290, notes the "peau ecailleuse"
as a generic character, see also Latreille, 1835, p. 511.

Acarus fischeri in Savigny, 1826 (see Iconogi-aphy).

Argas fischeri AuDOUiN, 1827, p. 427, probably mature nymph.

Argas pipistrellae Audouin, 1832, p. 412, larvae.

Argus pipistrellae in Ant. Dug^s and Milne Edwards (no date).

Caris vespertilionis Gervais, 1844, p. 227, larvae.

Caris elliptica Kolenati, 1857, p. 16, probably nymphs.

Caris longimaiia Kolenati, 1857, p. 16, probably nymphs.

Caris decussata Kolenati, 1857, p. 16, probably nymphs.

Caris inermis Kolenati, 1857, p. 16, probably nymphs.

Argas fischeri G'EORG'E., 1876, pp. 223-225, in Westwood, 1877, p. Ixii, referring to
George's paper. Probably nymphs i.

Argas piilchella George, 1876, p. 224. The author proposed this name tentatively
as he thought the " Blyborough tick " might be A. fischeri. He described
the mechanism of the foot and notes the spiracles between legs III and IV.

Argas pipistrellae Westwood, 1877, p. Ixii.

Argas vespertilionis (Latreille) Neumann, 1896, pp. 20-23 ; 1901, p. 254.

Caris vespertilionis Latreille, revived by Pocock, 1907, p. 189.

Iconography^; Savigny, 1826, PI. IX, Fig. 6 (1, 1', 2, 2') much better figures than
most of those published since ; large nymphs or possibly adults. — Audouin,
1832, PI. XIV, Fig. 1.— Ant. DuGfes and Milne Edwards, PI. XXVII, Fig. 3
(larva, poor).— Gervais, 1844, PI. XXXIV, Fig. 8, larva.— George, 1876, Pis.
XXI, XXII, figures of nymph : venter, part of capitulum, tarsus, dorsum,
intestinal caeca*. — Neumann, 1896, Figs. 22-26*: nymph, ventral surface
and detail thereof ; capitulum ; hypostome, digit ; larval hypostome, digit,
dorsal plate, anus. — Wheler, 1899, p. 12, Fig. 4, described as nymph ;
reprinted 1906, Fig. 8*, described as adult after balsam specimen of the
"Blyborough tick" (see George); also (1906) figures part of capitulum and
marginal integument.

1 Westwood (1877, p. Ixii), who is quoted in Neumann's synonymy (1901, p. 254),
refers incidentally to George's papers (1876, 1877) in his Presidential Address, stating
with regard to the name given by George: "It is given as Argas fischerii of Walcken.,
but it seems to me identical with A. pipistrellae, Aud."

2 Figures marked * are reproduced herein.

A. vespertilioms 35

There is no extant description of the adult of this species. Wheler
(1899, p. 12) described a specimen at the British Museum which he
believed to be adult, but we have examined it and find it to be a
nymph, probably in the second stage. Lounsbury has kindly lent us
a mature $ of the form, recognised by Neumann as at least only a
variety of A. vespertilionis, which infests penguins at Queenstown,
Cape Colony. The following description refers to this specimen :

Female : L. (hood not included) 7'4, W. 8"8 mm., broad oval, much
depressed, irregularly convex in the middle of the dorsum, red-brown
with narrow reflexed margin of reddish-yellow colour, the integument
entirely covered with very fine, conical, sharp granulations, except on
the discs, which are arranged as shown in the figure (Fig. 48). The

Fig. 48. Argas vespertilionis ? , L. 7'4 mm. Venter (see text). Original, N. and ^Y.

anterior margin projects in the form of a triangle on a level with the
dorsum, and from beneath this projection a rounded, yellow convex hood
proceeds forward and downward, visible dorsally, and concealing the
capitulum. Venter : anus at about the middle of the body ; supra-coxal
fold well-marked, short, extending fi-om the sides of the hood to the
posterior ends of coxae IV, Coxal folds well-marked, narrow ; very
numerous, small, radially arranged discs. Vulva: a very broad slit
between the posterior ends of coxae I, much wider than the basis
capituli and reaching the coxae on either side. Behind the anus, at a
distance of about three times its diameter, there is a remarkable paired
organ (see Fig. 49) consisting of two narrow, deep, crescentic clefts, on
each side of the middle line ; each cleft lies in an area of modified
iutegument, free from mammillae ; behind the cleft the area is finely
punctate, while in front, on its concave border, it is finely striate
at right angles to the direction of the cleft. (This structure is also
observable in the nymphs.) Capitulum very salient ventrally, with
very long base and extremely small appendages ; hypostome narrow,

3—2

36

Genus Argas

indented at the tip, with small inconspicuous teeth, apparently all
marginal. Palps : article 1 comparatively large and massive, the other
articles very small and narrow. Legs: coxae sub-conical, coxa I
separated from coxa II ; coxae II, III and IV contiguous ; tarsi abruptly
tapering, almost humped.

The descriptions of the nymph and larva given below are condensed
from Neumann (1896, pp. 22-24).

Fig. 49. Argas vespertilionis. One of two curious grooved organs lying posterior to
anus as indicated in key-figure above and highly magnified below. Present in adults
and nymphs. Drawn from large nymph (opaque object) and only indicated by a
line in Fig. 48 of ? . Original, N. and W.

Argas vespertilionis.
Fig. 50. Nymph, ventral aspect, x 28 (Nn. 1896, fig. 22).
Fig. 51. Nymph, ventral aspect, x 13 (Wheler, 1906, fig. 8).
We regard these as Ist and 2nd stage nymphs.

A. vesper tilionis

37

Nymph (Figs. 50-52) : Body nearly circular, 2*4 x 2'1 mm.
(3-7 X 3'8 mm. to 4*95 x 4*37 mm. according to Wheler, 1899, p. 12,
? 2nd stage nymph. We have examined large nymphs from the Cape,
measuring 5"5 mm. in length). Integument finely creased in zig-zags in
young individuals, in polygonal design in the more aged ; margin
(Fig. 53) with irregular folds in young individuals, quadrangular areas

Fig. 52. Argas vesper tilionis. Nymph's intestinal caeca.
After George, 1875, pi. II, fig. 2.

JS^

a-^Xci,

Fig. 53.

Fig. 54.

Fig. 55.

Fig. 56.

Argas vespertilionis. Nymph.
Fig. 53. Slightly schematic figure of part of ventral surface and margin, x 300.
Fig. 54. Capitulum, ventral aspect, x 65.
Fig. 55. Hypostome, x585.
Fig. 56. Digit, x 665.

(Neumann, 1896, figs. 23, 24, 25 a and b.)

in others, radiating, having very short hairs ; numerous small slight
depressions dorsally and ventrally, arranged similarly to the discs in
A. persicus. Venter: spiracles reniform (34 /a w.), less than half the
width of the anal ring (130 x 90 fx). Capitulum (Fig. 54) anterior, the
three distal articles of the palp together with the tips of the chelicerae
and hypostome visible dorsally (in younger nymphs) ; hypostome (Fig.
55) slightly indented, 2 | 2 rows of 5 or 6 teeth ; digit, see Fig. 56 ;

38

Genus Ai^gas

palps short, thick, with articles scarcely longer than broad, excepting
the first ; each article bears 4 short denticulated hairs dorsally ; two long
post-hypostomal hairs, two shorter post-palpal hairs more posterior.
Legs thick sub-equal ; coxae contiguous ; tarsi tapering, tarsus I alone
showing a slight dorsal protuberance. In some of the large 2nd stage
nymphs we have examined there existed an imperfect sexual orifice.

Fig. 57. Argas vespertiliotiis. Larva: A. hypostome, x575; B. digit, x865; C. dorsal
plate, X 110; D. anus, x 370 (Nn. 1896, fig. 26).

Larva (Figs. 57 A-D): Body short oval, 1'3 X|l mm. or 2 x 1"5 mm.
Some ten radiating furrows on the posterior half of the dorsum.
Integument with fine transverse parallel folds except on an oval
squamous shield anterior to the middle of the dorsum. Three pairs
of hairs on the anterior part of the dorsum. Twenty marginal hairs.
Radiating series of discs between the intestinal caeca. Capitidum,
almost entirely visible dorsally. Hypostome narrow, long, pointed ;
dentition 4 | 4. Palps slender, article 1 nearly as wide as long, articles 2,
3, and 4 twice as long as wide ; article 4 small. Hairs on the basis
capituli as in the nymph, but short. Legs slender, sub-equal ; coxae
nearly contiguous, elongated, a broad interval between the coxae of the
two sides.

Neumann's description of the nymph is based on three specimens,
one in the possession of M. Julien (Paris), and two specimens of the
" Blyborough tick " from R. Blanchard. The ticks were taken from
Vesperugo pipistrellus.

A. vespei'tilionis 39

It is probably Argas fischeri Audouin. {Carts elliptica Kol. was found
on Myotus murinus ; Caris longimana Kol. on Miniopterus schreibersi
and RJiinolophus clivosus ; Caris decussata Kol. on Myotus murinus and
Rhinolophns Jiippocrepis ; Caris inermis Kol. on BracJiyotus dasycnemus
and Synotus barbastelliis.)

Neumann's description of the larva is based on the examination of
four specimens ; two collected in Paris by E. Simon, from Vesperugo
pipistrellus, one taken by Dr Buysson from Plecotus auritus at Bronet-
Vernet (Allier), and one from Vesperugo kuhli at Chegga (R. Blan-
chard's Coll.). Latreille's Caris was taken from Vesperugo noctula.
Argas pipistrellae Audouin, and Caris vespertilionis Gervais, were
taken from V. pipist7'eUus. Argas fischeri Audouin, collected in Egypt,
is only known through Savigny's figure. There are larvae of A. vesper-
tilionis in the British Museum ; Mr E. G. Wheler's collection contains
larvae and nymphs collected at Puttenham (1903), and we have also
obtained larvae taken from V. pipistrellus from Histon, Cambridgeshire,
11 May, 1905 and 29-30 July, 1908.

Genus 2. ORNITHODOROS C. L. Koch, 1844.

Synon. and Lit.: Omithodoros Koch, 1844, p. 219; 1847, p. 11. — Fiirstenberg, 1861,
p. 208. — Canestrini and Fanzago, 1877, p. 194. — Canestrini, 1890, p. 531 ; 1892,
p. 582. — Trouessart, 1892, p. 47. — Neumann, 1896, p. 3 et seq. — Lounsbiiry,
1899, p. 240.— Ward, 1900 a, p. 196 ; 1900 b, p. 437.— Salmon and Stiles, 1901,
p. 407.— Neumann, 1901, p. 256.— Banks, 1904, p. 45.— Donitz, 1906, p. 145.—
Pocock, 1907, p. 189.— Banks, 1908, p. 16.

Argas {Omithodoros) Murray, 1877, p. 183.

Omithodoros Karsch, 1878, p. 321.— Marx, 1892, p. 233; 1895, p. 199.— Osborn,
1896, p. 255.— Morgan, 1899, p. 137.— Neumann, 1908, pp. 17, 26.

Rhynchopriwn Marx, 1895, p. 199.— Osborn, 1896, p. 255.— Neumann, 1896, p. 42
{Rhynchoprium a misprint).

Omithodorous Koch : Morgan, 1899, p. 136 {sic).

Alectorohius Pocock, 1907, p. 189 (see Omithodoros talaje).

Ornithodorxis of numerous writers since Koch.

Generic Characters. Body fiat luhen unfed, but usually becoming
very convex on distention. Anterior end more or less pointed and hood-
like. Margin thick and not clearly defined, similar in structure to the
rest of the integument, and generally disappearing on distention.
Capitulum sub-terminal, its anterior portions often visible dorsally in the

40

Genus Ornitlwdoros

adult. Discs- present or absent ; hut when present, not arranged radially

Urtam fairly constant grooves and folds on the venter, namely a coxal

fold internal to the coxae, a supra-coxal fold external to the coxae, a

TARSI I AND IV OF SPECIES OF ORNITHODOROS

savignyi
nwubata

savigrtyi

^..^^zzf:^

nwubata

turicata

talaje

talaje

laAorensis

iakor

megnjru

megnini

^iCZ]

furXOSLLS

Fig. 58.

J

farcosus
Original, N. and W.

Structures analogous to the " discs " of Argas are probably to be found in all species
of Ormthodoros under sufficient magnification, but in some species they are not visible as
such to the naked eye, or by the aid of a simple lens. In such cases the species is said in
the subsequent diagnoses to be without obvious discs.

Grenus Ornithodoros

41

transverse 'pre-anal and a transverse 'post-anal groove or furrow, and a
post-anal median groove'^. Eyes present or absent.

Type species Ornithodoros savignyi (Audouin).

N.B. 0. savignyi differed so markedly from the species of Argas
known when Koch established the genus Ornithodoros that the diagnoses
of the two genera presented no difficulties. More recently described
forms have, however, tended to narrow the gulf between them. Argas
brumpti, for example, has some features more characteristic of Ornitho-
doros, while Ornithodoros tholozani in some respects approaches Argas.

The genus Ornithodoros includes 11 well established species.

1. savignyi .

2. moubata

3. coriaceus

4. turicata .

5. talaje

6. pavimentosus

7. erraticus

8. tholozani

9. lahorensis

10. furcosus .

11. megnini ,

Doubtful species are :

12. canestrinii

13. papillipes

14. morbillosus ,

42

46
55
57
59
62
63
65
67
70
71

78
79
80

Suppressed are :

americana = turicata

miliaris = erraticus

coniceps = talaje var. coniceps

rudis = talaje

reticulatus Gervais, 1849-51, in de Gay, Zoologie, p. 44,
PL V, Fig. 8. Latin description in two lines
and figure. Species purely nominal (Nn.,
1896, p. 41).

1 Refer to Fig. 1, p. 7.

42

Genus Ornitliodoros

(0)
id)

(e)

(9)
(70

(i)
U)

Key for the determination of species of Ornithodoros.

(Exclusive of doubtful species q.v.

[Integument mammillated or granular
( „ not mammillated

(with eyes ....
(without eyes

(Integument pitted, no eyes
[ „ finely wrinkled

(with flat contiguous granulations
(with hemispherical granulations

(with body broad and rounded in front
I with body sub-conical in front .

, [with eyes of equal size

(with anterior pair of eyes larger

[with obvious discs ^ ....
(without obvious discs ^

[with movable cheeks to camerostome
(without movable „ „

l^coxae II larger than I ; with tarsi forked
' distally .....

I coxae II smaller than I ; with 3 humps on
y tarsi I — III .....

(tarsi slightly humped

{tarsi II — IV with prominent dorsal humps

p. 41.)

b.
c.

d.
e.

megnini, p. 71.
lahorensis, p. 67.

pavimentosiLS, p. 62.
/.
moubata, p. 46.

9-

savignyi, p. 42.

coriaceus, p. 55,

h.
i.

talaje, p. 59.

furcosus, p. 70.

turicata, p. 57.

erraticus, p. 63.
tholozani, p. 64.

ORNITHODOROS SAVIGNYI (Audouin), 1827.

PI. II and Figs. 58, 59-65, 70, 71.

Synon.: Argas savignyi Kwdiowm, 1827, p. 18.3.
Ornithodoros savignyi C. L. Koch, 1844.
? Ornithodoros morbillosus Gerstaecker, 1873 (see p. 80).
Argas schinzii Berlese, 1889.
Native names : see 0. moubata, p. 46, with which it must be included by natives

in the places where both species occur together, as will be seen by reference

to their geographical distribution (see pp. 46, 52).

1 See footnote, p. 40.

TICKS

PLATE II

Fig. 1. O. savUjmji ?. Dorsal aspect, x 8. Specimen from S. Africa.
Original, C. and R. phot.

Fig. 2. O. savignyi ? . Ventral aspect, x 8. (Same specimen as in preceding figure.)

0. savignyi 43

Lit. and Icon.: Savigny (1826, PI. IX, excellent figures of dorsum, venter, legs;
less good of capitulum and mouthparts), rei)roduced by Walckenaer and
Gervais (1837-47, PI. XXXI, Fig. 2).— Guorin-Menoville, 1829-43, PI. VI, Fig. 4,
mouthparts taken from Savigny without acknowledgement. — C. L. Koch, 1844,
p. 2. — L. Koch, 1875, p. 1, notes (only) three specimens from vicinity of Anseba
river.— Murray, 1877, p. 182.— Megnin, 1880, p. 134 (nothing original).—
Laboulbfene and Megnin, 1882, p. 324 (nothing original).— Berlese, 1889,
PI. VII.— Neumann, 1896, pp. 27-29, Figs. 27-32 ; here reproduced.— Pocock,
1900, p. 49.— Neumann, 1901, p. 256.— Lounsbury, 1899.— Braun, 1906, p. 372.
0. — Christophers, 1907, PI. I-V, two text figures, deals especially with internal
anatomy. Figure of larva here reproduced. — Donitz, 1906, p. 145, PI. Figs. 3
and 6 (here reproduced).

Adult : Body (PL II, Figs. 1 and 2) short oval, broadly rounded at
both ends and often slightly narrowed on a level with coxae III and IV.
Integument thick, and covered by distinct mammillae (100 to 200 /u, in
diameter, PL II and Fig. 59), except along certain tracts which appear as
furrows in unfed individuals, and are still recognisable by the absence
of mammillae in gorged specimens ; their arrangement is best seen
from the figure. They are transversely wrinkled and present no
obvious discs. Numerous short hairs between the mammillae, more

^

Fig. 59. Fig. 60.

Fig. 59. 0. savignyi. Granulations of integument, x 65 (Nn. 1896, fig. 27).
Fig. 60. O. savignyi. A "fossette" or disc, x 235 (Nn. 1896, fig. 28).

numerous on the venter. Venter (PL II, Fig. 2) : supra-coxal folds
well-marked, and bearing four eyes of equal size, one pair facing coxae I
and the other the intervals between coxae III and IV^ ; coxal folds
slight ; pre-anal groove well-marked ; behind it, three pairs of
longitudinal furrows, especially noticeable in large specimens ; a short
median depression in the form of an inverted Y ; anus wider than long
(400 X 450 /x). Spiracles above the supra-coxal folds, 600 /x wide,

1 See Figs. 1 and 83 of 0. coriaceus where position of eyes is similar.

44

Grenus Ornitliodoros

crescentic. Gapitulum stout, free; palps (Fig. 61) tapering, articles
1 and 2 equal in length ; article 3 the shortest ; hypostome (Fig. 62)
short with dentition 3 j 3 as regards the principal teeth, the external
teeth the stoutest; more numerous smaller teeth posteriorly; Chelicerae:
see digit (Fig. 63). Legs (Figs. 58, 64, 70 and 71) stout, the last
pair one and a half times the length of the first ; coxae contiguous
and diminishing from I to IV ; the protarsus (article 5) of leg I has
three large dorsal humps, the middle the largest, and the tarsus three
sub-equal humps equidistant ; the tarsus of leg IV has three humps.

Fig. 61.

Fig. 62.

Fig. 63.

Fig. 61. 0. savignyi. Palp, the articles measuring from base to tip 400 — 400 — 250 —

310 M (Nn. 1896, fig. 31).
Fig. 62. 0. savignyi. Hypostome, x 65 (Nn. 1896, fig. 30).
Fig. 63. 0. savignyi. Digit, 200 fi I. (Nn. 1896, fig. 29).

the two proximal near together, an interval three times as great
separating the second and third. The (f and $ resemble each other
except in the generally smaller size of the (^ and of the cT genital
orifice ^ Many long blunt hairs thickly beset the anterior margin
of the body giving it a downy appearance.

1 See Figs. 66, 67 of 0. rnoiibata.

O. savignyi

45

Nymphs : There are at least two nymphal stages, if not more. In
the nymphal stage preceding maturity, there is a great resemblance
to the adult and there is a rudiment of the sexual aperture.
Smaller nymphs, belonging to an earlier stage, are also encountered
which show no trace of a sexual aperture.

Fig. 64. 0. savignyi. Legs IV (A), III (23), I (C), x 17 (Nn. 1896, fig. 32).

Fig. 65. O. savignyi. Larva recently emerged from the egg, dorsum and venter.
After Christophers (1906, p. 51, about 1 mm. I., same size as the egg).

Larvae (Fig. 65) : These issue from the egg as in other Ixodoidea,
0. savignyi differing from 0. moubata (q.v.) in this respect, according to
Christophers (1906, p. 51), in India. This author informs us (1908) that

46 Genus Ornithodoros

the larva does not feed and does not increase in size before it moults,
but unlike that of 0. moubata it appears able to throw off the eggshell
before turning into a nymph.

Hosts : This species attacks man and Lounsbury (1899 ; he may
have included 0. inouhata in the statement) reports that it feeds on
various animals : fowl, dog, horse, goat, cattle, pig. We have fed it on
fowls and rabbits in Cambridge. It may be noted that it feeds like
0. moubata but for the fact, observed in one instance, that it inserts its
palps into the wound it inflicts.

Geographical distribution : Recorded from Nubia (at Bularli, Pell
Coll.), W. Somaliland, Abyssinia (Courbon Coll., Paris Mus.), German
East Africa, S. W. Africa (Nn. 1901, p. 256), and the Congo (Pocock,
1900, p. 49, and 1907). Its distribution in Africa overlaps with that of
0. moubata. Lounsbury has sent us specimens from Cape Colony, and
he records it (1899) as occurring in Rhodesia, Bechuanaland, Portuguese
E. Africa and Transvaal. Neumann records in detail (1896, p. 29) the
specimens found in different parts of Africa, possibly some of them
represent 0. moubata which at that time was not recognised as another
species. Neumann and Gerstaecker record it from Egypt (where it was
first found by Audouin) and Upper Nubia. Neumann has examined
specimens from Southern India, where its presence has been recorded
by Christophers (1906, p. 2). We have received specimens from Aden
and Egypt through the courtesy of Dr C. W. Daniels.

ORNITHODOROS MOUBATA (Murray), 1877.

Figs. 58, 66-80.

Synon. : Argas mouhata Murray, 1877, p. 182.

Ixodes mouhata, Cobbold, 1882, p. 78.

Ornithodoros savignyi var. caeca Neumann, 1901, p. 256 (referred to as var. caecus
or caecum by various authors).

Native names'^: bibo, Toro name in Uganda (Christy, 1903, p. 187).— Suaheli terms
are pasi (plural papasi) and Kupe' (We\\ma.n MS.). — gourat in Nubia and Soudan ;
mechger in Abyssinia ; cotirdoud in Galla ; oldirrho in Kilima-ndjaro (Brumpt,
1901, p. b18).—papazi in Mombassa (Karsch, 1878, p. 311, who calls it
0. savignyi). — tampan or tampCw on Lower Zambesi (Livingstone, 1857, p. 628 ;
Pocock, 1903 and 1907 ; Wellman, 1907, the last-named author informs us that
this name is employed by the Portuguese E. African colonists). — kufu or mhu'^

1 Without doubt some of these names include 0. savignyi.

^ Also applied to mosquito and to malaria, according to B. Koch, in parts of E. Africa.

O. moiibata 47

in Tote (misspelt "bu" in Mauson, 1903, p. 7l4).—carrapaio^ in Portuguese East
Africa (a loose term), Livingstone (1857, p. 382) spelt it carapato, often misspelt
"■arrapato. Wellman (in MS.) states that kxirapatti and garapata are Arabic and
Bantu corruptions of the Portuguese word. — tapazi ai Nyangwe, Lualaba river,
Livingstone (1874, il, p. 115). — hifundikala about Leopoldville Congo (Duttou
and Todd, 1905 h).—bimp^isi is the Batdk^ name (Button and Todd, 1905 b).—
mouyata at Popokabaka (Button and Todd, 1905 b). — moxibata in Angola
(Murray, 1877, p. 182). According to Wellman (in MS.) Dihata (singular) and
Mabata (plural) is the Kimbundu name for the tick, so that the name moubata
given by Welwitsch as the native name and adopted by Murray is incorrect, a
rarer Kimbundu name is Mbanze. — ochihopio (pi. ovihopio), native Umbundu
name in Angola, W. Africa (Wellman, 1906, p. 154). — liwuno^ is the Lovale
name in E. Angola and N. W. Rhodesia (Wellman, 1908, personal communica-
tion).— kvnpztto, common name in Eastern Congo Free State (Button and Todd,
1905 b).

Icon. : 1877. Murray, Fig. on p. 182 (wretched).— 1903. Christy, coloured PI. XV
and photograph, 4 figs. — 1905. Newstead, Pis. I and II. some figs, here
reproduced.— Button and Todd, PI. Ill, Figs. 2-4; PI. IV, Figs. 1-3, photo-
graphs of 0. moubata feeding, in a dish, profile of tick showing line of cleavage
of skin when moulting, details of internal anatomy. — 1906. Bonitz, plate
showing tarsi I and IV* — Wellman, p. 155, Fig. 2 ; dor.sal and ventral aspects
(very poor); 1906-07, Fig. 1.— Guiart and Grimbert, Fig. 410, sketchy.— 1907.
Pocock, text Fig. Ill, $ ; dorsal and ventral aspects. — 1908. Manson, PI. IV,
Fig. 3 ; dorsum of adult (?). Nuttall, vii, 1908, Fig. 1*

Lit.: 1857. Livingstone, pp. 382, 628.— 1874. Livingstone, Vol. ii, pp. 33, 115.—
1877. Murray, p. 182, gave a quite useless description. The species was
established by Pocock, 1900, who examined the types (Brit. Mus. collected by
Welwitsch in Angola).— 1882. Cobbold, p. 78. 0.— 1895. Bowson, p. 1201,
sent specimens from Tete to British Museum ; describes eflfect of bite, fever,
etc. — 1896. Neumann, p. 30.— 1900. Pocock, p. 222, first described some of
the specific diffei-ences between moubata and savignyi. — 1901. Neumann,
p. 256.— Brumpt, p. 578.— 1903. Christy, p. 187.— Manson, p. 713.— Pocock,
p. 188, determined specimens collected by Christy in Uganda, and states that
Bowson had also sent specimens of 0. moubata from Tete, Zambesi. — 1905.
Caiman, p. 124, determined specimens sent by Wellman from Angola. — Button
and Todd (b), pp. 123-127. — Feldmann, p. 64, Filaria perstans, supposed host
being 0. moubata. Author states it is an Argas, and it might be from his figure.
His description appears to apply partly to moiibata and partly to one of the
Ixodidae. — Kerr, p. 126, criticism of Feldmanu's observations.— Newstead,
pp. 1696, 1697.— Wellman, repr.— 1906. Bonitz, pp. 145, 148.— Werner, p. 776.
—Wellman, p. 155 (also 1906-7).— 1907. Pocock, pp. 194-196.— Mollers,
p. 278.-1908. Manson, p. 203.— Nuttall, p. 388.

1 Used in Europe to denote the castor-oil plant {lUcinus communis) according to
Wellman (ms.).

- Also applied to Sarcopsylla penetrans according to Wellman (ms.).

48 Genus Ornithodoros

This species closely resembles 0. savignyi, from which, however, it may
easily he distinguished by the absence of eyes, and the details of its leg
structure {compare Figs. 70-73). The humps on the protarsus of leg I are
sub-equal and toothlike, while the tarsus of leg IV is shorter and thicker
than in 0. savignyi, and its humps are nearly equidistant, the intervals
(proximal and distal) between them being as 4 — 5\ Less hairy than
0. savignyi'^.

Adults usually measure 8x6 or 7 mm. (Christy, 1903, p. 187 ;
Pocock, 1907, p. 195, and others), but the females, especially when
gorged, may attain I. 11 mm. (Dcinitz, 1906 ; Wellman, 1906), or
even 12 x 10 mm. when fully grown and filled with eggs and blood
(Button and Todd)l The colour varies fi-om dusty brown to greenish
brown in living specimens with occasional dull ochreus patches ; after
gorging it appears darker and turns reddish or blackish brown in
alcohol. Reference to the figures (Figs. 66-69) will give a better idea
of the appearance of the tick than a lengthy description. From the
observations of Newstead, Dutton and Todd, and of Donitz (1906,
p. 146 ; 1907, pp. 11, 20) and R. Koch, whom he cites, there appears to be
no doubt but that adults (8 — 9 mm. I.) may moult. Donitz states he
has observed females moult after ovipositing. Newstead, Dutton and
Todd (1907) raised a female from the egg and kept her under
observation for 25 months during which time she moulted six times.
Other specimens moulted 6 — 9 times during a year. Under favourable
conditions they may moult 2 — 3 times in two months. Mollers (1907,
p. 278) states that the nymphs moult after each feed of blood and that
they feed 6 — 7 times before reaching the adult stage, after which only
the males continue to moult. The females lay batches of eggs after
each feed but do not continue to moult. (This is contrary to what
Donitz states.) It is obvious, therefore, that there are several nymphal
stages, the number not having been clearly established. The largest
nymphs may equal adults in size, and we have noted that they show
a punctiform mark where the sexual orifice is situated in the adult,

1 A point established by Donitz, 1906. According to Newstead (1905, p. 1696) the
species can be further distinguished by the dorsal process of the digit being bidentate in
0. moubata, but this is not of practical importance.

" According to Mollers (1907, p. 278) ticks descended from the same parents may differ
considerably in size. The males are usually smaller than the females but the size is very
variable in both sexes. Nymphs of the same age which have fed 4 to 5 times may measure
3 — 4 or 8 — 9 mm. long. (This variability in size, also noticeable in other Ixodoidea, doubt-
less depends upon the amount of blood consumed.)

0. movhata

49

Fig. 66. 0. moubata ? , x 5. Dorsum and venter; specimen from
British Central Africa. (Nuttall, 1908, fig. 1.)

Fig. 67.

Fig. 68. Fig. 69.

Fig. 67. 0. moubata <? . Anterior portion of venter. Original, N. and W.
Fig. 68. O. moubata. Sketch of second stage nymph's cast skin, seen in profile and

showing line along which it splits running back above spiracle, x 6. Original,

N. and W.
Fig. 69. 0. moubata ? . Ventral aspect of capitulum omitting chelicerae, x 66.

Original, N.

50

Genus Ornithodoros

otherwise they conform to the adult in all essential characters. Nymphs
of intermediate sizes may be encountered down to the smallest or first
stage nymph after its ecdysis from the larval skin and the egg-shell.
In moulting (see Fig. 68) the skin of the adult or nymph splits along
a definite line on each side running from the front margin backward
above the spiracles, the tick emerging anteriorly, but the whole dorsal
cuticle may come off like a cap or adhere for some days to the tick
which has issued. Nymphs of the third stage attain 5 x 1h mm.,
whereas unfed nymphs of the second stage average 1'4 x 1"2 mm.

Fig. 70.

Fig. 72.

Fig. 71.

Fig. 73.

Figs. 70, 71. O. savignyi. Distal ends of legs of adult of large size, (70) leg I and

(71) leg IV.

Figs. 72, 73. 0. moubata. Distal ends of legs of adult of large size (11 mm. I.),

(72) leg I and (73) leg IV with tarsus 1-3 mm. 1.

(Donitz, 1906, pi., figs. 1, 3, 4, 6.)

The Nymph (1st stage, Figs. 74 and 75), immediately after it has
emerged from the larval skin and the egg-shell, measures on an average
1 X "87 mm. After some time it becomes somewhat flattened and
longer, its contour being sub-circular and the colour ochreus. The
minute spiracle is situate as in adults, between legs III and IV. The
mammillations covering the dorsum extend but partially over the
postero-ventral border, the capitulum is well developed, the hypostome

0. moubata

51

bears 2 | 2 rows of teeth, with two prominent hairs on its base as in the
larva. They are usually ready to feed 3 — 4 days after ecdysis^ and may
double in size after feeding.

Fig. 74. Fig. 75.

Fig. 74. Nymph, 1st stage, I. 1*3 mm., unfed, dead and shrivelled. Ventral aspect.

From specimen given by Mr Newstead. Original, N.
Fig. 75. 0. moubata. Nymph, capitulum, x 200. Original, N.

Fig. 76.

Fig. 77.

Fig. 78. Fig. 79.

Figs. 76-79. 0. moubata. After Newstead, 1905, pi. I, figs. 1-4, showing respectively
contour of egg on first day, x circa 40. — Egg on about 10th day with cuticle broken
away showing larva protruding in profile, x circa 40. — Larva removed from the egg,
ventral aspect, x circa 40 (anus, omitted in the original figure, has been added,
G. H. F. N.). — Tarsus of larva, x 250, seen between the foregoing figures.

Larva (Figs. 77-80). The larva can be extracted from the egg-
shell as was first observed by Dutton and Todd (1905 b, p. 126), "it
moves its legs but cannot crawl." It has been fully described by
Newstead (1905 b, p. 21), whose figures are here reproduced. It is ver}-

1 The nymphs were first recorded by Livingstone (1857, p. 382) and described as " the
size of a pin's head."

4—2

52 Ueniis Ornithodoros

imperfectly developed but shows all the essential structures seen in
larval ticks. It is sub-circular, dull purplish brown, and, as it matures,
a Y-shaped mass of excrement accumulates posterior to the anus and
the cuticle grows opaque. The chitin is exceedingly thin, and in shed
skins is colourless. 0. moubata and 0. savignyi are the only ticks at
present known which have an inert larval stage ^, the larva of moubata
appears however to be even more undeveloped than that of the closely
allied species 0. savignyi, for the latter casts its egg-shell before turning
into a nymph.

Fig. 80. O. moubata. Capitulum of larva, ventral aspect and dorsal aspect of left palp.
Whole length of capitulum including palps, 180 fi. The figure differs from that given
by Newstead (1905). Original, N.

Eggs (Fig. 76) : slightly ovoid, glistening, golden yellow when newly
laid, they measure about "9 x "8 mm. (Button and Todd) ; closely
examined, they show " an irregular faint whitish polygonal reticulation
and interrupted radiating streaks" seen through the cuticle (Newstead).
{Biology, etc. see further under Section II.)

Hosts : Besides man, they may attack domesticated animals. Thus,
Wellman (1906, p. 154) states that they bite pigs, sheep, goats and dogs,
etc. in Angola. He has found them in pig-styes. In the Laboratory in
Cambridge, they readily fed on fowls, rabbits, rats and mice. They also
feed on monkeys. (Laboratory experiments in Liverpool and London.)

Geographical distribution : 0. moubata is widely distributed in
Africa, it is chiefly recorded from British East Africa to the Transvaal in
the East, and across the Continent to the Congo and southward to German
South West Africa and Cape Colony. It has been collected in Egypt"
by Boue, Abyssinia by Courbon, to the south of Lake Tchad by Closel
(Nn. 1901, p. 256). Brumpt states (1901, p. 578) that he encountered
it in vast numbers at Biocobaba, Somaliland. In German East Africa
it is found in all places along the caravan route from Dar-es-Salaam to
beyond Kilossa in the direction of Mpapua and on the way from Kilossa

1 This has been confirmed by Donitz (1906, p. 146 ; 1907, p. 20), by E. Koch (cited by
Donitz) and by Nuttall. The large size of the eggs in these species may be noted in this
connection.

2 Brumpt says similar ticks are common in dirty prisons in Cairo, possibly the ticks
he refers to are 0. savignyi.

0. mouhata 53

to Iringa. It is also found in villages in the Rubeho Mountains and in
places off the caravan routes (R. Koch, xi. 1905, p. 1866). Specimens had
been previously collected by Kramer (labelled A. schinzii in Hamburg
Mus.) and Stuhlmann in German East Africa (identified by Nn. 1901,
p. 256). A native from Kilima-ndjaro told Brumpt (p. 578) that the
tick and fever were well known in his country, as was also the case with
the natives of Galla. Dutton and Todd (1905, map) note its presence
at Tabora, and Donitz (1906, p. 145) at Pokomani, Wituland. There are
specimens from Quango in the Berlin Museum (Nn. 1901, p. 256).
Zanzibar Island : it is common in prisons there according to Brumpt,
and this locality is mapped by Dutton and Todd.

In British East and Central Africa, Christy (1903, p. 187) notes it as
fairly common in Usoga, Uganda and Buda, and at Wadelai on the Nile
(North of Lake Albert). Neumann (1901, p. 256) and Pocock (1907)
record it from Namaqualand and Transvaal. Dutton and Todd (1905)
state it is reported to be present in Shescheke, Rhodesia, and Wellman
(ms.) states it occurs in Bechuanaland, Cape Colony and German South
West Africa. In Portuguese East Africa at Tete (Livingstone, 1857,
p. 382) where it occurs to-day. Wellman (ms.) states it occurs at
Mozambique. In West Afnca its presence was noted in Angola, at
Ambaca, and it was stated to be common in native huts of the country
by Livingstone (1857, p. 382). It is from this country that Murray's
types came. Specimens from Angola were identified by Neumann
(1901, p. 256). It is still found at Ambaca and in Malange, Bihe,
Bailundo, Andulo, Caconda, Chiyaka, Benguella, Lovaleland, Moxico and
Landana, according to Wellman (in MS.). In the Congo Free State its
presence was recorded at Nyangwe, on the Lualaba river, by Livingstone
(1874, II. p. 115). Dutton and Todd (1905, p. 123 and map) state that
it certainly occurs at Lokandu, Ukungwa, Mulamba, Mwana, Maketa,
Nyangwe and Kasongo, whilst it is reported to occur in Kimpudi and
from Popokabaka to Francis Joseph Falls, at Katola on the Kasai River,
at points along the route from Kasongo to Baudovinville (west of Tan-
ganyika) and along the western frontier to the shores of Lake Albert
Edward. It certainly occurs northward at Beni. Neumann (1901,
p. 256) recorded it from the Congo and from the upper Zambesi at
Loango, Landana, where it was collected by Foa. Dutton and Todd
consider that the tick came into the Free State from the East Coast
with Arabs, and into the Oriental Province and Cataract Region with
traders from Portuguese territory to the South where the tick existed in
Livingstone's time. The rivers are the present highways. "A glance at

54 Genus OrnitJiodoros

the map will show that ticks are found particularly along much travelled
roads. Although plentiful in many Arabised villages along the Congo
between Kasongo and Ponthierville, they are quite unknown in native
villages an hours' walk inland."

One of us has received specimens collected at Dowa, Lilongwe, and
at Blantyre, British Central Africa ; at Tete, Portuguese East Africa,
and in the District of Benguella, Angola. These, and other specimens
of uncertain origin, have reached us through the courtesy of Messrs
Daniels, Leishman, Newstead and Wellman.

Habitat : Livingstone (1857, p. 628) noted their presence in native
huts at Ambaca, at Tete and " wherever the Arabs go."

Karsch (1878, p. 311) writing of Mombassa, states that " papazS" are
found especially in the " Fort " and the huts of the Wanika and
Wataita.

Christy (1903, p. 187) states that the "bibo" is most easily collected
in Uganda by " searching the dust and straw on the floors of the huts
erected for the caravan porters, or the houses of the natives, though in
the latter it is not so easily found when the floors are kept clean. Near
Kampala the natives collected them around the bases of the vertical
supporters of the roof" He further states that they are " frequently
carried long distances in mats or bedding, or in porters' loads which
have been piled for safety in the rest-huts at night." Some specimens
he collected in Toro had been carried in bags of salt from Kative at the
north end of Lake Albert Edward.

Dutton and Todd (1905 b) suggest that "Perhaps one of the reasons
for which ticks are more often found in Arab than in native houses is
that the Arabs make better, drier buildings, and live in permanent
villages. Native huts are temporary affairs and a slight cause, one or
two cases of sickness, is often enough to make a community leave their
houses and build a village elsewhere." Along the Congo the rest houses
for native travellers were always the most infested. "In infested houses
the ticks are found in the dust and cracks of mud-floors, particularly in
dry places near the hearth, in bed-platforms, or immediately inside the
door-sill, just where the natives are accustomed to sit down. They hide
themselves in the cracks and crevices of mud or grass walls, and even in
the thatched roofs."

Specimens sent to us by Dr Wellman were collected in cracks in
the floor in native kraals. District of Benguella, West Africa. Wellman
states he has found " as many as 100 in one hour in an old native
hut."

0. mouhata

55

See further under Section II, regarding observations on the liatehing
of eggs, oviposition, longevity, natural enemies, feeding habits, effects of
bite and the part played in the transmission of disease.

ORNITHODOROS CORIACEUS C. L. Koch, 1844.

Figs. 1, 58, 81-84.

Icon.: C. L. Koch, 1847, PI. I, Fig. 2, 5 (coloured) ; Fig. 3, ventral aspect in out-
line (very good figures).— Nuttall, vii, 1908, Fig. 2*.— Banks, 1908, PI. I,
Figs. 5, 6, legs.

Lit. : C. L. Koch, 1844, p. 219 ; 1847, p. .31.— Berlese, 1888, p. 193.— Neumann, 1896,
p. 31 ; 1901, p. 258.— Banks, 1908, p. 18.

Body narrow and more pointed in front than in 0. savignyi; integu-
ment with mammillae white in living specimens, reddish or dark in old
preserved specimens ; large reticulated areas where mammillae are
absent (see Fig. 81) not showing up as definite discs. Four eyes
(Figs. 1, 82, 83) placed as in 0. savignyi, hemispherical, greenish, the
anterior large, prominent, the posterior one half smaller. Venter :
spiracles rather large, circular, prominent; pre- and post-anal grooves
well marked. Capitulum closely enveloped in a camerostome ; base
long, four equal post-hypostomal hairs at the same level ; two post-palpal
hairs; palps long, tapering, 1, 2, 4, 3^; long white hairs on dorsal sur-

Fig. 81.

FiK. 82.

Figs. 81, 82. 0. coriaceiis ? , x 4. Dorsum and venter. Drawn from living specimen
a few days after feeding to repletion. Original, N. and W.

1 These numbers refer to the relative lengths of the articles.

56

Genus Ornithodoros

faces of articles 1, 2, 3 ; hypostome spatulate, emarginate at the tip,
extending as far as the middle of article 3 of the palp; a corona followed
by a few large teeth, 2 | 2, about 3 teeth per file, then a few squamous
teeth ; only the distal third toothed. Legs (Figs. 58 (p. 40) and 83)
longer and thinner than in 0. savignyi ; coxae I and II separated, II —
IV contiguous, diminishing posteriorly. Tarsi as in Fig. 58. Closely
allied to 0. savignyi.

Fig. 83. Fig. 84.

Fig. 83. 0. coriaceus ? . Living specimen, lateral aspect (same specimen as in fig. 81),
shows size and position of eyes and spiracles. Original, N. and W.^

Fig. 84. 0. coriaceus d" , x 8. Venter. (The eyes are hidden by the legs, see fig. 1, p. 7.)
Original, N. and W.

C. L. Koch (1844 and 1847) describes this species as follows : '' Shaped
like the sole of a shoe, thick margined, roughly shagreened, yellowish
earthy colour, spotted rusty red, legs toothed dorsally. Length 9*3 mm.
Body about twice as long as wide, width fairly uniform, indented on the
sides, pointed above the mouthparts, rounded posteriorly, a thick turned
up border all around-; the whole surface, above and below, thickly
granulated like fish skin (shagreen), the granules flat above, consequently,
the whole leathery ; on the back unequal folds and grooves. Beneath
in the front of the body a deep groove running to the stigmata and on
the inner protrusion the rather large round quite clearly marked eyes. The
coxae gradually thicken towards the distal extremity and are somewhat
bent; the other articles somewhat compressed and clearly notched or

^ This figure was drawn several days after feeding and the body has shrunk somewhat,
especially in front. Immediately after feeding the body, viewed in profile, appears much
more rounded behind, and the anterior margin is almost obliterated.

- This applies to an unfed specimen.

0. coriaceus 57

round-toothed. The whole surface, above and below, dirty yellowish
earthy colour, rusty red spots irregularly distributed throughout.
Capitulum and palps light yellow. Legs grey-brown. Female.
Male : unknown. Habitat : Mexico." (Translation from the original.)

Berlese (1888) states that he saw a specimen found by A. Balzan at
Rio Apa, Paraguay. We have recently determined specimens received
both from California and Mexico. Those from California^ were collected
by Mr H. A. Moran at Los Olivos in August, 1904, and comprised
7 cf cT and 9 $ $ (Stanford University Coll.); the largest $ measured
138 X 8'2, the smallest 9'5 x 5"3 mm. ; the largest </ measured 8'6x4"6,
the smallest 6'4 x 3"4 mm. Those from Mexico were collected by Mrs
Zelia Nuttall in January, 1908, the lot including 1 (/, 2$?, and
3 nymphs (resembling the ^T). Both females were received alive and
the better specimen was drawn, as also the </ (Figs. 81-84). In respect
of colouration, the female agreed very closely with Koch's figure, the
other specimens are all dull brown like the soil in which they were
found. As noted above, the males are smaller than the females and are
similar in structure but for the sexual aperture (see Figs. 82, 84). The
nymphs were as small or smaller than the males and showed no trace of
sexual " Anlage." The ticks are called " talajas " by the natives, and
were found whilst exploring some rock inscriptions near San Geronimo,
on a hill in a sandy plain (Isthmus of Tehuantepec). Our description is
based on our Mexican specimens.

See further under Section II on feeding habits and effects of bite.

ORNITHODOROS TURICATA (A. Duges), 1876.

Figs. 58, 85-87.

Synonymy : Argas turicata A. Dugfes, 1876.
Ornithodoros americanus G. Marx, 1895.
Native name: "turicata" in Mexico (Dng^s, 1876).

Lit. and Icon.: Dug^s, 1876.— Megnin, 1885, p. 463, PI. XX, Figs. 1-4.— Marx, 1895,
posthumously published plate, appeared in Osborn, 1896, PI. Ill, Figs. 2, 2^ ;
good figures of ^ dorsum, venter, etc. — Neumann, 1896, p. 31, Fig. 33, digit*. —
Ward, X. 1900 a and b. O.— Braun, 1906, p. 373. 0.— Banks, 1908, p. 18,
PI. I, Figs. 13-16, integument, palp, legs 1 and 4.

Adult (Figs. 85, 86) : Body with sides almost straight and parallel,

the anterior extremity narrowed to a round point, the distal portions of

1 Banks, 1908, p. 19, has seen specimens from San Francisco and from Santa Clara
County which had been taken from cattle and cattlemen.

58

Genus Ornithodoros

the capitulum more or less visible dorsally in the replete adult. Integu-
ment thick, with hemispherical, brilliant contiguous granulations, larger
posteriorly. No obvious discs, but tracts where mammillae are absent,
arranged as in 0. savignyi. Numerous clubbed hairs between the
mammillae. No eyes. Venter : coxal and supra-coxal folds well
marked in unfed specimens; a pre-anal groove reaching to the supra-
coxal fold ; a post-anal groove parallel to it, and midway between the
anus and the posterior border ; median post-anal groove reaching the
posterior border. Anus almost as wide as long, with numerous hairs ;
anal frame nearly quadrangular, 225 /a broad. Spiracles circular (180
to 200 IX wide), with crescentic perforate plate. Capitulum with integu-
ment finely honey-combed ; hypostome slightly lanceolate with a crown

Fig. 85. Fig. 86.

Figs. 85, 86. 0. turicata ^ . L. 3*5 mm. Dorsum and venter,
from Guanajuato, Mexico. Original, N. and W.

Specimen

of numerous small teeth, followed by 2 | 2, 11 or 12 teeth per file ; two
hairs at the base of the hypostome ; digit (see Fig. 87); palps long and
only slightly tapering ; articles 1 and 2 equal in length (</ 230 fi,
$ 300 fi) and articles 3 and 4 equal (</ 150 fi, $ 170 fi); numerous
pennate hairs on dorsal surfoce of palps and basis capituli. Legs : coxae
contiguous, decreasing in size from I to IV ; bristling with very fine
spines, reinforced by granulations, especially on the posterior border;
tarsi cylindrical, slightly tapering at their tips; three dorsal humps,
more marked the more anterior the leg, scarcely visible on leg IV
(invisible in our specimen. See Fig, 58, p. 40); similar tubercles,
less salient, on the protarsi.

0. tnricata 59

Males average 3*5 X 25 mm.; females ()x4mm. ; nymphs 2'2xl"omm.

Neumann's description, from which the above is partly taken, is
based on numerous specimens collected in Guanajuato, Mexico, by
A. Duges^

Fig. 87. 0. turicata. Digit, 90 tj. I. (Nn. 1896, fig. 33, modified).

This species attacks pigs, cattle and man. Its bite is painful and is
said to produce grave effects. The 0. americaniis of Marx was found in
Texas on the nostrils of the horse, in South America on the llama
and in Florida on the tortoise, Xerohates polyphemus, whose burrows it
appears to infest (see Hubbard, 1894, p. 306). Banks, 1908, p. 18, has
moreover seen specimens from Florida (in gopher holes). New Mexico,
Arizona and California (on cattle).

ORNITHODOROS TALAJE (Guerin-MeneviUe), 1849.

Fig. 88.

Synonymy : Argas talaje Guerin-MeneviUe, 1849, pp. 342, 343.

Oniithodoros rudis Karsch, 1880, p. 141 ; identified by Neumann, 1901, p. 259,
as 0. talaje after examination of type.

{1 Argas coniceps Canestrini, 1890, p. 535. Regarded as A. rejlexus $ by Canestrini
and Fanzago, 1897, p. 193. From Canestrini's description and figure (see Icon.)
Neumann, 1896, p. 36, appears almost justified in referring it to 0. talaje, of
which he makes it a variety : 0. talaje coniceps (Canestrini). The determination
is, however, doubtful.)

Alectorobius talaje Pocock, 1907, p. 189.

Lit. and Iconography : Guerin-Meneville, 1849, PI. II ; figures inaccurate, rej^ro-
duced by Megnin, 1885, PI. XX, and by Murray, 1877, p. 183. — Canestrini,
1890, PI. XLI, Figs. 1-1 d ; dorsal and ventral aspects, palp, tarsus (like
0. talaje), digit ; sketchy. — Blanchard, 1890, p. 883 et seq. — Neumann, 1896,
Fig. 34*.— Braun, 1906, p. 373. 0.— Banks, 1908, p. 19, PL I, Figs. 7, 8, 17,
palp, legs 4 and 1.

1 We are indebted to Professor Neumann for the gift of one of these specimens from
which our figure was drawn.

60

Grenus Ornithodoros

Body : L. 5 — 6, W. 3 — 3"5 mm., with sides almost straight and
parallel, conical anteriorly, bluntly rounded posteriorly, earthy yellow or
dirty brownish red when replete, dorsum and venter covered by large
glossy mammillae (150 — 200 /i in size); dorsum bearing large discs..
When unfed the dorsal border is raised and thick and the dorsal surface
undulating with a prominence over the capitulum, flanked by four sym-
metrical depressions; all of these, and other depressions posteriorly,
disappear when the tick is gorged. Pre-anal and post-anal grooves,
supra-coxal and coxal folds well marked, the coxal folds passing between
coxae I and II. Post-anal groove with fine transverse striae. Anal
frame almost as wide as long (225 x 200 /i), the valves bearing two to
four hairs each. Spiracles conical, 100 yu, broad, plate crescentic,
situated dorsally on the supra-coxal folds. Eyes absent. Capitulum
lying in a deep camerostome or pit, with prominent lateral flap-like
borders which move as upon a hinge and protect the capitulum ; ventral

Fig. 88. 0. talaje. A, hypostome (xl60). B, digit, 60 m Z. (Nn. 1896, fig. 34).

base of capitulum finely wrinkled transversely, permitting (according to
Megnin) the mouthparts to be protruded and retracted. Chelicera :
digit 60 fji I. (Fig. 88 B) with tridentate dorsal process near base of
internal article and external article bidentate. Hypostome (Fig.
88 J.) emarginate, with many fine teeth distally, followed by dentition
2 1 2 covering half its length ; two very long post-hypostomal hairs.
Palps with articles 1 — 4 measuring respectively 134, 114, 80, 87 fx I.
Numerous pennate hairs especially on dorsal surface of articles 1
and 2, a stout hair at the antero-internal angle of article 1. Legs
long and slender; coxae contiguous, decreasing in size from pair I
to IV, covered with granulations or mammillae posteriorly; tarsi
tapering without marked distal protuberance, unless on pair I ; all
the articles bear semi-pennate hairs, which are longest on tarsi.

The foregoing description (condensed from Neumann, 1896, pp. 34-
36) is based on the examination of three specimens from Cumana

TICKS

PLATE

Fig. 1. 0. talaje, var. capensls. Dorsal aspect, x 16. (Lounsbury Coll.)
Original, C. and R. phot.

'#

Fig. 2. 0. talaje, var. capeitsis. Ventral aspect, x 16. (Lounsbury Coll.)
Original, C. and E. phot.

O. talaje 01

Venezuela (Paris Mus.), and two specimens collected by Sall^ in Guate-
mala. Guerin-Meneville's specimens were collected on the route from
Guatemala to Zacapa. Megnin's specimens were sent from Mexico by
Duges. Karsch's specimens (in Berlin Mus., identified by Neumann)
were collected in New Granado. by Goudot. According to Neumann
(1901, p. 258) the Paris Museum possesses specimens collected by Gay
at Santiago de CJiHi and by Steinheil in Colombia, and Shauinsland
found a nymph on the Island of Laysan, Hawaii (Poppe Coll.). We
have examined specimens, determined by Neumann for the British
Museum, from native houses at Santana-Totima, Colombia (nymphs
2'5 mm. I. ; adults 5 x 2*5 mm.). Guerin-Meneville's specimens measured
5 — 7 mm. l.\ those of Karsch 5"5 x 3 mm. Banks (1908, p. 19) records
specimens from Florida, Texas and San Clemente Island, California.

Canestrini (1890, p. 536) found his Argas coniceps, together with
A. rejiexus, in the interstices of the mosaic of the Church of San Marco,
Venice. Birula states that several specimens taken near the Sea of A ral
are in the Zool. Mus, of the Acad, of Sc, St Petersburg. Canestrini's
specimens measured 5 — 6 mm. I. The determination of this species or
variety is doubtful.

This species approaches 0. turicata, but is distinguished especially
by the structure of the capitulum and legs.

Habitat : native houses, attacks man. (See further under Section II
for effects of bite, etc.)

ORNITHODOROS TALAJE var. CAPENSIS Nn., 1901.

Figs. 58, 89 and PI. III.

Lit. : Neumann, 1901, p. 258 ; 1907, p. 193.

According to Neumann this variety differs from the American type
chiefly in (1) the cheeks or flaps being less developed in width, more
separate at their attached borders, which do not appear capable of com-
pletely hiding the hypostome and palps and do not attain the level of
the ventral surface of the base of capitulum ; (2) by the longer and more
abundant hairs on the movable articles of the legs. This variety is
based upon numerous specimens collected in Penguin nests, on guano,
on Islands off the coast of Cape Colony (Lounsbury Coll.). It readily
attacks fowls when given the opportunity. It abounds in its native
home. (Nn. 1901, p. 258.) Through the courtesy of Mr Lounsbury
we have been able to examine his specimens. Moreover, the British

62 Greniis Ornithodoros

Museum possesses numerous specimens of nymphs and adults collected
by the Challenger Expedition, 28, viii. 1876, at St Paul's Rocks from
birds' nests, where they were found together with small spiders, etc.
The specimens, preserved in spirit, were mostly yellowish or reddish
brown in colour. Neumann (xii. 1907, p. 193) records specimens found
on the ground at Cargados Carajos (Siren Island) by Mr J. Stanley
Gardiner.

Fig. 89. 0. talaje var. capensis s . Anterior portion of venter (same
specimen as figured on PI. III). Original, N. and W.

ORNITHODOROS PAVIMENTOSUS Neumann, 1901.

Figs. 90-92.

Lit. and Icon. : Neumann, 1901, p. 257, Fig. 1, of legs I and II*— Donitz, 1906,
pp. 144-148, PI., Figs. 2 and 3, of legs I and IV*

Male : Unknown.

Female : Body short oval, rounded at both ends, 12x8 mm. ;
covered by contiguous, fiat granulations (whence " pavimentosus "),
smaller on the depressed areas; coarse hairs, especially anteriorly.
In other respects like 0. savignyi, except that the protarsi and tarsi
of legs I, II and III are much shorter, with the dorsal protuberances
arranged close together (see Fig. 58 and compare with Figs. 70, 71).

Neumann's description is based on a single dried ^ % taken by
Schenk at Bethany, Great Namaland, S. Africa (Berlin Mus.). In his
original description, Neumann pointed out the differences in the leg
structure between this species and 0. savignyi (should be 0. moubata as
pointed out by Donitz). According to Donitz, who has examined many

This may account for the " black " eyes.

O. imvimentosiis

63

specimens collected in Namaland by Schultze, 0. savignyi generally has
larger humps on the tarsi, but in two specimens of savignyi from
Dongola, no differences in leg structure were observable. According
to Schultze, 0. pavimentosus has the same habits as 0. moubata (q.v.),
being common in places where travellers rest, and attacking men when
they lie upon the ground (apparently its attack is not confined to the
night time).

Fig. 90.

Fig. 91.

Fig. 90. 0. pavimentosus. End of leg I of a specimen measuring 12 mm. I. (Donitz,

1906, pi., fig. 2.)
Fig. 91. 0. pavimentosus. End of leg IV of same specimen as preceding, tarsus

2-4 mm. I. (Donitz, 1900, pL, fig. 2.)

Fig. 92. 0. pavimentosxis. Ends of legs I and II. (Nn. 1901, fig. 1.)

ORNITHODOROS ERRATICUS (Lucas), 1849.

Syn. and Lit.; Argas ermticus Lucas, 1849, p. 316, gives a fairly good description,

considering the date.
Ornithodoros miliaris Karsch, 1880, p. 141: brief Latin description of three lines.

Neumann, 1901, p. 256, examined the type and referred it to 0. erraticus.
Ornithodoros erraticus (Lucas) Neumann, 1896, p. 37.

Icon.: No figures published.

64 Genus Ornitliodoros

. Body 5x3 mm., oval (lateral borders parallel in the young), tapering
anteriorly to a rounded point. Integument thin, with closely crowded
hemispherical granulations of unequal size (75 to 100 /i), with some
short claviform hairs between them ; large symmetrical discs, of which four
in a transverse series across the middle of the dorsum ; another transverse
series in a groove which limits posteriorly a dorsal prominence corre-
sponding to the capitulum ; a broad undifferentiated margin. No eyes.
Venter : well-marked pre-anal groove, concave behind ; post-anal groove
midway between the anus and the posterior border; post-anal median
groove extending beyond the post-anal, but not reaching the posterior
border ; anal frame 230 x 200 /x. ; anus with hairs on the valves.
Spiracles semicircular (?), 100 ju. wide. Capitulum free, exposed ven-
trally ; palps 1, 4, 2, 3' ; numerous pennate hairs on the dorsal surface of
the palps and basis capituli ; hypostome resembling that of 0. talaje, a
crown of denticles being followed by 2 | 2 files of 5 or 6 stout teeth, then
squamiform teeth 3 j 3, then 4 | 4 ; two pennate hairs at the base of the
hypostome. Legs medium ; coxae contiguous, the first pair only being
slightly granulated ; article 2 conical, the others cylindrical ; tarsi only
slightly humped.

Neumann's description, from which the above is taken, is based on
three specimens from Algiers. Pavesi (1884, p. 485) records two
specimens, 3'5 mm. I., found by Doria in the vicinity of Tunis, one
from Nemours and two from Marnia (?). Lucas received his three
specimens (4 x 2*5 mm. in size) from the plain of Lake Houbeira, near
La Calle. They were found beneath stones where Bu/o pantherinus was
also in hiding. Karsch's 0. tniliaris came from Bengal (type in Berlin
Mus., Karsch gives the size at 2*5 x 1*7 mm.). Neumann states that it
resembles 0. talaje, but does not possess the lateral expansions of the
camerostome. These specimens also differ in the details of the digits of
the chelicerae, and in the tegumentary characters of the legs.

^ The relative lengths of articles.

0. tliolozmii 65

ORNITHODOROS THOLOZANI (Laboulbene and
Megnin), 1882.

Figs. 93, 94.

Syn., Lit, and Icon. : Argas tholozani Laboulbfene and Megnin, 1882, pp. 335-337,
Pis. XXII and XXIII ; the figures include : adult dorsum and venter, capitulum,
hypostomes ( ,^ and 9 )> digits, palp, integument, tarsus ; larva (showing intes-
tinal caeca apparently passing into the legs), the ovum. — Blanchard, 1890.
—Neumann, 1896, p. 38, Fig. 35*. — Dubreuilh and Beille, 1896, p. 80, Fig. 8
(copied from Laboulbene and Megnin). — Braun, 1906, p. 373. O.

{Argas papilUpes Birula, 1895, pp. 38-44, Fig. 35 ; digits of J and $ ; referred here
by Neumann, 1901, p. 259. See 0. papilUpes, p. 79.)

Native name: "Kene"or "sheep bug" in Persia (Laboulbfene and Megnin, 1882,
p. 329).

Adults : Body (the (/ varies from 4x2 mm. to 6 x 4 mm. ; the %
from 8x4 mm. to 10x5 mm.), with lateral borders nearly straight
and parallel, tapering abruptly, but sinuously, to a rounded process
in front, and broadly rounded behind; a broad margin is marked
off from the rest of the dorsum by a well-defined marginal groove.
Integument coarsely shagreened, then, with numerous short hairs,
longer and more apparent anteriorly; hemispherical granulations 40 —
60 jj, in diameter ; giving the effect of a network of meshes limited by
creased folds ; one mesh out of every 5 or 10 bears a long hair. Discs
similar to those of 0. erraticus, but relatively smaller; seven on the
anterior median prominence arranged in three rows (3, 2, 2), surrounded
by an irregular circle of similar discs whence a median row starts
posteriorly, double at its origin and reaching the posterior border. A
short row on each side, and other scattered discs. On the ventral
surface discs occupy the depressions and grooves. No eyes. Venter:
pre-anal groove deep, concave behind; post-anal groove nearly straight;
median post-anal groove nearly reaching the posterior border; sexual
orifice in ^ and $ between coxae II ; anal frame wider than long
(300 X 350 /Lt); anus with seven or eight long hairs on each side. Spiracles
(350 yu,), crescentic. Capitulum : palps tapering, and not much longer
than the hypostome; chelicerae (digits. Figs. 93, 94). The hypostome
differs in the sexes: in the f^, it is somewhat spatulate and indented, a
crown of small teeth being followed by two rows of three teeth on each
side, the middle rows not ftir apart ; then three or four rows of simple
squamiform teeth ; in the $ , it is somewhat lanceolate, and the dentition

66 Genus Ornitliodoros

is 2 I 2 throughout, the median rows being far apart. Four long hairs
(post-hypostoraal and post-palpal) on the base of the capitulum, as in
0. avignyi. Legs fairly long and slender; coxae II — IV contiguous ;
tarsi 2 — 4 with terminal dorsal protuberance prominent, pointed and
directed distally, the terminal portion tapering. Short hairs on all
articles, longest on the tarsi.

Fig. 93.
Figs. 93 and 94. O. tholozani. Left digits of <? and ? . (Nn. 1896, fig. 35.)

Nymph : resembles the male but has no sexual orifice.

Larva : 1 x "5 mm. when newly hatched ; a circular dorsal area
(disc ?) ; the intestinal caeca pass into the legs (Laboulbene and Megnin).

Egg: yellowish brown, 'Sx'Gmm.; about 100 laid by each $
(Laboulbene and Megnin).

This species is allied to 0. erraticns, from which it may be distin-
guished by its tarsi and the dimorphism of the hypostome.

Origin : Persia, where Neumann states they were collected by
Tholozan, mostly in a chicken yard at Teheran, others were found on
camels (specimens in Paris Mus.).

The above descriptions of the adult and nymph are condensed from
Neumann, who has communicated certain particulars by letter (6 April,
1908), stating that Laboulbene and Megnin figure the tarsus inaccurately,
and that his description is based on the examination of specimens
received at the Paris Museum from Dr Tholozan at the time when
Laboulbene and Megnin received theirs from the same source. We have,
unfortunately, not succeeded in obtaining a specimen of this tick.
Neumann's description does not agree in some respects with Laboulbene
and Megnin's figures, but he assures us that these are inaccurate.

Laboulbene and Megnin (1882, p. 329) state that they were fed on
man and rabbits in Paris and (p. 337) that unfed specimens survived
four years.

0. lahorensis

67

ORNITHODOROS LAHORENSIS Neumann, 1908.

Figs. 95-100.
Lit. and Icon.: Neumann, 1908, p. 17, Figs. 12-15 (here reproduced, p. 68).

Adult (Figs. 95-97). Body: ^ 8x4-5 mm., ? 10x5-6 mm.;
lateral borders parallel with a faint constriction behind coxae IV ; the
anterior extremity narrowed to form a conical hood, more pointed in the
^ ; posterior border broadly rounded. Colour earthy yellow, legs paler.
Dorsum convex anteriorly, irregularly concave posteriorly, grooves and
discs as in figure, the principal discs being 2 oval, parallel, near together
in front, and 4 circular forming a trapezium, narrower anteriorly, in front
of the middle of the dorsum, the integument between the discs not

Fig. 95.

Fig. 96.

Figs. 95, 96.

0. lahorensis ? . After feeding and ovipositing, 10 mm. I.
Original, N. and W.

granular but finely wrinkled, the wrinkles radiating from the discs. A
few very short and scattered hairs. No eyes. Ventei' : anus about half
way between the basis capituli and the posterior border ; pre-anal
groove shallow and incomplete ; post-anal groove immediately behind
the anus, a well-marked ano-marginal groove, occupied by a file of discs ;
on either side of it several depressions with discs ; camerostome tri-
angular, deep, narrow, longer than broad ((/) or about as long as broad
and less pointed ( $ ), its sides converging and uniting in part in a
transverse lip. Hairs longer and more abundant, especially in front,

6—2

68

Genus Ornitliodoros

than on the dorsum. Spiracles semi-lunar, equal to (?) or larger (j/*)
than the anal ring.

Fig. 97. 0. lahorensis ? . Nn. 1908, fig. 12 (circa 10 mm. I.).

Fig. 98.

Fig. 99.

Fig. 100

Fig. 98. 0. lahorensis. Adult, digit, x 215. Nn. 1908, fig. 13.

Fig. 99. 0. lahorensis. Adult, hypostome, x 120. Nn. 1908, fig. 14.

Fig. 100. 0. lahorensis. Adult, tarsi I— IV, x 28. Nn. 1908, fig. 15.

y/.

///

/1^

0. lahorensis 69

Gapitulum ( $ about 1*2 mm. long) ; chelicerae (see digit, Fig. 98) ;
hypostome (Fig. 99) long, lanceolate, indented at the end, 2 | 2 files of
eight or ten teeth, the internal files far apart ; palps cylindro-conical, the
articles 300 /i, 250 jju, 150 jj, and 200 yu. respectively ; long hairs, curved
forward, on dorsal surface of articles 2 and 3. Legs medium, coxae
(Fig. 96) sub-conical, coxae I and II slightly separated, the others
contiguous ; tarsus I (Fig. 100) with three blunt dorsal prominences ;
tarsi II, III and IV" with very short proximal false articulation, promi-
nent dorsally, and with progressively longer blunt spurs near the distal
end of the distal false-articulation.

Nymph : Neumann has recognised two nymphal forms :

First nymph : o"5 x 2*7 mm. Integument with very fine folds, with
a few spiniform hairs, discs hardly visible ; the ventral grooves only
faintly marked, except the ano-marginal. No trace of sexual aperture ;
camerostome shallow ; legs short and thick.

Second nymph : attaining 11x6 mm. Integumental folds coarser
and polyhedric ; discs visible, hairs almost absent ; a punctiform sexual
orifice^ ; camerostome deeper, but less completely filled by the capitulum ;
legs like the adult, but with spurs less pronounced.

Taken from Ovis aries in Lahore by E. Montgomery (all were collected
in the nymphal stage in January, 1906, but six weeks afterwards some
gave rise to adults. Collection of the Liverpool School of Tropical
Medicine). Our figure is drawn from a living $ kindly sent to one of
us by Mr Newstead (Liverpool). We received a ^f and $ . The cT fed for
25' on a fowl 7. v. 1907, for 45' on 2. xii. 07 and was killed and preserved
27. iii. 08. The ? refused to feed 7. v. and 11. vi. 07, she began to
oviposit 27. xi. 07, fed for 2 hours 20' on 2. xii. and began again to
oviposit 9. xii. She lived until iii. 08 when she was lost. All the
eggs, about 80, proved sterile. The ticks refused to feed on other
occasions (G. H. F. N.). Our specimens agreed so closely with the de-
scription of 0. tholozani given by Laboulbene and Megnin that we
referred it to this species, from which, however, as Neumann informs us,
it differs in the structure of the integument and tarsi, L. and M.'s figures
of the tarsi being inaccurate.

^ Doubtless but an " Anlage," in any case imperfect.

70 Genus OrnitJiodoros

ORNITHODOROS FURCOSUS Neumann, 1908.

Fig. 101.

Lit. and Icon.: Neumann, 1908, p. 21, Fig. 16 (here reproduced).

Female : 10 x 5 mm. ; lateral borders nearly parallel with anterior
conical hood ; posterior border broadly rounded ; blackish-brown, capi-
tulum and legs paler. Dorsum irregularly convex ; integument granular,
the granulations larger than the intervals between them ; a few long
hairs more abundant in front. No eyes. Venter convex, the folds not
well-marked ; anus nearer to the posterior border than to the vulva ;
camerostome shallow, without lateral folds ; vulva between coxae I.
Spiracles circular, prominent. Capitulum (about 1*4 mm. I.); base flat
ventrally, very convex dorsally, nearly as deep (dorso-ventrally) as broad ;
hypostome somewhat lanceolate, 2 | 2, the inner files far apart ; palps
long, tapering, forwardly curved hairs on the dorsal surface of articles 1,
2 and 3. Legs : long, especially pair IV ; coxae sub-conical, contiguous
with wrinkled integument ; coxa II larger than I ; coxa III smaller
than I ; coxa IV still smaller ; article 2 sub-cylindrical, rather longer
than broad ; articles 3, 4 and 5 very long and swollen at their distal
extremities ; tarsi forked, owing to a sharp terminal dorsal spur ; tarsus I
has three low, rectangular dorsal prominences in addition ; tarsus IV very
long.

Described from one ? in bad condition, taken at Riobamba(Equador),
by Dr Rivet (Paris Mus.).

//.

J— U-^.^^^

rc

IK

Fig. 101. 0. furcosus ?. Tarsi I, II, IV, x 20. Nn. 1908, fig. 16.

0. megnini 71

ORNITHODOROS MEGNINI (Duges), 1883.

Figs. 102-112.

Synon. : Argas megnini Dug6s, 1883, p. 196.

'■^ Argas americana Packard" : Townsend, 1893, p. 50.
Rhynchoprium spinosum Marx, 1895, p. 199.
Ornithodoros megnini (Dugfes) Neumann, 1896, j). 42.
The " Spinose Ear Tick " of American authors.

Lit. and Icon.: Dugfes, 1883, p. 195i.— Megnin, 1885, p. 460, PI. XXI, Figs. 1-8.—
Neimaann, 1888, p. 96'-^; 1892a, p. 101- ; 1892b, p. 105.— Blanchard, 1890,
p. 883 et seq. 0.— Eailliet, 1895, p. 718.— Townsend, 1893, p. 49.— Dolly,
1894, p. 9802.— Marx, i895, p. 195, Figs, la-i, reprinted by Osborn, 1896
(p. 255), PI. Ill; Figs. 1 and la reproduced by Salmon and Stiles,
1901, and in this book. — Neumann, 1896, p. 42, Fig. 36 (here reprinted).
— Ward, 1900a, p. 199, Fig. 3 (good figures of nymph, dorsal and ventral
aspects, but termed " full-grown " adult by author ; agrees very well with
Marx's figures). — Stiles and Hassall, 1901, repr. 2 pp. — Salmon and Stiles, 1901,
p. 408, PI. LXXIX (coloured, of nymphs and adults (?) natux'al size and x 2),
two figures after Neumann's Fig. 36 ; Fig. 98, integument of nymph ; Figs. 103,
105-107 rejjresent the larval capitulum, the "pupa-like stage," the larval tarsus ;
the remaining (essential) figures here reprinted. — Simpson, 1901, repr. 7 pp.,
3 Figs. (2 photographs of spinose nymph, dorsal and ventral aspects ; rough
sketch of spiracle, both by S. G. Wheler).— Wheler, 1901, p. 61 ; 1903, p. 49.—
Banks, 1904, Fig. 72, after Marx.— Braun, 1906, p. 374. 0.— Hooker, 1908 a,
pp. 40, 42, 45, 51.— Banks, 1908, p. 17 ; PI. I, Figs. 9-12, palp, legs 4 and 1,
integument.

Adult (Figs. 102, 103) : Body panduriform, slightly attenuated
anteriorly ; broadest at legs 2 or 3, constricted behind legs 4, broadly
rounded posteriorly ; colour brown to violet or black. Dorsum : sym-
metrical depressions, one arciform, longitudinal, from each side of the
projection corresponding to the capitulum ; lateral and slightly anterior
to this an infundibuliform depression ; posteriorly along the lateral
border, a longitudinal groove of variable length, sometimes interrupted ;
a median groove occupying the greater length of the posterior half
Venter : pre-anal groove absent ; replaced by a first post-anal groove at
a tangent to the anal frame, straight and joining the coxal groove ; the
true post-anal groove near posterior border ; a broad median anal groove
terminating at this posterior groove. Supra-coxal and coxal folds broad,

1 Original inaccessible, passages quoted in Salmon and Stiles, 1901, and referred to by
Neumann, 1896.

2 Unimportant, cited in bibliography by Salmon and Stiles, 1901.

72

Genus OrnUhodoros

well marked. Spiracles circular, 250 /x in diameter, stigmal plate
semilunar (resembling Fig. 10). Anus: anal frame broader ^than long
(150 to 200/ix200 to 250 /i), each valve provided with four short
hairs. Ei/es absent. Integument (see Fig. 102) with small circular,
shallow pits (not mammillae) with short central hairs, the whole surface

Fig. 102.

Fig. 103.

Figs. 102, 103. 0. megnini c? , x 8. Dorsum and venter. From specimen, loaned by
Prof. Neumann, from New Mexico, U.S.A. Original, N. and W.

Fig. 104. 0. viegnini. A, digit of nymph, 85 yit I. B, digit of 2 , 75 fi I.

being finely granular like sandpaper. Reticulate fossettes occupy the
ventral and dorsal depressions ; others scattered here and there. On
the ventral surface, between the two post-anal grooves, extremely small
and crowded spines. Capitulum very small and very short; 150 yu,
(from base of palps to tip of hypostome). Chelicerae: see Fig. 104 B.

O. megnini

73

Hypostome short, broad basally, rounded distally, unarmed'^. Palps thick,
300 IX long (90 [x, 85 /u., 65 /x, 60 yu,, from first to fourth article), with
articles relatively broad and short, the second 110 At broad; on dorsal
surface of the articles, especially on the first and dorsally on the basis
capituli, posterior to the palps, numerous pennate hairs. Two post-
hypostomal hairs. Legs comparatively thinner and a little shorter than
in the nymph ; coxae disposed as in the other members of the genus ;
a dorsal tuberosity on tarsi (see Fig. 58). Size : $ Z. 5 to 6 mm., W. 3
to 4 mm. ; (^ somewhat smaller. We have received 2 </ and 1 $ from
Texas (W. D. Hunter), the (/•(/ measured 67 and 7 mm. L., the ?
7*5 mm. L.

Fig. 105.

Fig. 106.

Figs. 105, 106. 0. megnini. Unfed. Nymph, dorsal and ventral aspects. Salmon and
Stiles, 1901, figs. 94, 95 after Marx, 1895, pi. II, figs. 1 and 1 a. 17th Ann. Eep.
B.A.I. , U.S. Dept. Agric.

Nymph (Unfed. Figs. 105-107). Body somewhat lozenge-shaped,
slightly longer than broad, greatest breadth at about the third pair of
legs, anterior to this the outline is rounded ; posteriorly, suddenly con-

Note observations on this point in Section II.

74

Genus Oi^nithodoros

tracted at the fourth pair of legs ; border thick, dorsal surface more or
less undulating. Integument, both ventrally and dorsally, is beset with
posteriorly directed spines or bristly hairs; the spines on the dorsum are
much crowded and form a crescentic area anteriorly with concavity
directed posteriorly, the horns thereof extending laterally to legs 2 and 3
(Neumann) or to legs 3 and 4 (Salmon and Stiles) ; the rest covered by
bristly hairs ; ventrally, the spines extend to or slightly beyond the

Fig. 107.

Fig. 108.

Fig. 107. 0. megnini. Unfed. Nymph, ventral aspect. Salmon and Stiles, 1901, fig. 100.

17th Ann. Kep. B.A.I., U.S. Dept. Agrie.
Fig. 108. 0. megnini. Capitulum of nymph, ventral aspect. Salmon and Stiles, 1901,

fig. 99. 17th Ann. Eep. B.A.I., U.S. Dept. Agric.

anus ; integument finely striated, pits absent. Colour, unfed, earthy
yellow ; replete, violet brown ; when replete, the contours more rounded,
integument glossy, the spines more separated owing to stretching of
integument between them. Anus very small, slightly broader (110 to
120 yu,) than long ; two long hairs on each external border. Spiracles
300 to 350 /i in diameter, on salient tubercles^ (seen in Fig. 105) near

1 Wheler (1901) states that in the living specimen he has seen a "pointed organ,"
protruded and retracted from the apex of the tubercle, and he gives a rough figure of it in
Simpson's paper. The structure requires further study.

0. megnini 75

the dorsal border, between coxae III and IV ; stigmal plates covered by
numerous hairs. Capitulum (Fig. 108) subterminal in first-stage nymphs,
inferior in later stage nymphs, elongate, base as long as palps ; about
375 /J, long (fi'om base of palps to tip of hypostome) ; chelicerae (see
Fig. 104) A); hypostome lanceolate, dentition 4 | 4, with 7 to 9 teeth per
file; palps thick (articles 1 to 4 measure 140, 125, 110, 180 /a); second
article 110 yu, broad; simple hairs, not very numerous, on articles 2 — 4,
six to eight short terminal bristles; some short thick hairs, symmetrically
placed, on basis capituli. Legs long and strong ; coxae very short, but
broad, distant from median line, almost marginal ; other articles cylin-
drical, relatively short compared with their diameter ; terminal portion
of tarsi elongate, claws strong, pulvillum absent, hairs sparsely distri-
buted on all articles. Size: 3 to 4 mm. long, when unfed, to 8%5 mm.,
when maturer and gorged, the latter 5*5 mm. wide (Neumann).

The foregoing description of the adult is partly based on that of
Neumann (1896, pp. 43, 44), that of the nymph on the descriptions of
Neumann (ibid., pp. 42-43), and Salmon and Stiles (1901, pp. 409-410).
According to Hooker (1908, p. 40 et seq.) there is but one nymphal
stage. The full-sized nymphs we have seen conform to the adults in
general appearance but differ in having no sexual aperture and having
the hypostome armed as in Fig. 108. We have verified the descriptions
by the examination of specimens in our possession, and have added
matter of our own.

Larva^ (Figs. 109-110): hexapod, elongate oval, with long terminal
capitulum, measuring 0'61 mm. from tip of hypostome to posterior ex-
tremity. Integument finely striated with symmetrically arranged pits,
from which bristle-like hairs issue. Cajntultim : hypostome 2 j 2, almost
as long as palps; palps: articles 2 and 3 equal, 1 and 4 short; large pits
In inter-coxal spaces 1 and 2. Legs : pulvillum small (see Fig. Ill),

Gorged larvae attain 4x2 mm. (Salmon and Stiles' " pupa-like
stage," see Fig. 112); they appear as "rounded white pyriform struc-
tures with one end elongate," the "distended smooth hexapod skin
with terminal capitulum " contains the spinose nymph " with subter-
minal capitulum, and with well-developed claws on the tarsi, and four
rows of denticles on each half of the hypostome." (Condensed and
modified from Salmon and Stiles, 1901, pp. 410-411.) This is obviously
the stage which Townsend (1893, p. 49) interpreted as an egg.

"This tick is remarkable for the difference in appearance between

^ Also described by Stiles and Hassall, I'JOl.

76

Genus Ornitliodoros

Fig. 110.

Figs. 109, 110. 0. megnini. Larva, dorsal and ventral aspects. Salmon and
Stiles, 1901 (figs. 101, 102. 17th Ann. Rep. B.A.I., U.S. Dept. Agric).

O. iiiegnmi

77

the young spiny stage and the adult form, a difference that is so great
that the two stages have been described as belonging to different
species " (Salmon and Stiles). The identity of the different stages
has, however, been fully established by Neumann and Salmon and
Stiles.

Hosts : occurs chiefly in the ears of the horse, ass, ox, and not
infrequently in the human ear in Mexico (Duges). Simpson (1901) has
reported a case in which two spinose nymphs occurred in the ear of a
gentleman who came to England from Arizona where he had been
camping in June. One tick came away and the other was removed in
the end of August.

Fig. 111.

Fig. 112.

Fig. 111. 0. megnini. Leg of larva, tarsus I ia profile.
Fig. 112. 0. viegnini. Larva, gorged, so-called " resting- or pupa-stage."
Both figures from Salmon and Stiles, 1901 (figs. 108 and 104.
17th Ann. Eep. B.A.I., U.S. Dept. Agric).

Geographical distribution : it was first described from Mexico by
Duges (1883), who states it is very abundant in the State of Guanajuato,
and it has been reported from various parts of the United States, i.e.
Louisiana, Texas, New Mexico, Arizona, California, Nevada, Idaho, Iowa,
Kentucky, Kansas and Nebraska (Salmon and Stiles, 1901, p. 411;
Banks, 1908, p. 17).

78

Genus Oimithodoi^os

ORNITHODOROS CANESTRINII (Birula), 1895.

Fig. 113.

Synon.; Argas canestrinii Birula, 1895, p. 353. — Neumann, 1901, p. 260.

Icon.: Birula, 1895, PI. I, Figs. 1-3. Anterior part of venter, tarsus I, anus (Figs.
1 and 2 here reproduced).

Body ((^ 10 X o mm., $ 14 x 8 mm.) elongate, with sides sub-parallel,
rounded posteriorly, cone-shaped anteriorly. General colour fawn or
blackish fawn-coloured, lighter on venter ; palps and legs lighter. Eyes
absent. No grooves on venter. Integument finely wrinkled dorsally, with
rounded depressions, shallow, confluent in places ; on the ventral surface,
integument smooth on the median line and on coxae ; sexual aperture
between coxae I (in ^ and $ ) ; on each side of the base of the capitulum.

Fig. 113. 0. canestrinii $ ; anterior part of venter, and tarsus 1 ; slightly modified
from Birula, 1895, pi. I, figs. 1, 2.

a fold half as long as capitulum. Capitulum : base rectangular, scarcely
wider than long ; hypostome elongate, rounded and not indented at its
apex, one and a half times as long as the base; dentition 2 | 2, two rather
long hairs ventrally on its base ; palps longer than hypostome, bearing
hairs on their dorsal border ; article 1 very thick ; article 2 of the same

0. canestrinu

79

length, more slender ; article 3 a half shorter. Legs stout ; coxa I
divided in two short, rounded teeth ; tarsus 1 bearing three successive
protuberances on dorsal border ; a single one near distal extremity of
the other tarsi.

Found at Teheran, Persia, by E. v. Keyserling (1839) and at Tasch-
Burun, Caucasia (1885). (After Birula's description with the aid of his
figures.)

We are not acquainted with this tick but admit it provisionally as a
good species since in the detailed figure it resembles no species known
to us.

ORNITHODOROS PAPILLIPES Birula, 1895.

Fig. 114.

Lit. and Icon.: A. Birula, 1895, p. 354, PI. I, Figs. 4-6, anterior portion of venter,
including capitulum, tarsus IV, Hallers organ (Figs. 4 and 5 here reproduced).

Body 7x4 mm. elongated, oval, rounded behind, prominent in front
(uncleft) ; reticulate-rugose, grey or greyish-yellow. No eyes. Lateral
folds of camerostome constricted in the middle. (Apparently free in
figure.) Venter : genital orifice behind coxae I. Capitulum (1 mm. I.),
base almost square, 3 small setae on each side ; hypostome 2 | 2, about
as long as the base ; palps short and thick ; two long hairs at base of

Fig. 114. 0. papillipes ?. Anterior part of venter, and tarsus IV ; slightly modified
from Birula, 1895, pi. I, figs. 4, 5.

80 Genus Ornithodoros

hypostorae ; two hairs posterior to the palps. Legs (pair I, 4 mm.,
pair IV, 5 mm. I.) ; a single hump on all the tarsi.

Neumann is probably correct in considering 0. papillipes a s3monym
of 0. tholozani, since, as he informs us in a letter (6. iv. 08), in the copy
of Birula's paper which he had received from the author the latter had
crossed out the name papillipes and substituted " tholozani (Laboul-
bene and Megnin)." He points out, also, that the integument is similar
" corporis derma reticulato-rugosum." Birula's figures, however, are
difficult to reconcile with the description of 0. tholozani given above,
especially as regards the sides of the camerostome and the tarsi, so that
we have thought it best to insert them here, together with his descrip-
tion of 0. papillipes.

Birula's specimens were collected in the Caucasus by Motschulsky.

ORNITHODOROS MORBILLOSUS Gerstaecker, 1873.

Lit. and Synon.: Ornithodoros morbillosiLS Gerstaecker, 1873, p. 464.

Ornithodoros savignyi (Audouin) ; referred to this species by Neumann, 1896,
pp. 27-29, but Donitz, 1907, p. 23, who has examined the tyjie, states that
it differs in the leg structure as noted below.

Species based on a single specimen, measuring 6*3 x 4"3 ram., which
may be an abnormality. Donitz (1907) states that it differs from
0. savignyi in having 2 humps on tarsus IV instead of 3. Gerstaecker
only had Savigny's figure (1826) to consult in determining the differ-
ences. Origin : Lake Jipe, German East Africa, October 1862.

81

SECTION II.

DEALING WITH THE GENERAL BIOLOGY OF THE ARGASIDAE— THE
EFFECTS OF THEIR BITES —THEIR RELATION TO THE SPREAD
OF DISEASE, ETC.

Argas persicus (p. 81), reflexus (p. 91), bmmpti (p. 95).

Ornithodoros savupu/i^, coriaceics {-p. 102), turicata (p. 102), talaje (p. 103), pavi-
mentosus^, tholozani (p. 103), lahorensis^, megnini (p. 103).

ARGAS PERSICUS.

The life-history has been carefully studied by Lounsbury (1903,
pp. 6—11).

The eggs are laid in the hiding places of the adults, in cracks and
crevices in the walls, etc. Fuller (1896, p. 593) and Brown (1902, p. 86)
state that oviposition takes place occasionally on fowls. The eggs are
laid in batches of about 20, 50, 100 more or less, usually about a week
after feeding in warm weather, and hatching begins in about three
weeks.

The larvae which issue from the eggs are almost colourless, and in
Lounsbury 's experience, die if unfed within 8 weeks. When the
chitin has hardened they attach themselves to a host within a few
minutes, as we have observed. After about 3 days they swell visibly,
appearing as small dark spheres on the skin of the fowl. On the 5th
day they drop off, but if the weather is cool they may remain attached
for 10 days. The body has now flattened (Fig. 20, p. 16) and assumed
somewhat the form of that of the adult. In summer, moulting follows
after about 8 days (Lounsbury).

First nymphal form: the ticks feed quickly from now on to the
adult stage, taking 1^ — 2 hours to gorge-. After about two weeks, in
warm weather, they moult and the second nymphal form appears.
These in turn feed upon a host and after some weeks moult and

1 The few facts known regarding the biology of these species are included under
Section I : savignyi (p. 42), pavimentosus (p. 62), lafioreitsis (p. 67).
'^ See feeding experiments in Cambridge, noted below.

N. I. 6

'341

82 Biology

appear as Adults: the female feeds more plentifully than the male
and increases in size, whereas the male may be mistaken for a second
nymphal form. Copulation was observed four times by Lounsbury, the
male inserting his mouthparts (not the palps) into the vulva— after the
manner observed in Ixodidae.

Feeding : the adults feed about once a month in hot weather, at
longer intervals when it is cool. No feeding may take place during the
winter. Experimenting with adults, Lounsbury found that females
were ready to feed after each batch of eggs had been laid, remaining
practically stationary prior to oviposition. Batches of eggs were laid
after each meal. He liberated the hungry ticks in the bird cages at
night. Few eggs were laid after the fourth feeding and fewer still
after the fifth and sixth. It took about 10 months to raise one lot
of ticks from egg-stage to egg-stage, thus completing the life-history.
Although under natural conditions they appear to feed only at night
(the larvae excepted), nymphs and adults have fed readily upon fowls
and pigeons etc. in the Laboratory at Cambridge when protected
from strong light. Oken (1818) also notes that they occasionally
attack man during the day-time in Persia, they are not, therefore,
" strictly nocturnal " as stated by Riley and Howard (1895). Lounsbury
(1903, p. 7) is correct in stating that persicus is "always uneasy in
strong light, and tends to crawl away from it." When disturbed, they
often sham death, lying still with their legs retracted.

The following observations were made by one of us (N.) in Cam-
bridge upon specimens received alive from India, S. Africa, Egypt and
N. and S. America :

Larvae were placed on fowls in cages and observed from day to day.

Experiment I. 20. iii. 1907 larvae placed on fowl, some attached themselves at

once.

23. „ „ larvae measure 1 mm. in length, appear as blueish

black points, somewhat rounded, hanging by
capitulum, like Ixodidae.

24. „ „ larvae measure over 1 mm., appear spherical, blackish.
25-26. „ „ an equal number dropped off each day (8 in all), they

appeared blackish, but flattened i, with hood pro-
jecting quite over the capitulum ; measured
2x1-5 mm.

27. „ „ one gorged larva found in cage.

13. v. 1907 Nymphs of first stage appeared.

1 Hooker (1908 a, p. 39) states that the larvae are globular in shape up to within a few
hours of their abandoning the host. They then become flattened like the adult ticks which

A. persicus

83

Experiment II. 4. iv. 1907 larvae placed on fowl.

9. iv. 1907 larvae (29) found gorged.

10. iv. 1907 larvae (17) found gorged.

2. vi. 1907 Nymphs of tirst stage appeared.

The larvae in both experiments remained attached 5 — 6 days and
first-stage nymphs issued after a further interval of 48 to 53 days, the
ticks being kept at about 20^ C.

First Stage Nymphs feed rapidly; thus 14 specimens were observed
to drop off gorged within 35 minutes. Second Stage Nymphs were on
one occasion observed to cast their skins 15 days after feeding, having
been kept at 24' C.

Second Stage Nymphs and Adults appear to feed still more rapidly,
but at times the process of feeding may be much prolonged, especially
if the ticks have grown feeble or been starved too long. Where they
gorge rapidly (in 5 — 10 minutes), they often discharge clear fluid from
the 1st inter-coxal space, as observed in Ornithodoros moubata, 0. savignyi,
etc. The following table contains a record of the time consumed in
feeding by all the ticks experimented with :

4 fed in 50 minutes

103

23

16 fed in 5

minutes

22 „ 10

9 „ 15

22 „ 20

20 „ 25

U4 „ 30

[8 „ 35

7 „ 40

.8 „ 45

22/3

3

„ 55

6

„ 60

1

„ 65

3

„ 75

2

„ 80

1

„ 90

1

„ 110

1

„ 120

From the foregoing, it appears that | of the ticks fed in half-an-hour
or under.

It was noted that

2 fed for 95 minutes, after starving 4 months
1 „ 90 „ „ „ over 5 „

1 „ 85 „ „ „ 10

permits them to "crawl rapidly and to secrete themselves in cracks and crevices protected
from the wily fowl." The replete larvae usuallj' drop at night when the fowls are roosting,
consequently when they emerge as nymphs they have no difficulty in finding birds upon
which to feed.

6—2

84 Biology

Nevertheless, they may feed more rapidly after fairly long starvation,
if they are in good condition. On the other hand, they appear to have
some difficulty, possibly a mechanical one, in feeding on mammals.
Thus of 7 hungry specimens placed on a rat, 4 fed for 2 hours and 3 for
4| hours, and 1, placed on a mouse (having starved 3 months), fed
for 5|- hours. A meal of blood does not always appear to agree
with them. We do not know upon what this depends. Thus
Lounsbury (1903) saw an adult turn black and die soon after having
sucked his blood, and Donitz (1907, p. 28) saw larvae which had fed on
white mice die rapidly after they had abandoned the host. Donitz
supposes that the death of his larvae may have been due to the blood
of the mouse being toxic for the tick, but he brings no proof in support
of the hypothesis.

Note relating to oviposition and hatching of larvae.

Female
fed on

Oviposition
began

Oviposition
ceased

Eggs kept at 24° C.
hatched out

Larvae
alive

23. i. 07

6. iii. 07

—

19. iii. 07

—

19. ii. 07

26. ii. 07

6. iii. 07

11. iii. 07

—

31. V. 07

18. vi. 07

—

—

7. ix. 07

19. vi. 07

25.vii.07

—

—

7. ix. 07

24. vi. 07

25.vii.07

—

—

1. ix. 07

30. i. 08

6. ii. 08

15. ii. 08

—

—

31. i. 08

6. ii. 08

15. ii. 08

—

—

3. ii. 08

11. ii. 08

18. ii. 08

—

—

From the foregoing, oviposition appears to last 7 to 9 days, and
larvae hatch out in 11 — 13 days, when the eggs are kept at 24° C.
The period when oviposition takes place after feeding was 6 — 8 days in
four cases, 18 — 42 days in four cases. Retarded oviposition may be
due to fertilization not having taken place.

Longevity {unfed) : Lounsbury observed unfed adults to survive in
captivity for a little over 2 years. He believes that under natural con-
ditions both larvae and nymphs can survive unfed for long periods.
Laboulbene (1881, and Laboulbene and Megnin, 1882, p. 337) observed
the survival of unfed specimens (from Persia) for over 3 years. Robertson
(1905, p. 561) kept adults unfed in pill-boxes for 2 years and 3 months,
nymphs survived 2 months without food. Borrel and Marchoux (1905,
p. 362) kept adults and nymphs alive for a year unfed and in a dry
place.

Resistance to immersion in fluids: they may, in our experience,
survive for over an hour in spirit. In von Loder's letter to Oken (1818,

A. persiciis 85

p. 1567), it is even stated that a specimen survived 24 hours' immersion
in spirit !

Seasonal Prevalence: Riley and Howard (1895, p. 2G7) state that
the plague of persicus continues through the winter in Texas, their
numbers being greatest in dry, hot years. Lounsbury (1903, pp. 9, 11)
found about Cape Town, that " a few specimens of all stages may be
found on almost any night in the year prowling about roosts in search
of hosts." Complaints of their ravages are, however, most frequent in
November and December. " This is undoubtedly because in these
months great numbers almost simultaneously take their first feed after
their long winter fast, and because of the brood of young ticks which
then appears."

Effects of the bite of Argas persicus on man and animals^.

Argas persicus has a rather formidable reputation in Persia. Dupre
(1819, pp. 323 — 324) seems to have been the first to write regarding it,
stating that its bite is at times dangerous, causing prolonged sickness ;
he speaks of it as a " teigne." Oken (1818, p. 1567) cites a letter from
v. Loder stating that the bite of persicus is fatal to man within 24
hours. He reports the case of an Englishman who kept a starved
specimen one year in a glass, and died 24 hours after he allowed it to
bite him. Kotzebue (1819, p. 180) says that it behaves like a bed-bug,
and may so infest villages as to drive out the inhabitants. The natives,
he relates, are comparatively immune, but foreigners suffer severe pain,
delirium and convulsions, and even death, within 24 hours in consequence
of its bite. Fischer de Waldheim (1823, pp. 269—273) also says that
the bite of this species may prove fatal owing to some complication.
Heller (1858), who examined their anatomy, denies that they have a
poison gland, and ascribes the effects to the mechanical injury (!) inflicted
by the parasite. Taschenberg (1873) thinks that the effects ascribed to
Argas persicus are really due to a fever which prevails in Miana ("dem
in Miana herrschenden Faulfieber "). Taschenberg (1874, p. 171) records
persicus as attacking man in Egypt as it does in Persia. Schlimmer
(1874), of Teheran, considers that the relative immunity of the natives
is acquired by their having been bitten at some time or other by the
Argas, and that such bites act like a preventive inoculation with vaccine
against smallpox. He says the symptoms are like those of " remittent

^ The portions dealing with the effects of the bite of Argasidae is reprinted from
Nuttall (1899, pp. 42—49) with additions to date.

86 Biology,

fever, extreme lassitude, disinclination to work, yawning, fever, per-
spiration, not accompanied by much thirst, increasing and decreasing at
stated hours in the day," so that many think it is only malaria acquired
during a short stay at Miana. Schlimmer does not share in this opinion,
and denies that the natives are subject to malaria. He says that
fatigued travellers, and those who have undergone privations, are
especially susceptible. A. 2')ersicus is also found at Chahroude and
Bestham on the main road from Teheran to Khoragan. No malaria
exists in these parts, but the effects of the Argas bite are the same as
at Miana. Schlimmer relates that he once (1858) treated 400 soldiers
who claimed to have been bitten by these parasites at Miana, but many
were unable to state on what part of their body they had been bitten.
The soldiers suffered from the symptoms above described, and were
promptly cured by the aid of " la poudre minerale de Bondin," or, when
the cases were refractory, by the administration of quinine. Bordier
(1882), who reprints the part of Schlimmer's publication from which
the above data are quoted, inclines towards the supposition that the
effects of the Argas bite are due to a poison, and, referring to the
reported immunity of the natives, says that this reminds him of the
fact that in many countries it is the strangers who are especially
attacked by mosquitoes, the natives having apparently acquired a
resistance towards the poison of these insects. Megnin (1882, p. 305)
denies the statement generally made by medical zoologists that the
bite of A. persicus is dangerous. He refers to a letter of Tholozan's to
Laboulbene (1881) which says that it is the belief among the common
people in Persia that the bite of Argas is dangerous and fatal to
foreigners, intermittent and remittent fevers being attributed to it.
Fumouze repeatedly placed a female Argas on a rabbit's ear from which
it sucked blood, but no pathogenic effects followed. Laboulbene and
Megnin (1882, p. 338) made a similar observation. Brandes (1897), in
view of his experience with A. reflexus cited later, considers that the
effects of the bite are due to a poison. Lounsbury (1900) at Cape
Colony, allowed himself and Davidson to be bitten by A. persicus, the
experiment gave rise to no further symptoms than a slight itching at
the seat of the bite ; the latter healed within 10 days in Davidson's
case and in 3 weeks in Lounsbury 's. There may well be some exag-
geration in earlier accounts regarding the effect of Argas bites, but
there is sufficient evidence to prove that they produce evil effects. It
has not been demonstrated that persicus conveys any infectious disease
to man as has been proved in the case of fowls.

A. 2)et'sicus 87

Economic Importance: the effect of the bites of A, j)ersicns on
fowls, when the birds are attacked by many ticks, may certainly be
serious. Thus Riley and Howard (1893, p. 267) published a note by
A. Turpe of Kinney Co., Texas, who states that these ticks kill pullets
" by creeping in masses under their wings, under their shoulders, and
actually suck their lives out until they die." Hoehr (1893, p. 348)
wrote from the same locality, stating that he had been acquainted with
the pest since 1888 and had seen 25 — 30 "/o of the chickens succumb in
a short time on a ranch. " They spread very rapidly. Last year I built
a new chicken house, but in 8 days it was literally full." Hoehr lays
stress upon its being the young Argas (larvae) that inflict the chief
injury. If numerous, they even cover the bird's skin, and kill off old as
well as young birds. Railliet (1895, p. 718) states that it kills off
fowls in Mauritius ("^. mauritianus"), and Osborn (1896, p. 256)
confirms the statements of the American observers. Salmon and Stiles
(1901, p. 405) report " We have seen one case from Florida in which a
chicken was literally covered with the hexapod stage," and one of us
has received a piece of a fowl's skin from Barbados, preserved in alcohol,
the skin being crowded with larvae oi i^ersicus in all stages of repletion.
Hart (1899, p. 180) of Trinidad, states that the infested birds sit down,
drop their wings, and show fever. Dressing with petroleum kills the
ticks and the birds recover. Fuller (1897, p. 590) of New South
Wales, states that fowls may die from exhaustion due to intense tick-
infestation. According to Lounsbury (ix. 1903, p. 11) the fowl tick "is
regarded by many as the most pernicious poultry pest that we have in
South Africa. It is directly responsible for the death of numerous fowls,
far more, indeed, than is generally suspected. It also seems to be the
worst of poultry parasites in many parts of Australia and in the
Southern States of America." Lounsbury, moreover, writes : " Paralysis
is commonly attributed to the attack of the ticks but I am not certain
that any disease is caused or transmitted by them at the Cape, and I
incline to the belief that the victims die entirely from the loss of blood
and the inflammation produced by the excessive parasitism. Vast
numbers of the creatures take their fill from the legs, and the after
irritation and the soreness probably account for the inability to walk
often noticed on the part of many fowls in a suffering flock ; and birds
thus lamed and thus made unable to mount the roosts at night become
doubly troubled by the pest. Young chickens, being also easily acces-
sible, suffer more than their share, and it is not uncommon to meet with
a whole brood almost black from the presence of adhering larvae. Sad

88 Biology

to relate even such destructive infestation often fails to attract the
attention of the owner to the pest. The losses are most striking when
a fowl house is first occupied after having stood empty for a few months,
as frequently happens in and about towns. The hungry ticks swarm to
the birds and may kill half or more of them before their presence is
discovered."

Reaney (1907), in Central India, saw fowls die within 24 — 48 hours
from intense infestation with persicus, after being placed in a fowl house
which had been disused for a fortnight. He attributed the death of the
birds to a poison emanating from the tick and noted that the bites
produced extensive extravasations of blood beneath the wings.

From the foregoing it appears reasonable to conclude that loersicus
may kill animals by massive infestation comparable to the "tick worry"
seen in cattle infested with Ixodidae. Apart from this there appears
to be evidence pointing to the ticks giving off some poisonous substance
into the wounds they inflict, and evidence in this respect will be also
- found in the notes relating to other Argasidae. Experiments conducted
in the laboratory at Cambridge have recently proved that the salivary
glands of A. -persicus contain a substance which retards or prevents
blood coagulation, i.e. anticoagulin (Nuttall and Strickland). Again it is
a not infrequent occurrence that tick bites become secondarily infected,
the wound serving as a point of entrance for pathogenic bacteria. Finally
we know that some Argasidae and Ixodidae may transmit certain
blood parasites to their hosts in the manner considered below for the
Argasidae.

Argas persicus in Relation to Disease.

Spirochaetosis in Foivls is a disease whose cause, the Spirochaeta
marchouxi^, was first discovered in Brazil by Marchoux and Salimbeni
(1903). The disease may be very fatal, since it is capable of destroying
all the fowls in a yard in the course of a few days. The disease begins
with diarrhoea, followed by loss of appetite, the birds appearing som-
nolent; the feathers being ruffled and the comb pale. The birds cease
to perch, lie down with the head resting upon the ground, and death
takes place during a convulsive attack. At times the disease runs a
slower course, the legs become paralysed, then the wings, and the bird
grows thin and dies in 8 — 15 days. Recovery may take place, but it
is rare after paralytic symptoms have appeared. At autopsy, during
the acute period of the disease, the spleen appears much enlarged and

1 Sp. marchouxi Nuttall, 1904, p. \& = Sp. galUnarum Blanchard, 1905.

A. persicus 89

the liver swollen with more or less fatty degeneration, at times the liver
is dotted with focal necroses. In chronic cases both of these organs
may appear atrophied. The blood is fluid and dark. Spirochaetes are
plentiful in the blood until shortly before death, and they disappear as
recovery sets in.

This disease is transmitted by Argas 'persicus {= miniatus), as was
proved by Marchoux and Salimbeni. By means of infected Argas
persicus sent to one of us (N.) in England by Drs Marchoux and Borrel,
of Paris, the disease was reproduced in fowls at Cambridge. Balfour
(1907) has observed the same disease in Anglo-Egyptian Soudan, and
has also transmitted it by means of infected A. persicus. Blood films
or infected ticks or both have been received from Dr Balfour (Khartoum),
Dr Bitter (Cairo), Captain Greig, I.M.S. (Punjab, India), and Dr Johnson
(Adelaide, S. Australia), from which it appears that the disease is very
widely distributed. Reaney (1907) has observed spirochaetosis in fowls
at Agar-Malwa, Central India, and also records the presence there of
persicus. He demonstrated that the ticks conveyed the disease.
Marchoux and Salimbeni (1903) and Borrel and Marchoux (1905)
found that when A. persicus sucks blood containing Sp. niarchouxi
the latter multiplies within the body of the tick (maintained at
30 — 85" C.) and it is capable of transmitting the disease for 6 months
or more afterwards if it bites a susceptible animal. When the ticks are
maintained at 15 — 20" C. after feeding upon infected blood they are not
capable of transmitting the spirochaetes to clean fowls. The spirochaetes
seem to disappear in ticks kept at a low temperature, but they reappear
if the ticks are placed at 30 — 35" C. although a period of 3 months may
have elapsed since the ticks fed upon infected blood. The spirochaetes
may be demonstrated in the coelomic fluid of the infected ticks by
cutting ofl" a leg and collecting the fluid on a slide. The spirochaetes
do not appear to injure the ticks. The disease usually breaks out 4 — 5
days after the ticks bite the birds.

Spirochaetosis in Geese was observed in the Transcaucasus by
Sakharotf (1891), of Tiflis. The disease is as fatal as Spirochaetosis
in fowls, and it is probable that it is transmitted by A. persicus,
since this tick occurs in Southern Russia.

Note: Proof is still lacking that A. persicus plays a part in human
pathology as a carrier of infection, or that the fever attributed to its
bite in Persia is relapsing fever. Manson (1908, p. 196) states that
" miana fever " is " certainly communicated " hy A. persicus, but there is
no scientific evidence in support of the statement.

90 Biology

Destruction of Argas persicus.

We have referred to the fact that the Persians migrate from their
villages or burn them when the tick grows too aggressive (Oken, 1818,
p. 1569, etc.).

With regard to fowls, Riley and Howard (1893, p. 267) have recom-
mended spraying the hen-coops and poultry yard and washing the
poultry with kerosene-emulsion. Hoehr (1893), on the other hand, did
not find the emulsion satisfactory and obtained better results with lime
and sublimate. Ehrhorn (published by Packard, 1895, p. 418) reported
that a spray of " creozozone " instantly killed the ticks and gave good
results. Riley and Howard (1895, p. 348) state that oil of sassafras
kills the ticks quickly but is dangerous when applied to young birds.
Fuller (1897, p. 590) recommends white-wash, and scalding the fowl's
nests, boiling water being effective. The ticks are killed by turpentine.
Lounsbury (ix. 1903, p. 12) discusses the value of various remedies and
describes an instance in which the following procedure was successfally
adopted :

All the old roosts and nests were burnt, the wooden walls of the
fowl-house were well brushed over with hot coal-tar. New roosts were
suspended by wires from the roof so that they did not touch the walls
(into the crevices of which ticks retreat ; poles with bark on them
should not be used for roosts for the same reason). Paraffin was freely
applied to nests when ticks were encountered in them, the floor was
regularly swept clean and sprinkled with wood ashes and lime. In
other cases frequent spraying of fowl-houses with various sheep-dips
proved effective. The ends of the roosts may be wrapped around with
oiled waste or be insulated in a deterrent fluid. Corrugated, or better,
sheet iron fowl-houses have the advantage of being readily cleaned if
any ticks are about ; they may be tarred inside. Loose bark on trees
in fowl-runs and wooden fences are also hiding places for ticks.
Instead of dipping or treating young birds, Lounsbury recommends
isolating them for some days in crates when any young ticks on them
drop off and may be destroyed by burning the crates.

A. reffexus 91

ARGAS REFLEXUS.

Longevity (unfed): Hermann (1804, p. 70) fed a specimen and then
kept it unfed in a glass in which it survived 8 months. Bianconi (1867)
reports that it survived in a disused hen-coop for fully 8 months.
Gulliver (1872) states that specimens survived 5 months unfed in a
tin box, and FuUager (1874, p. 121) kept them alive 22 months in a
glass-topped box ; unfed larvae lived 6 months. He was informed by
Austin that the latter saw them (later stages of the tick) survive 4 — 5
years unfed in a box in which they had been placed and forgotten.
Berlese kept an unfed specimen alive for about 6 years in a corked
bottle. Ghiliani informed Perroncito (1901, p. 568) that he had seen
unfed specimens survive for 22 months. Brandes (1897) states that
living sjDecimens were found in an abandoned pigeon-coop after the
lapse of 2 years.

Seasonal Prevalence: Schellack (1908, p. 487) in Magdeburg,
Germany, found rejiexus in fairly large numbers in autumn, but they
were scarce in March.

Feeding Habits, Economic Imjjortance, etc.: Taschenberg (1880,
p. 153) states that it remains motionless during the day-time or when
exposed to lamplight, this being in accord with the statements of other
writers, A few observers record the time occupied by the tick in
feeding (meaning nymphs and adults); thus Alt (1892) saw them feed
20 minutes on man, and Boschulte (1860), who allowed himself to be
bitten by a specimen, states that it took 27 minutes to feed. Obviously,
in this respect its habits are similar to those of A. persicus. The larvae
are stated by Braun (1895) to remain "some oime " on their hosts, from
which we may gather that they stay attached for some days as do
A. persicus larvae (q.v.). According to Perroncito (1901), the larvae
(we assume) occurred in large numbers on the skin of the young
pigeons he saw succumb to their attacks. Bianconi (1867) placed
4 pigeons in a reflexus-inksted hen-coop, which had been disused for
8 months. Two of the pigeons (young birds) died the first day, and
the other two (adults) died on the third and fourth days respectively.
The pigeons were literally covered by the ticks, and Bianconi attributed
their death to exhaustion and loss of blood. Fowls similarly placed did
not appear to suffer. The injury they inflict on pigeons are also noted
by Megnin, 1880 ; Laboulbene and Megnin, 1882 ; Railliet, 1895 j

92 Biology

Osbom, 1896 ; Brandes, 1897. Young birds are the chief sufferers,
and at times it is impossible to raise them owing to the pest.
Schellack (1908, p. 487) writes that reflexus attacks pigeons especially
on naked parts of the skin, beneath the wings and about the anus.

Effects of the bite of Argas reflexus on man and animals.

When numerous, they may, through their bites, cause the death of
pigeons. They have been observed to wander into chicken-houses and
dwellings. They do not seem to annoy chickens, but they occasionally
attack man and cause much trouble. Raspail (1839) attributed a severe
erythematous eruption on a child's neck to the bites of this species,
consequent on his going into a pigeon-coop. Boschulte (1860) describes
the case of a family, several members of which were bitten by reflexus,
only pain and slight swelling following in all cases excepting that of an
old man. The latter was bitten on the lower part of the thigh, with
the result that a deep circular suppurating wound about the size of the
head of a pin marked the spot where he had been bitten. There was
extensive oedematous swelling and redness of the surrounding parts.
Boschulte allowed himself to be bitten by reflexus. The pain was like
that of a mosquito-bite. A small drop of coagulated blood subsequently
covered the puncture. Nothing especial was noticed, and three days
later the wound had healed. Ten days after he had been bitten the
spot began to itch and showed a nodular swelling, which grew red and
increased to the size of a pock. No exudation of serum occurred, but
the itching was very annoying. This subsided after six days, a small
scab was cast off at the point bitten, and the skin resumed its normal
appearance. Boschulte (1879) reported, nearly twenty years later, that
the place where he had been bitten still showed a sharply-defined
circular flattened elevation with a central cicatrix, and that in the
interim several similar, but smaller, elevations had appeared in its
vicinity.

Taschenberg (1873) wrote that reflexus attacked some children in
Friedeberg a. d. Saale. In a later paper, Taschenberg (1880, p. 153)
states that in all cases where reflexus has attacked human beings, the
latter slept in rooms adjoining pigeon-coops. The bites occur chiefly
on the hands and feet and appear as small red points which cause much
itching, extending up the limb. A bite on the hand produced itching
up to the shoulder, a bite on the foot itching up to the hip and back,
there being less irritation about the bite itself Scratching aggravates

A. reflexus 93

the trouble especially in children, where there may be inflammation.
In a girl 5 years old, blisters formed over the hand, wrist and forearm.
The itching may last 8 days. Chatelin (1882) reports the case of a
child that was bitten by reflexus, which had wandered from the pigeon-
house into the dwelling. The pigeon-house had not been used for
6 years. The bites were followed by pain and oedematous swelling,
which j^ersisted for some time. Other persons who were bitten at the
same time exhibited no such symptoms. Brandes (1892 d, p. 10)
describes the case of a man who was bitten at Aschersleben in 1883,
the bite being followed by much swelling. Alt (1892) saw a case
which occurred under similar circumstances, where the bite was
followed by urticaria factitia and general erythema, which subsided
in a few hours. Brandes (1897) also describes this case — that of a man
who had been bitten five times in four years. Hauch, who attended
him, stated that he woke at night with pain about the wrist, on which
he discovered the Argas. Within half an hour an erysipelatoid swelling
spread from point of the puncture all over the body, increasing, par-
ticularly about the head, until the eyes were hidden by the swollen
lids. During this time the patient suffered from shortness of breath,
palpitation, dulness, etc. for an hour, when the symptoms began to
subside with the appearance of profuse perspiration. The swelling
gradually subsided during the following 10 to 15 hours. The patient,
who seems to have been particularly susceptible to the bite of the
Argas, had previously kept pigeons in his house, but the pigeon-house
had been walled up two years before. As Brandes states, this latter
proceeding seems to have caused the migration of the parasites into
the dwelling. Alt (1892), and two other persons, allowed themselves
to be bitten by j'eflexus obtained from the abandoned pigeon-house.
Slight pain, that came and went, followed, but nothing in particular
occurred, excepting in one case, where, after four to five days, a painful
nodule, the size of a pea, appeared at the seat of the puncture, but this
disappeared soon afterwards. Two persons who suffered from urticaria
also allowed themselves to be bitten ; one of them remained unaffected,
whilst the other developed general erythema after four hours, which
subsided again in an hour. Brandes reports a case which was observed
in 1884 at Aschersleben, where a man became so oedematous after four
to five hours that his clothes had to be cut off". The oedema is said to
have lasted three days in this case. The effects here noted seem to
depend on a peculiar idiosyncrasy. Brandes believes that a poison is
probably elaborated in the salivary glands of Argas. Alt, who injected

94 Biology

three of them, which had been crushed, subcutaneously into a dog,
produced symptoms of intoxication in the latter which were similar to
those which are produced by small quantities of snake ("Puffotter")
venom.

Terrenzi (1893, pp. 73 — 76) observed the effects of the bite on
the hand as follows : After a few hours a yellowish vesicle appeared,
and lymphangitis extended up the forearm, the course of the lymphatics
appearing red and feeling hard and the skin rough (scabrosa) ; when
morning had arrived the vesicle had grown five times in size and the
lymphangitis worse, and the arm discoloured. The effects diminished
and disappeared in 15 days, but a scar was visible a long time after.
Gibert (1896, cited by Mosler and Peiper, 1904, p. 345) observed
general symptoms follow the bite: nausea, vomiting, diarrhoea, irregular
pulse, dyspnoea, etc. Ajutolo (1898) states it tormented persons in
Bologna, and refers to it as " this terrible destroyer of poor pigeons."

Argas reflexus in Relation to Disease.

Until quite recently there has been no evidence to prove that
reflexus is capable of conveying disease. Schellack (1908, p. 487)
however, reports upon two experiments which he carried out with
A. reflexus and Spirochaeta marcliouxi. Experiment I: 4 reflexus
were placed on a fowl with spirochaetes in its blood, and 7 days
later they were placed on a clean fowl. Spirochaetes appeai'ed in this
fowl's blood after 16 days and it died of the disease. Experiment II :
6 infected reflexus were allowed to bite a clean fowl ; the bird showed
spirochaetes on the 8th day and it recovered; the ticks had fed on a
sick fowl about 64 days before. It is interesting to note, therefore,
that Sf. marchouoci may be transmitted by two species of Argas
{persicus and reflexus) as well as by Ornithodoros moubata (q.v.).

That this tick may transmit other pathogenic micro-organisms is
indicated by the following instance reported by Tonnel (1906, p. 552) :
A family moved into an apartment which had remained unused for four
years having been previously occupied by a man who kept fowls and
pigeons in the rooms. The father of the family had suffered from
general furunculosis before moving into the dwelling. He was bitten
by reflexus and so were his two children, who in consequence also
developed abscesses and phlegmons starting at the points bitten by the
ticks. The family moved out of the apartment because they considered
it bug-infested (they took reflexus for bed-bugs), after which the trouble
subsided. Within 48 hours another family moved into the apartment.

A. hriimpti 95

with the result that two members of this family (father and child)
suffered from the same effects as did the previous family. On searching
the apartment rejiexus were discovered, and, following a thorough
disinfection of the premises the trouble ceased. It appears clearly
demonstrated by thi^ interesting case that reflexus may transmit
pyogenic bacteria to healthy persons when they have previously had
access to the skin of a person suffering from furunculosis.

ARGAS BRUMPTI.

Effects of Bite : Dr Brumpt communicated the following particulars
to Professor Neumann (1907, p. 229) and to us :

The bite is somewhat more painful than that of 0. mouhata ; it ma}^
wake the sleeper and causes pruritus, lasting several days. The puncture
bleeds slightly and becomes covered by a small crust of dried blood
beneath which a small drop of extravasated blood gathers. Within
about 2 hours the puncture becomes surrounded by a violet-coloured
circular ecchymosis 6 — 8 cm. in diameter resembling the mark follow-
ing a leech-bite. The mark goes through the gradations of colour
usual in ecchymoses, or bruises, and disappears in 6 — 8 days ; but the
centre remains indurated for a long time. Of 17 bites which were
inflicted on Dr Brumpt in July 1901, five still exhibited an indurated
centre in April 1908 (Letter to G. H. F. N.). He states that the small
subcutaneous nodules may be felt on palpation and seem to be dis-
appearing with the exception of one in the umbilical region, this nodule
measuring 7 mm. in diameter. From time to time he still suffers from
pruritus at the seat of these bites ; one bite was inflicted on the neck,
and this place is particularly subject to pruritus, owing, apparently, to
the friction with the collar. All traces of the other 12 bites have
disappeared.

Habitat: Dr Brumpt (9. v. 1908) informs us that he found
A. brumpti in dusty sheltered hollowed out places beneath over-
hanging calcareous rocks, into which opened porcupine {Hystrix sp.)
burrows. The ledges of rock were situated along the rocky bed of a
torrent, and the floor of the hollows, which were about 5 ft. deep,
consisted of fine dust about 15 cm. in depth, numerous bones of small
mammalia lying upon it. After A. brumpti had fed it promptly buried
itself in the dust. The ticks wander about at night and Dr Brumpt
was unable to catch any during the daytime.

96 Biology

ORNITHODOROS MOUBATA.

Period required for the Egg to hatch: eggs kept at 29° C. (Newstead,
1905, p. 1697) became flattened on the 6th day and the legs and
capitulum of the larva could be seen through the shell ; on the 9th day
the shell split and the larva was fully formed ; on the 10 — 15th day the
young nymph hatched out and on the 16th day the nymphs were
observed to escape anteriorly from the larval exuviae. Some hatched
out in 8 — 13 days. Wellman (1906-7) observed hatching after 15 days
in Angola. Dutton and Todd (1905, p. 124) observed hatching after
18 — 23 days in the Congo (temperature 19'5 — 32° C), whereas two or
three days more were required when the eggs were kept at 19 — 22° C.
(Laboratory experiments in Liverpool ; Newstead, Dutton and Todd,
1907.)

Oviposition: as in other Argasidae we have studied (A. persicus,
0. savignyi, 0. lahorensis) the eggs are laid in batches, the eggs being
agglutinated in masses and deposited upon the sand or in hollows
burrowed out by the female. Dutton and Todd (1905 b, p. 125)
note that the number of eggs is increased when the female has
taken a large meal. They observed females which laid batches of
10 — 20 eggs at intervals of 1 — 2 weeks, although they were frequently
disturbed. The largest total number of eggs laid by one female was
139. Newstead (1905, p. 1697) placed a female in a bottle with sand in
which she rapidly buried herself after feeding. Placed at 29° C. in a
dry incubator, she laid three batches of eggs, oviposition taking place at
night. She rested upon the successive batches of eggs for a time ; the
batches numbered 17, 51, 26 (total 94), intervals of 3 and 8 days
elapsing between the layings. Wellman (1906) observed oviposition
in two females which were kept unfed in a dish with floor dust at the
bottom : Tick I oviposited after 57 days, laying 49 eggs ; she was
removed from these eggs and laid 29 more by the next day. Tick II
oviposited after 64 days, and laid three batches of eggs totalling 88.
According to Wellman (in MS.) a fecundated female lays no eggs until
she has had a meal of blood. Mollers (1907) saw a female lay 80 eggs,
she buried herself and the eggs were pushed out so that they appeared
upon the surface of the sand.

Longevity : under natural conditions 0. moubata doubtless may live
for several years. Specimens have been kept alive, though unfed, for

0. moiibata 97

4 — 6 months or more by Newstead, Button and Todd, the ticks, never-
theless, being capable afterwards of infecting animals with the Spirochaeta
duttoni. Unfed specimens have been kept alive for similar periods in
Cambridge. Wellman (in MS.) saw females survive unfed for 4 to
nearly 6 months after ovipositing and Mollers (1907, p. 278) states that
adults may survive unfed for a year. Although fed regularly in
captivity adults were seen by Mollers to gradually die off after the
lapse of 2 years.

Natural enemies: as Livingstone (1857, p. 382) wrote: they are
difficult to kill, " their skin is so tough and yielding, that it is im-
possible to burst it by any amount of squeezing with the fingers."
This, combined with their colour and life habits (hiding in cracks and
burying themselves in sand or dust), unquestionably affords them
much protection. Nevertheless, under natural conditions in the Congo,
Button and Todd (1905 b, p. 127) and Wellman (in MS.) state that
they are devoured by chickens, rats and mice, and that ants carry off
young ticks ^ and eggs. Wellman (1906 and 1907) has seen Phonergates
hicoloripes Stal. attack and suck the blood out of 0. vioubata, and has
figured the manner in which the bug seizes upon its prey. He has sent
specimens of this tick-enemy to the British Museum (see Austen, 1906,
p. 113) and to Cambridge. Button and Todd (1905 b) noticed, as have
others, that when disturbed they often curl up their legs as if dead.
" So lifeless do they seem that one might easily be deceived, especially
since they sometimes lie motionless for hours." This habit doubtless
affords them protection against enemies. Wellman (1906) states that
in the Bihe Bistrict, Angola, he observed what appeared to be their
destruction by a parasitic mould.

Feeding: Button and Todd (1905b, p. 124) state that a large ?
may remain attached to a monkey for 2 — 3 hours, others feed for half
an hour. In feeding, the tick braces itself on the forelegs, depresses the
capitulum and bores in its mouthparts. It may expel faecal matter,
and it exudes clear fluid, in fairly large amount, from the 1st intercoxal
space whilst attached to the host or after it has dropped ofi". According
to Newstead, Button and Todd (1907) they will not feed well more
often than every 7 — 10 days. Newstead weighed a female before and
after feeding and notes that her weight increased ten times. Mollers
(1907, p. 278) states that mouhata feeds for ^ to 4 hours. Fed
on laboratory animals in Cambridge during the day-time, the adults

' Wellman (in ms. ) once saw a swarm of driver ants {Doryliis nigricans lUiger) bearing
away mouhata from a native kraal in Angola.

N. I. 7

^8 Biology

and nymphs were observed to feed for very variable periods; thus 15
ticks, of which accurate records were kept, fed respectively for 20,
20, 25, 30, 30, 40, 40, 40, 45, 55, 60, 80, 100, 100 and 125 minutes.
Nymphs (1st stage), of which 5 were timed whilst feeding, took 10,
10, 25, 30 and 60 minutes before they dropped off the host. They
usually feed at night under natural conditions, but they may attack
persons by day as Wellman has observed in native huts in Angola
(personal communication).

The effect of the bite has been repeatedly described :

Livingstone (1857, p. 383), who was bitten by the " tampan," states
that the tingling sensation at the point bitten lasted for about a week.
He refers to the fever which may follow the bite as being well-known to
the natives. Murray (1877, p. 182) quotes Dr Welwitsch as stating
" that the pain of the bite is not felt until two hours after it has been
inflicted, but it makes up for the respite by continuing painful and in-
flamed for from 12 to 24 hours thereafter."

Brumpt (1901, pp. 578 — 580) was the first to study the effects of
their bites, allowing himself to be bitten by 43 moubata collected in
Somaliland, where "tick fever" prevailed. He suffered no ill effects and
states that the bites were not troublesome except when they attack man
in large numbers, producing anaemia consequent upon loss of blood.

Dutton and Todd (1905 b, p. 123) say that the bite even of a small
tick is painful; they allowed a tick to bite a monkey: "Immediately after
feeding, a small crust of serosanguinolent fluid forms at the site of the
bite. Surrounding it is a roseola about 2 mm. in width. Two hours
later the central clot is surrounded by 2 concentric zones, each 2 mm. in
width ; the first colourless, the second ecchymotic. Six hours later the
clot has become almost black, and is placed at the apex of a slight,
colourless wheal, bordered by an ecchymotic zone about 1"5 mm. in
width."

Wellman (1906, 1907) writing from personal experience in Angola,
states that the bite is very painful, the swelling and irritation (especially
in Europeans) not subsiding for days. "The wheals are hard, raised, and
itch and swell most disagreeably if scratched, and this even a week
after being bitten. The bite of young ticks (nymphae) is said by
the natives to be more severe than that of the adults." Wellman
experimented upon himself and is convinced that this is generally
the case. He states (in MS.) that in some natives, usually aged
individuals, the bite may scarcely leave a mark. (From this it is
evident, as is the case with mosquito bites, that immunity to its effects
is acquired.)

0. moiihata 99

Treatment of Bites : Wellman (in MS.) recommends prolonged bathing
in very hot water, followed by the application of a strong solution of
bicarbonate of soda, which is allowed to dry upon the skin. He states
that this treatment is comforting. For severe itching he advises
smearing the bites with vaseline, which is slightly impregnated with
camphor or menthol. Medical aid should be sought when complications
arise.

Prevention against being bitten : Livingstone (1857, p. 628) noted
the danger of sleeping in native huts at Tete and (p. 382) wrote : " I
had felt the effects of its bite in former years, and eschewed all native
huts ever after." He was, nevertheless, bitten at Ambaca in a European
house. Again he states (1874, p. 33) " the human tick, which infests all
Arab and Suaheli houses," and, writing at Nyangwe, the ticks " that
follow wherever Arabs go, made me miserable, but the Arabs are in-
sensible to them ; Abed alone had a mosquito curtain, and he never
could praise it enough."

Manson (1903, p. 714) writes that the natives in some districts
" protect themselves against the tick by plastering the walls and floors
of their huts with mud and cow-dung ; a practice adopted by the Boers,
the Bechuanas and nearly all the cow-keeping native tribes. They
frequently smoke their huts to drive the ticks from their lodgement in
the thatch."

The Portuguese always warn newcomers not to place beds on the
ground and to search their mosquito nets before retiring at night.

Wellman (1906 and 1907) says the natives of Angola are at times
forced to burn their huts to get rid of these ticks. The use of insecticide
powders (Pyrethrum) has been recommended.

Wellman reports to the Government of Angola four recommendations,
of which we quote three :

" (1) The tick in question should be regularly destroyed in crowded
centres by disinfecting native houses, barracks and other permanent
quarters, and by burning old camps, huts, etc.

" (2) Soldiers, labourers on plantations, etc., should be made to keep
their houses clean, and to sleep in hammocks or in beds well raised from
the floor and away from the wall. Natives should never be allowed to
sleep in or near the quarters of Europeans.

"(3) Soldiers, porters, servants, plantation labourers, and other
controllable bodies of natives should be compelled to observe regulations
regarding regular bathing and washing of clothes."

Naturally whites should avoid sleeping in or near native kraals and

100 Biology

servants' quarters should be established at a distance from the white
lines. In travelling, old camp sites and resting places should as far as
possible be avoided. " Native servants who are allowed to enter sleeping
apartments of whites should be compelled to change their clothing on
coming from native quarters." (Wellman MS.)

Ornithodoros mouhata in Relation to Disease.

African Relapsing Fever in Man or " tick fever " occurs in German
and British East Africa, in Central Africa, in the Congo Free State and
Angola. The disease was referred to by Livingstone, and it has been
mentioned by various travellers in Africa ^ The main symptoms are
headache (especially at the back of the head), vomiting, abdominal pain
and purging, with severe fever, a pulse of 90 — 120, dry hot skin,
congested eyes and shortness of breath. After a period of fever lasting
about two days, there is a fall of temperature, but a fresh attack soon
follows. These relapses occur more frequently than in European
relapsing fever, being usually 5 — 6 in number, but there may be
more. The attacks leave the patient in a weak condition for a long
time after recovery which usually follows, but death occurs in about 67o
of the cases. The fever attacks are due to protozoal parasites, the
Spirochaeta duttoni, which multiply in the blood, the greatest number
being present in the blood during the attacks of fever. The disease
lasts 1 — 3 weeks or longer, depending upon the number of relapses.

The Spirochaeta duttoni is transmitted to man by the bites of
infected 0. mouhata, as has been proved by scientific experiments. The
natives of parts of Africa (Angola, Congo, Uganda, Abyssinia, Somali-
land, German E. Africa) have for a long time attributed the fever to the
bites of this tick. In describing the effect of the tick's bite, according
to Christy (1903, p. 187), the natives in Uganda " invariably go through
a pantomime indicative of vomiting, with pain in the head and abdominal
region,"

The attack of fever usually follows 5 — 10 days after the susceptible
person has been bitten. The Spirochaeta duttoni was discovered by
A. R, Cook (Jan, 1904; this author took it for Sp. recurrentis) and by
Philip Ross and Milne (Nov. 1904, in Uganda), and the part played
by the tick was demonstrated by Dutton and Todd (Feb. 1905, in
the Congo), and subsequently by R, Koch (Nov. 1905, in German East

1 See Hinde (1897, p. 3), Plehn (1902), etc., who state that the disease is referred to
tick-bites by the natives.

0. moubata 101

Africa)^ When a female 0. muiihata sucks blood containing Sp.
duttoni, the latter pass into the ovaries of the tick and penetrate the
undeveloped eggs, within which they multiply. They persist in the
tick which developes from the egg and pass out of its mouthparts when
it feeds in the 1st nymphal stage upon the blood of a fresh host.
Monkeys and rats in England have thus been infected with the disease
through the agency of infected ticks brought from Africa. Dutton and
Todd, in addition, found that the spirochaetes persist in the gut of the
tick up to 5 weeks after it has ft>d. The tick, once infected, may harbour
the parasite for months and transmit it when it has occasion to feed.
Finally, MoUers (1907, p. 277) finds that the spirochaete is transmitted
to the third generation of ticks, the second generation having been fed
on blood free from spirochaetes; such ticks may infect animals (rats,
monkeys) by their bites.

Filariasis in Man: Christy (1903, p. 187) considered that 0.
monhata is capable of transmitting Filaria perstans to man. In this
disease the filarial embryos circulate in the blood. Feldmann (190.5,
p. 64), whose statements have been criticised by Kerr (1905, p. 126),
advanced the extraordinary hypothesis that the ticks infected with
filariae lay their eggs in bananas stored in native huts and in some way
give off the worms which are eaten with the bananas by the natives.
Wellman (1907 and MS.) states that he has observed a certain degree of
development of F. perstans embryos in moubata. His results are very
suggestive, since he worked with moubata which he raised from the egg.
The matter requires further investigation.

Spirochaetosis in Foiuls. It is interesting to note that Fiilleborn and
Mayer (1908, p. 31) have found that they could transmit Spirochaeta
marchouxi (see p. 88) by means of 0. moubata in experiments con-
ducted in Hamburg. In the positive experiments which they report,
the ticks had fed twice before upon infected fowls. The ticks were
infective for 103 days after feeding on a fowl harbouring the spirochaetes
in its blood.

Brumpt (1901, p. 578) observed that the parasites of tertian malaria
degenerated inside the gut of moubata and ticks fed on himself after
feeding on malarial blood produced no ill effects.

1 Massey(1905, p. 225) and Wellman (1905, p. 97) also observed the spirochaetes in
Angola. The epidemiology and history of the disease in German E. Africa are discussed
by Werner (1906, p. 776).

102 Biology

ORNITHODOROS CORIACEUS.

Effects of bite and feeding habits : two females bit Mrs Z. Nuttall
through her clothing and inflicted painful wounds, " their bites were
intolerably sharp and painful, and both wounds bled a good deal — but
notwithstanding, there has been intermittent irritation ever since"
(this persisted after 4 months, and the seat of the bite was still dis-
coloured and the puncture covered by a scab). Eight months after
the bite was inflicted there remained a nodule which occasionally
itched. The natives of Tehuantepec, Mexico, fear this tick for the
reason that the bites are severe and often do not heal for a long
time. The females immediately proceeded to feed, on arrival in
Cambridge, when placed upon a fowl. They fed for 45 minutes and
1 hour 40 minutes respectively, and drew a large amount of blood. The
bites caused intense ecchymosis, measuring about one inch in diameter.
Whilst feeding the palps did not penetrate the wound as once observed
in the case of 0. savignyi, but both specimens exuded clear fluid as
observed in 0. moubata.

ORNITHODOROS TURICATA.

Effects of bite : the " turicata," as the Mexicans call it, may cause
serious injury by its bite. According to Duges (1876) it has been
known to be fatal to pigs. This author also states that chickens fed on
turicatas died about the third day. The effect of the bite in man is
especially bad if the turicata's capitulum is torn off, and, where this
occurs, Duges recommends the use of the cautery, otherwise it causes
severe itching, and an ulcer forms at the spot bitten, and this may persist
for months, or there may develop erysipelatoid dermatitis, lymphangitis,
the formation of bullae containing serum about the puncture, at times
gangrene, subcutaneous abscesses, etc. In three cases he reports general
symptoms following the bite. In two of these a vein had been punctured
by a turicata. One patient had difficulty in speaking and swallowing,
swelling and numbness spreading over the whole body, accompanied by
vomiting and diarrhoea. In another patient all these symptoms subsided
within an hour, when an urticaria made its appearance, accompanied
by profuse perspiration. Duges says people are reported as having died
from the bites of turicatas, the noxious effects of which he attributes to
a venom, a peculiar idiosyncrasy existing in certain individuals.

Biology ' 103

ORNITHODOROS TALAJE.

Habitat, Effect of bite, etc. : Salle (1849, p. 342) and his companion,
Jules, were severely bitten by 0. talaje in May, 1847, at Casa Vieja de
Gastoya. They were awakened out of sound sleep by "atrocious itching
on the hands and face," and on lighting a candle found their hands were
" covered with blood and blotches like large bites of bugs." The mule-
teer said the bites were due to "talajas." The ticks infest old houses,
retreating into the crevices of the walls, which are built of bamboo and
covered with mortar. The talajas bite at night and disappear by
morning. Salle states "my hands and ears were much swollen, and I
suffered horribly " ; a fortnight elapsed before he recovered from the
effects. 0. talaje, according to Guerin-Meneville (1849), causes intoler-
able itching and pain by its bite. Megnin (1885) says its saliva may
be venomous like that of a mosquito or tarantula. This stands in
direct contradiction to his previously expressed views regarding A.
persicus (q.v.).

ORNITHODOROS THOLOZANI.

Effects of bite: Megnin (1882, and 1892, p. 66) claimed that the
bite of this tick is harmless. He allowed one which had starved
for years to bite his hand. It sucked itself full in about half an hour,
the pain produced being less than that of a leech. The only effect was
the formation of a violet ecchymosis 6 mm. in diameter about the bite.
As Johannessen (1885, p. 347) very properly remarks, one experiment by
Megnin (in France) with a tick which had been kept starving for years,
has no value as proving that its bite is innocuous under normal
conditions.

ORNITHODOROS MEGNINI.

The life history of this species has been recently studied by Hooker
(1908 a, pp. 40, 42, 45, 51) who placed bags over the ears of infested
cattle and safeguarded the bags by cords tied about the horns so as to
prevent their being displaced. The larvae, having gained entrance to
the ear of the host, attach themselves deep down in the folds of the skin
and gorge themselves. They moult upon the host after about 5 days
and the nymphs continue feeding sometimes for months. In one case
a nymph abandoned the host's ear 35 days after the larva had been

104 Biology

introduced, in other cases the nymphs still remained attached after
98 days had elapsed. "After leaving the ears as nymphs, these ticks
usually crawl up several feet from the ground and secrete themselves in
cracks and crevices, where in about 7 days in September, after leaving
the ear, they shed a membranous skin and appear as adults without
spines." Fertilization then takes place and oviposition commences, after
which the female dies. Eggs are not laid by unfertilized females and
the latter may live a long time. In summer the larvae hatch out after
11 days.

As far as we know the life history of megnini is unique amongst the
Ixodoidea since but one moult (larva to nymph) takes place upon the
host and the nymph stores up enough food to make it unnecessary for the
adult to feed before fertilization and oviposition take place. Hooker
believes that the adults probably never feed, and we would note that
this view gains support from the fact that the adult capitulum (Fig. 103)
is very small and that the hypostome (see p. 73) is unarmed, no similar
structure being known to us in other ticks. Even the structure of the
digit appears modified in that the external article does not bear the
usual outwardly directed teeth (see Fig. 104, digit of female). The
peculiar habit of the replete nymph of creeping upward several feet
from the ground before moulting appears to Hooker to be correlated
with the tick's parasitic habits, for when the adults mate and the
females oviposit, the larvae which issue from the eggs are placed in an
advantageous position where they can readily gain access to the ears of
their hosts.

Injurious Effects: Salmon and Stiles (1901, pp. 413, 414) write:
"Judging from letters received by this bureau (The Bureau of Animal
Industry, Washington, D.C.), the ear tick is accused of causing a variety
of troubles. Some correspondents report sickness and even death among
cattle as having been caused by the parasite, while others are not
inclined to attach so much importance to its presence. Owing to their
position in the ear, it is not possible to use any very drastic measures
against them, but, as a rule, if any bland oil, such as linseed or olive oil,
is poured into the ear, the ticks will soon vacate ; they are not killed by
this treatment, hence every tick caught should be crushed in order to
prevent increase." Simpson (1901) readily removed a nymph from the
ear of a gentleman by introducing a pledget of cotton containing a little
chloroform into the ear.

Longevity {unfed): Megnin (1885) states that he kept some 0.
megnini alive unfed for two years.

BIBLIOGRAPHY

The titles (about 300) given in the following bibliography refer to the Argasidae.
Throughout the book the Harvard System of References to the bibliography has
been adopted. Wherever an author has been cited in the text the date of publication
(and usually the page) has been added in brackets. In the following bibliography
the names of the authors are given in alphabetical order, the names being followed
by the date of publication, the title of the paper and the Journal, etc. where the
paper was published. The Roman numerals after the name of the Journal or book
indicate the number of the volume, and the figures printed in ordinary type refer to
the page. In some cases a short comment upon the contents of the publication
accompanies the reference. Unless otherwise stated the quotations are from the
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Ixodidae.)

4

I

Bihliograpliif s

Austen, E. E. (1906), An insect enemy of the disseminator of human tick fever in
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Banks, N. (1901a), Acarina in "Some Spiders and other Arachnida from Southern
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(1901b), Some Arachnida from New Mexico. Proc. Acad. Nat. Set., Phil-
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(1902b), Papers from the Hopkins-Stanford Galapagos Expedition, 1898-1899,

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(1904 b), A treatise on the Acarina or Mites. Proc. U.S. National Mus., xxviii.

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(1889), Acari africani tres illustrati. Atti d. Soc. veneto-trentina di sc. naturali,

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(1890), Traite de la Zoologie mcdicale, ll. 883, Figs. 388-883. (Refers to

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8*— 2

Bihliography 5

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BUMof/raphy 7

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Bihliof/raphy 9

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BihliodmpJni 11

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Bibliof/raiyhi/ 13

Gervais, P. (1844), Acferes Phryneides, Scorpionides, Solpugides, Phalangides et
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i

Bibliofjraphy 15

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I-XXX.

Bihliofirapliy 1 7

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Kiichenmeister and Ziirn (1879), Parasiten, p. 538, et seq. {Argas, cit. Huber,
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Kiichenmeister, F. (1855), Die in und an dem Korper des lehenden Menschen vorkom-
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Bihllofjraphy 35

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Important for Medical Men, Economic
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The remarkable development of parasitology in recent years, and the
increase in our knowledge of the part played by parasites in human and
animal diseases, demand a means of publication, in the English language, of
original papers dealing with the subject in its widest sense. It is proposed in
future to relegate all such papers to Parasitology.

Papers on the subjects we have mentioned are now scattered in journals of
widely different character, into some of which they are but grudgingly admitted.
We trust that Parasitology will fulfil the purpose the editors have in view,
of encouraging the study of parasitology, especially in relation to disease, by
providuig a means for the publication of papers relating to pathogenic and
disease-transmitting parasites.

It is therefore hoped that this publication will appeal not only to
medical men at home and abroad, but to veterinarians, zoologists and
agriculturalists.

Parasitology, though a supplement to the Journal of Hygiatie, will be
issued in the form of a separate yearly volume of four to five hundred pages.
The parts will be issued when sufficient material has accumulated. The sub-
scription price is .£1. Is. per volume (post-free), payable in advance; single
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scribers at the reduced rate of Ids. Subscriptions may be sent to any
Bookseller, or to Mb C. F. CLAY, Manaqbr, Cambridge University Press
Warehouse, Fetter Lane, London, E.C. (The Contents of the first three
numbers of Parasitology are given on the next page.)

PARASITOLOGY (continued)

CONTENTS OF VOL. I. NO 1. (March 1908.)

Jordan, K., and Rothschild, The Hon. N. C. Revision of the Non-
Combed Eyed Siphonaptera (Plates I — VII and 1 text figure),
pp. 1—100.

CONTENTS OF VOL. L NO. 2. (June 1908.)

Castellani, a. Note on a Liver Abscess of Amoebic Origin in a

Monkey (Plate VIII), pp. 101—102.
Imms, a. D. On the Larval and Pupal stages of Anopheles maculipennis

Meigen (Plates IX, X), pp. 103—133.
NuTTALL, G. H. F., and Graham-Smith, G. S. The Mode of Multipli-
cation of Piroplasma bovis and P. pifkeci in the Circulating Blood

compared with that of P. cards, with Notes on other species of

Piroplasma (Plate XI and 4 diagrams), pp. 134 — 142.
NuTTALL, G. H. F. Note on the Behaviour of Spirocha€ta£ in Acanthia

lectularia, pp. 143 — 151.
NuTTALL, G. H. F., Cooper, W. F. and Robinson, L. E. The Structure

and Biology of ffaemaphysalis punctata Canestrini and Fanzago, I.

(Plates XII — XVI, of which two are coloured, and 9 text figures),

pp. 152—181.
Masterman, E. W. G. Hirudinea as Human Parasites in Palestine,

pp. 182—185.
Harding, W. A. Note on a Gnathobdellid Leech (Limnatis sp. ?) from

Angola (1 figure), pp. 186—189.
Shipley, A. E. Note on Cystidicola farionis Fischer, A thread- worm

Parasite in the Swim-bladder of a Trout, pp. 190 — 192.
Leiper, R. T. Note on the Anatomy of Gystidicola farionis, pp. 193 —

194.

CONTENTS. OF VOL. I. NO. 3. (October 1908.)
Turner, G. A. Bilharziosis in South Africa, pp. 195—217.
Cleland, J. Burton. Note on Spirochaetes in Castration Tumours of

Pigs, pp. 218— 219.
Nuttall, George H. F. and Graham-Smith, G. S. Notes on the

Drug Treatment of Canine Piroplasmosis, pp. 220—226.
Durham, Herbert E. Notes on Nagana and on some Haematozoa

observed during my travels, pp. 227 — 235.
Minchin, E. a. Note on the Polymorphism of Trypanosoma Gambiense

(Plate XVII), pp. 236— 237.
Nuttall, George H. F., Cooper, W. F. and Robinson, L. E. On the

Structure of " Haller's Organ " in the Ixodoidea (Plate XVIII and

1 text figure), pp. 238—242.
Nuttall, George H. F. and Graham-Smith, G. S. The Development

of Piroplasma canis in Culture (Plate XIX and 1 text figure),

pp. 243—260.

Part II, Izodidae]

[May, 1911

TICKS

A MONOGRAPH OF THE

IXODOIDEA

By
GEORGE H. F. NUTTALL

M.A., M.D., Ph.D., Sc.D., F.R.S.

Fellow of Magdalene College

Quick Professor of Biology in the University of Cambridge

CECIL WARBURTON

M.A., F.Z.S.

Christ's College, Zoologist to the Royal Agricultural Society

W. F. COOPER and L. E. ROBINSON

B.A., F.Z.S., F.L.S. A.R.C.SC. (Lond.)

PART

II

By

GEORGE H. F.

NUTTALL

and

CECIL WARBURTON

CAMBRIDGE
AT THE UNIVERSITY PRESS

LONDON: FETTER LANE. C. F. CLAY, Manager
AND H. K. LEWIS, GOWER STREET

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BOMBAY AND CALCUTTA : MACMILLaN & CO., LTD.

I

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THE

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EDITED BY

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CONTENTS OF VOL. XI, NO. 1 (March 1911)

MoRdAN, H. DE R. The Differentiation of the Mannite-fermenting
Group of B. dysmiteriae with Special Reference to Strains Isolated
from Various Sources in this Country.

BAiNbRiDGE, F. A. and Dudfield, R. An Outbreak of Acute Gastro-
enteritis caused by B. paratyphosus (B.).

PenfOLD, W. J. Studies in Bacterial Variation. With special reference
to the chemical functions of the members of the typhoid-coli group.
(With Plate I, and 9 Charts.)

Bainbridge, F. a. and O'Brien, R. A. On the Paratyphoid Group
of Bacilli.

Trommsdorff, R., Rajchman, L. and Porter, Agnes E. A Severe
Outbreak of Food Infection Caused by a Paratyphoid Carrier.

Hewlett, R. Tanner, Villar, Sidney and Revis, Cecil, On the
Nature of the Cellular Elements Present in Milk. Part III.
The Milk of Animals other than the Cow. (For the British dairy
farmers' association.)

Porter, A. E. The Precipitin, Complementbinding, and Antiopsonic
Tests in Tuberculous and Normal Cattle.

Seidelin, Harald. The Quantitative Estimation of Indole.

Chick, Harriette, and Martin, C. J. The Fleas Common on Rata
in Different Parts of the World and the Readiness with which they
Bite Man. (With Plate II)

TICKS

A MONOGRAPH OF THE

IXODOIDEA

PART II

CAMBKIDGE UNIVERSITY PRESS

3Lontion: FETTER LANE, E.G.

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3Sombag anti Calcutta: MACMILLAN AND CO., Ltd.

All rights reserved

TICKS

A MONOGRAPH OF THE

IXODOIDEA

By
GEORGE H. F. NUTTALL

M.A., M.D., Ph.D., Sc.D., F.R.S.

Fellow of Magdalene College

Quick Professor of Biology in the University of Cambridge

CECIL WARBURTON

M.A., F.Z.S.
Christ's College, Zoologist to the Royal Agricultural Society

W. F. COOPER and L. E. ROBINSON

B.A., F.Z.S., F.L.S. A.R.C.Sc. (Lond.)

Cooper Laboratory for Economic Research, Watford

PART II

Cambridge

at the University Press

191 1

IXODIDAE

Section I. CLASSIFICATION
Section II. THE GENUS IXODES

by

GEORGE H. F. NUTTALL

and

CECIL WARBURTON

PREFACE TO PAET II

WE greatly regret the delay which has occurred in the publication
of this the second part of our work. The delay has not been
due to any lack of diligence on our part but to various difficulties which
we encountered in the preparation of our manuscript. For, when we
came to make a careful study of Ixodes, we found it necessary to re-
write almost all of the specific descriptions after personally examining
such types as were accessible. This necessitated journeys to Berlin,
Paris, Toulouse and London (British Museum), and a considerable
correspondence with heads of Museums, owners of type specimens, and
others in many places throughout the world.

We commence Section I of this part with (1) an historical review
dealing with the classification of ticks. This, and the succeeding
chapter (2), dealing with the superfamily Ixodoidea, might, perhaps,
with more propriety have appeared at the commencement of Part I,
but it is chiefly with regard to the Ixodidae that taxonomic difficulties
have been encountered, and, moreover. Chapter 2 forms a necessary
connecting link leading to the matter which follows. In Chapter 3 we
have gone with some detail into the generic diagnoses of Ixodidae, these
being illustrated in a manner not attempted hitherto if we except the
crude figures accompanying the publication of Lahille.

Section II, which forms tlie bulk of Part II, concerns itself solely
with the genus Ixodes, but for some of the matter contained in the
appendices. At the commencement of the section, under Synonymy,
we give our reasons for suppressing various generic and subgeneric
names which have been applied to forms included by us in this genus.
We definitely condemn many species which are merely nominal, and we
have referred others to the category of " Doubtful Species," as will be
seen by reference to the lists at the end. Under doubtful species we
include such as are founded on immature forms or which have been

vi Preface

imperfectly described ; we propose to retain them in this category until
they may have been properly described and shown to be valid by their
authors or other workers. We do not consider it justifiable, in the
present state of our knowledge, to found species on immature forms, a
proceeding which might be warranted if the immature forms of all
established species were known, whereas this is only true of 8 out of 51
species in the case of Ixodes. We agree in the main with Neumann in
the synonomies of species, both valid and condemned, but we have in
some instances arrived at different conclusions. Our lists of condemned
genera and species and of doubtful species, which have been compiled
by one of us (G. H. F. N.) with great labour, sufficiently demonstrate
the difficulties we have encountered in separating the wheat from the
chaff. These lists should at least prove useful to those desiring to
avoid the giving of already preoccupied names to new species o^ Ixodes,
Some of the synonyms must, from the nature of things, necessarily be
regarded as tentative.

Of the innumerable species of Ixodes that have been described, we
are only able to recognize 51, and there may be but 48 which are valid.
As will be seen from the text, a number of species and their various
stages are adequately described and figured by us for the first time.

The notes on Biology refer to matters of special interest relating to
some of the species. We have added thereto two appendices dealing
(I) with oviposition in Ornithodorus moubata, and (II) with the adaptation
of ticks to the habits of their hosts. The appendices are reprinted from
papers by Nuttall and Merriman and by Nuttall, which have recently
appeared in Parasitology, Vol. iv (1911), and which bear directly upon
the subject in hand.

Illustrations.

Curiously enough, some of the commonest species have never been
accurately figured before, or figured at all : we have endeavoured to
make good this deficiency.

The 94 new illustrations in the text, like our earlier ones, are in most
cases reproduced from large drawings made from unmounted specimens
preserved in alcohol. The drawings were made with the aid of a
camera-lucida mounted on a Zeiss binocular microscope and used in
conjuuction with a Zeiss drawing-board. A scale drawn to one side of
the figure usually indicates the magnification employed. Certain
highly magnified figures of the capitulum, hypostome and other structures

Preface vii

were drawn from specimens mounted in balsam, using a monocular
microscope, any errors of interpretation due to the transparency of the
chitin in the mounted specimen being corrected by reference to the
opaque object. We lay great stress on the avoidance of errors due to
the examination of mounted specimens; figures of ticks drawn from
such specimens are exceedingly misleading. Where a figure is re-
produced from a freehand drawing, it is referred to as a "sketch " in the
accompanying legend. We attach much importance to the illustration
of the main structural characters in conjunction with the descriptions,
for they convey a much more accurate and rapid impression to the
mind than any amount of description. There are certain structures
which should be figured in nearly all instances, and we have included
them in most of our illustrations.

The Part contains 202 text-figures (several being marked A and B)
and 14 figures comprised in four plates, the total number of figures
being 216.

Text-Figures.

Of the 94 new figures 84 were drawn by G. H. F. Nuttall^
6 ,, ,, F. M. Hewlett I

3 ,, ,, C. Warburton ( *

1 was drawn by L. E. Robinson j

Of the 107 previously 1 u,- ^- i m

,,.,,,. !■ 48 are from pubhcations by Neumann

published figures J

32 ,, ,, Nuttall and Warburton, jointly,

or as separate authors

4 „ ,, Wheler

2 „ ,, Salmon and Stiles
1 is from a publication by Evans

1 ,, ,, Lewis

2 are from publications by Nuttall, Cooper and Eobinson,

and Nuttall and Merriman
18 are reprinted from Parts I and II to illustrate matters
treated of in the appendices
202

Plate-Figures.

Of the 4 new figures 2 are from photographs by E. G. Wheler
2 ,, ,, L. E. Robinson

The 10 published figures are all from Wheler

14

* We have made it a rule to consult together about most of our figures, with a view to
ensuring their accuracy.

viii Prejace

The figures reproduced from Neumann are mostly printed from the
original blocks. These have all been purchased from the publishers of
his papers, or they have been duplicated from the original blocks with
the kind consent of Professor Neumann and of the editors of the various
journals in which they were published. Mr E. G. Wheler has generously
presented us with the blocks used in the illustration of his papers,
together with numerous unpublished photographs of ticks. Of the
figures by other authors, we have only chosen such as appear suitable
for the purposes of this book.

We have decided to publish our whole Bibliography as it stands
in the form of a separate fasciculus ; should further references be
necessary in connection with succeeding parts, short supplementary
lists of references will be appended to the respective parts. We shall
be much indebted to any of our readers who may draw our attention
to errors and omissions.

A cknowledgments.

We are specially indebted to Professor L. G. Neumann, of Toulouse,
for the generous aid he has given us throughout, not only in placing his
valuable collection of types at our disposal, but also in helping us with
his advice and counsel. Our thanks are also due to Geheimrath
Professor W. Donitz, of Berlin, for friendly encouragement and ready
aid repeatedly given.

The following Institutions and gentlemen have helped us very
materially by the loan of types and, in some cases, the gift of co-types :
The Berlin Museum (gift of /. schillingsi Neumann co-types, through
the courtesy of Geheimrath W. Donitz and Professor Dahl) ; the Paris
Museum (gift of " /. transversalis Lucas" co-types, through the courtesy
of Professor Bouvier) : the Hamburg Museum (loan of various types,
through the courtesy of Professor Kraepelin) ; the Bureau of Entomology,
U. S. Department of Agriculture, Washington, D.C. (gift and loan of
various specimens (some of which were identified by Mr N. Banks)
thanks to the courtesy of Dr L. O. Howard). The Hon. N. C. Rothschild
presented us with a very extensive collection of ticks from all parts of
the world, many specimens not having been identified. We, how^ever,
discovered that the collection included Neumann's /. australiensis
(types) and /. nitens (types). The Rev. O. Pickard- Cambridge, F.R.S.,
sent us, on loan, the unique remaining type of /. putus (a nymph) in his
collection. Professor A. Birula, of St Petersburg, presented a specimen

Preface ix

of /. signatus (type locality). Dr S. Hadwen, of Vancouver, British
Columbia, presented various stages of /. angustus (types of cT, o and
larvae). Dr J. H. Ashworth and Mr W. Evans, of Edinburgh, presented
us with our /. caledonicus (types of $, o and larva). Mr E. G. Wheler
presented a valuable collection, including some types of /. tenuirostris.
Numerous specimens have been sent to us by Mr C. P. Lounsbury,
Union Entomologist, S. Africa. Much material has reached us in
the form of collections sent for determination ; we may especially
cite the following sources : The Leland Stanford Junior University,
California (courtesy of Professor Vernon L. Kellogg) ; the Dublin Museum
(courtesy of Professor G. H. Carpenter and Dr Scharff ) ; the Entomo-
logical Research Committee for Tropical Africa (our thanks are due
to the Committee's Secretary, Mr Guy A. K. Marshall, for his uniform
courtesy) ; the Indian Museum, Calcutta (courtesy of Dr N. Annandale).
We have been permitted to retain specimens from these various
collections.

"We, are, moreover, personally indebted to numerous gentleman for
aid, as stated in the notes on the specimens in our collection appended
to the scientific descriptions.

The original literature on various species of Ixodes, whilst very
extensive, is in nearly all instances quite useless to the systematist,
because many descriptions are short and vague and others, whilst very
lengthy, omit to mention the specific characters whereby the species can
be distinguished from others. We, of course, except the excellent descrip-
tions of Professor Neumann, from which we have obtained much help.
Almost all recent papers dealing with ticks are merely compilations
from Neumann. The publications of Donitz are a marked exception to
the rule. The papers by Banks have been of some assistance to us,
though we have too often found his descriptions imperfect, and his
illustrations greatly wanting in accuracy.

G. H. F. N.

C. W.

Cambridge,
May, 1911

a5

EERATA.

107, lines 1 and 26,

for Sarconyssus
read Sai-conissus

134, line 4 from below,

for Sarconyssus Kolenati, 1857, p. 21
read Sarconissus Kolenati, 1856 and 1857, p. 21
line 5 from below

for Dermanyssus Kolenati, 1857, p. 20

read Dermanissus Kolenati, 1856 and 1857, p. 20

CONTENTS

OF
PART II

THE IXODIDAE

SECTION I
DEALING WITH THE CLASSIFICATION OF TICKS

PAGE

Historical Review relating to Classification, with Summary and Conclusions 105

The Superfamily IXODOIDEA

Synonymy and litei'ature . 113

The superfamily characters defined . . . . . . 114

Ihe Family IXODIDAE

Synonymy and literature . . . . . . . . 115

The family characters defined 115

Classification of the Ixodidae^ togetlier with an illustrated descrip-
tion of the main characters of the different genera belonging to
the family ........... 116

Explanation of the terms and signs used in the descriptions . 127

SECTION II

I. THE GENUS IXODES LATREILLE, 1795.

Synonymy and literature .......... 133

The generic characters defined . . . . . . • . . . 135

Keys for the determination of species of Ixodes :

Males 135

Females ............. 137

Nymphs 140

Larvae ..... ...... 141

Xll

Contents

Specific descriptions of valid species and of their varieties and subspecies
N.B. The signs $ , $ , o , L, alongside the specific names indicate
the various developmental stages of each species which are
known to science.
1. ricimcs (^ , 9 > o , L

„ with three varieties : scapularis ^ ,
ovatus 9
californicus $

2. nigricans $

3. acuminatus

4. dentatus 9

5. diversifossus

6. minor ^, $

7. holiviensis ^ , $ ,

8. fuscipes 9

9. scidphis $

10. spinicoxalis $

11. marxi $.

12. pratti $, $

13. ruhidus $ , o

14. hexagomis ^, $, o, L

„ with one variety : cookei ^ ,

15. nitens $ .

16. bicornis $

17. auritidus $

18. hrunneus $, o, L

19. cavipalpus $ , $

20. angustus ^ , $ , o , L

21. caledonicus $ , o , L

22. fossulaius ? .

23. acutitarsiis 9 •

24. gigas $ .

25. rubicundus (J, $

„ with one var:

26. simplex $ , o .

27. japonensis $ .

28. canisuga ^ , $ , o ,

29. texanus $

30. stilesi ? .

31. neumanni 5 , o

32. percavatus $ .

„ with one variety : rothschildi

33. pilosus $ , $ , o , L

„ with ? one variety : howardi $ ,

34. lunatus $

35. rasws (J, $

?

iety : limbatus

Contents

Xlll

c^, ?

36. ugandanus (J, ? •

,, with one subspecies : djaroiuntsit

37. cordifer $

38. holocyelus (^ , ? , o

39. sehillingsi c? , ? . o

40. ornithorhynch ?' ? , o

41. tasmani $

42. tenuirostris cJ, $,o, L

43. fecialis $

„ with one variety : aegrifossiis $

44. australiensis $

45. vestitus $ , o , L

46. putus (J , ? , o

47. signatus ^ > o , L .

48. unicavatus $ , o , L

49. loricatus (J, 9

„ with one variety : spinosus $

50. coxaefiircattis $ . . . .

51. vespertilionis (J, 95 0)L

Geographical distribution of the Genus Ixodes

List of condemned and doubtful species of Ixodes, including their synonymy

and literature, by G. H. F. Nuttall
Notes on doubtful species
Notes on the biology of Ixodes, by G. H. F. Nuttall

with a special index and including two appendices :

I. The process of copulation in Ornithodorus moubata, by G. H. F

Nuttall and G. ]\lerrinian .

II. On the adaptation of Ticks to the habits of their hosts, by
G. H. F. Nuttall

Index to valid species of Ixodes, together with a list of the collections in
which the types are to be found

PAOB

230
233
233
235
238
242
244
246
248
250
250
252
256
261
264
266
269
270
271

278

280
291
294

318
324
346

LIST OF ILLUSTEATIONS

PLATES

Plate IV.

Ixodes ricinus.

Fig. 1.

cj, dorsum x 12.

Fig. 2.

(5^, „ (mounted specimen).

Fig. 3.

?, » X 12.

Fig. 4.

$ , venter x 9.

Fig. 5.

o, dorsum x 12.

Fig. 6.

L, „ X 12.

Fig. 7.

Copulation.

facing p. 144

Plate V. Ixodes tenuirostris and Ixodes putus. facing p. 246

Fig. 1. /. tenuirostris (J, venter.

Fig. 2. „ $, „ X 11 (mounted specimen).

Fig. 3. /. putus (J, dorsum x 7.

Fig. 4. „ ? , „ X 7.

Plate VI. Ixodes vespertilionis. feting p. 276

Fig. 1. $ (unfed), dorsum x 7.
Fig. 2. ? (replete), „

Plate VII. Ixodes ricinus, females ovipositing in a glass dish, x |.

facing p. 310

IN THE TEXT

(Figs. 115-138 with the names of parts indicated in accordance
with the generic descriptions in the text.)

FIGURE PAGE

115. Illustrating Ixodes: I. ricinus ^, dorsum and venter . . . 117

116. „ ,, /. hexagonus 9 (replete), dorsum and venter . 117

117. „ „ /. cavipalpus ^, capitulum, dorsal and ventral

aspects 118

List of Illustrations

XV

FIGURE

118. Illustrating Ixodes: I. cavipalpus $ , capitulum, dorsal and ventral

aspects ......

„ /. bicornis $ , capitulum, ventral aspect

Haemaphysalis : H. tcellhigtoni ^ , dorsum and venter
$ , dorsum ......

Dermacentoi- : D. variegatus var. kamskadalus ^, dorsum

and venter

„ D. variahilis ^ i capitulum and scutum

Rhipicentor : R. bicornis ^ and $ , dorsal aspects .
„ R. bicornis ^, venter ....

Rhipicephahis : Rh. bursa (J, dorsum

„ Rh. sanguineus ^ , venter

„ Rh. sirnpsoni 5 » capitulum and scutum

Margaropus: M. vnnthemi ,^ , venter

„ M. winthemi $ , capitulum and scutum

Boophilus : B. decoloratus ^ , dorsum and venter

,, B. anmdatus $ , capitulum and scutum.

Hyalomma : Hyal. aegyptium ^ , dorsum and venter

„ Hyal. aegyptium $ , capitulum, dorsal and

ventral aspects
Amblyomma : A. cooper i ^, dorsum and venter
„ A. variegatum. $, part of venter

„ A. cooperi $ , dorsum

136 a and b. Digit of chelicera of Haemaphysalis pimctata, $
137. Spiracles of 15 species of Ixodidae ....
Tarsi (4th) of 10 species of Ixodidae
Ixodes ncinus ^ , dorsum and venter

„ „ (J, capitulum, dorsal and ventral aspects

„ „ (J, tarsi 1 and 4 ....

„ „ ^ and 9 ) digits and hypostomes

„ „ 5 , dorsum, part of venter, etc.

„ „ 5) capitulum, dorsal and ventral aspects

„ ,, o, dorsum aud venter

„ „ o, capitulum, dorsal and ventral aspects

„ ,, L, dorsum and venter

„ ,, L, capitulum, dorsal and ventral aspects

Ixodes ricinus var. scapidaris ^ i capitulum and scutum, part of

venter, etc

Ixodes nigricans $ , essential parts ....

151. Ixodes acuminatus $ , venter .....

152. „ ,, 9, hypostome and digit

153. Ixodes dentatus $ , capitulum, ventral aspect .

154. „ „ ? , coxae and trochanters .

155. Ixodes minor ^, venter

156. ,, „ (J, venter

157. „ „ S, basis capituli, dorsal aspect .

119.
120.

121.

122.
123.
124.
125.
126.
127.
128.
129.
130.
131.
132.
133.

134.

135 a.
135 b.

138.
139.
140.
141.
142.
143.
144.
145.
146.
147.
148.
149.

150.

118
118

119

120
120
121
121
122
122
122
123
123
124
124
125

125
126
126
126
128
131
132
148
149
149
149
150
150
151
152
153
153

157
160
161
161
163
163
164
164
165

XVI

List of Illustrations

FIGURE

158.

Ixodes

159.

)»

160.

Ixodes

161.

»

162.

»

163.

Ixodes

164.

j>

165.

Ixodes

166.

»

167.

Ixodes

168.

Ixodes

169.

Ixodes

170.

n

171.

Ixodes

172.

>>

173.

175.

176.

>)

177.

J?

178.

Ixodes

179.

Ixodes

180.

Ixodes

181.

■)■>

182.

Ixodes

183. „

184. Ixodes
185.

186.

187. Ixodes

188.

189.

minor ^, hypostome .

„ $ , essential parts

holiviensis ^, dorsum, venter, hypostome, spiracle, tarsus 4
„ $ , capitulum and scutum, part of venter, spiracle

,, o, dorsum, venter, tarsus 4 .

fuscipes 9 , capitulum and scutum, capitulum in ventral
aspect, coxae, spiracle and tarsus 4 .
„ 5 J capitulum and scutum, coxae with part of

capitulum ......

spinicoxalis $ , basis capituli and scutum, coxae with par
of capitulum ....

„ $ , hypostome ......

marxi $ , capitulum and scutum, part of venter, etc. .
pratti 9) capitulum and scutum, capitulum in ventral aspect

coxae, spiracle and tarsus 4 .
ncbidus 5 , dorsum, part of venter, spiracle and tarsus 4
„ o, capitulum and scutum, part of venter, spiracle

and tarsus 4 .....

hexagonus (J, dorsum and venter, spiracle and tarsus 4
„ 9 , (gorged), dorsum and venter, spiracle an<

tarsus 4

„ 9 ) capitulum and scutum, parts of venter,

spiracle and tarsus 4 .
„ $ , capitulum, dorsal, ventral and lateral aspects

„ 9 5 tarsi 1 and 4, highly magnified

„ o , capitulum and scutum, part of venter, spiracle

and tarsus 4 .... .

„ L, dorsum, venter, tarsus 3, hypostome .

nitens $ , capitulum and scutum, parts of venter, spiracle and
tarsus 4 ...... .

hicornis $ , part of venter, hypostome ....

auritulus ^ > capitulum in ventral and dorsal aspects, with

scutum

5) ? > hypostome .......

hrunneus $ , capitulum and scutum, coxae, capitulum in
ventral aspect .....

„ $ , digit, hypostome, ventral aspect of basis capituli

with coxae I .... .

cavipalpus ^ , dorsum, capitulum in dorsal and ventral aspects
„ ^ and $ , spiracles and digits
„ $ , capitulum in dorsal and ventral aspects

angustus ^, dorsum and venter, spiracle and tarsus 4

„ 9 > capitulum and scutum, parts of venter, spiracle

and tarsus 4 .... .

„ o, capitulum and scutum, parts of venter, spiracle
and tarsus 4

PAGK

165
166
167
168
168

169

169

172
172
173

174
176

176

178

179

180
180
180

181
181

185
186

188
188

191

191
193
193
194
195

196

197

List of Illustrations

xvii

FIGURE

190. Ixodes

191. Ixodes

192.

193.

>'

194.

Ixodes

195.

Ixodes

196.

Ixodes

197.

Ixodes

198.

J)

199.

Ixodes

200.

Ixodes

201.

55

202.

Ixodes

203.

)5

204.

55

205.

55

206.

55

207.

55

208.

55

209.

55

210.

Ixodes

211.

212. Ixodes

213. Ixodes

214.

215.

Ixodes

216.

55

217.

Ixodes

218.

55

219.

55

220.

55

221.

55

222.

Ixodes

223.

55

224.

Ixodes

angustus L, dorsum, venter, tarsus 3 .... .

caledonicus 9 5 capitukun and scutum, parts of venter, spiracle
and tarsus 4 ..... .

„ o, capituhnn and scutum, parts of venter, spiracle

and tarsus 4 ......

„ L, capitulum, scutum, coxae, tarsus 3, anal grooves

fossiilatus 9 , capitulum and scutum, parts of venter, spiracle

and tarsus 4

acutitarsus 9 ) dorsum, digit, coxae ....

gigas ^, dorsum and venter ......

rubieundv^ cJ, venter, hypostome .....

,, 9 1 capitulum and scutum, parts of venter, etc.

simplex 9 5 ca[)itulum and scutum .....

japonensis 9 5 capitulum in dorsal and ventral aspects .

,, 9 5 tlofsal and ventral aspects of the body, spiracle

and tarsus 4 . . . .

canisuga ^, dorsum and venter ....

„ ^, capitulum in dorsal and ventral aspects

55 ^5 hypostome

^5 legs 1-4

„ 9 , capitulum and scutum, capitulum in ventral

aspect . . ...

,, 9 , (gorged), venter, spiracle and tarsus 4

„ o , capitulum and scutum, part of venter, spiracle

and tarsus 4 ....

„ L, dorsum, venter, tarsus 3 .

texanus 9 5 capitulum and scutum, parts of venter, spiracle
and tarsus 4 . . . .

„ 9 5 capitulum and scutum, parts of venter, spiracle
and tarsus 4 ....

stilesi 9 5 capitulum and scutum, palp in profile, digit
neumanni 9 5 capitulum and scutum, parts of venter, spiracle
and tarsus 4 . . . .

„ o, capitulum and scutum, parts of venter, spiracle

and tarsus 4 . . . .

percavatus 9 s capitulum and scutum

,, 9 5 f'^'i'e part of venter ....

pilosus (^, dorsum and venter, spiracle and tarsus 4

„ 9 5 dorsum (without capitulum), venter, spiracle and
tarsus 4 .....

,, 9 5 capitulum in dorsal and ventral aspects
„ o , dorsum and venter, spiracle and tarsus 4
„ L, dorsum and venter, tarsus 3 and digit
lunatiis 9 5 dorsum .......

„ 9 5 venter .......

rasus (^, venter, digit, hypostome ....

PAGE
197

199

200
200

201
202
204
205
206
208
209

209
210
211
211
211

212
212

213

213

215

215
217

218

219
220
220
222

223
224
224
225
227
227
228

XVlll

List of Illustrations

FIGUKE

225.

Ixodes

226.

»5

227.

Ixodes

228.

))

229.

Ixodes

230.

Ixodes

231.

11

232.

11

233.

Ixodes

234.

11

235.

236.

Ixodes

237.

11

328.

11

239.

Ixodes

240.

11

241.

Ixodes

242.

11

243.

))

244.

)>

245 rt

and b.

246.

Ixodes

247.

Ixodes

248.

11

249.

11

250.

Ixodes

251.

11

252.

11

253.

11

254.

Ixodes

255.

11

256.

11

257.

11

258.

11

259.

11

260.

11

rasics ? , dorsum, venter, tarsus 4, spiracle aud figures showing
variability ....

„ $ , digit, hypostome, tarsi 1 and 4
ugandanus $, venter, hypostome .

„ $ , capitulum and scutum

cordifer (J, venter, tarsus 4 .
holocyclits (J, venter, hypostome .

„ 5 , capitulum and scutum, venter (replete), hypo

stome .......

„ o , capitulum in dorsal aspect

schillingsi (^ , venter, tarsus 4, digit ....

„ $ , capitulum and scutum, venter, spiracle and

tarsus 4

„ o , capitulum and scutum, parts of venter, spiracle

and tarsus 4

ornithorhynchi $ , (two specimens) dorsum and venter, spiracle
„ 5 ) tarsi 1 and 4

?, digit, hypostome.
tasmani $ , capitulum and scutum, hypostome, end of tarsus
1 with foot .
„ $ , dorsum, venter, spiracle

tenuirostris $, capitulum

11 6i P*''^ of venter .

„ $ , capitulum and scutum

„ $ , ventral aspect of capitulum, with coxae I

Ixodes fecialis $ , capitulum and scutum, ventral aspect of

capitulum with coxae I
fecialis var. aegrifossus $, capitulum and scutum
australiensis $ , capitulum and scutum, capitulum in ventral
aspect ......

,j 5 » venter, spiracle and tarsus 4

J, 5 1 ventral aspect of capitulum and coxae I

vestitus 5 , capitulum and scutum

11 ? 1 capitulum in ventral aspect, and essential

parts

„ o, dorsum, parts of venter, leg 4, etc.

„ L, dorsum, venter, leg 3

putus Si dorsum, venter, side view, hypostome, spiracle
„ 5, dorsum, venter, side view of palp, spiracle
tarsus 4 ....

„ $, dorsum

„ $, capitulum in dorsal and ventral aspects
„ 5 , scutum, digit, hypostome, tarsus 4

11 ? 5 spiracle

„ o , capitulum and scutum, capitulum in ventral aspect,
coxae with trochanters, spiracle, tarsus 4

229
230
231
232
234
235

237

238
239

240

241
243
243
243

245
245
246
246
247
247

249
249

251
251
251
253

254
255
255
257

258
258
259
259
259

259

List of TllnMrations

XIX

261. Ixodes signatus 5 , capitulum and scutum, capitulum in ventral

aspect, palp in jjrofile, coxae, spiracle,

tarsi 1 and 4, hypostome .... 262

262. „ „ 5 ) venter of gorged specimen 262

263. ,, „ o , capitulum and scutum, capitulum in ventral

aspect, coxae, anal groove, spiracles and

tarsus 4 263

264. „ ,, L, capitulum and scutum, parts of venter, tarsus 3 263

265. Ixodes unicavatus 9 ? capitulum and scutum ..... 265

266. „ „ o , capitulum and scutum, parts of venter, spiracle

and tarsus 4 ..... . 265

267. ,, ,, L, capitulum and scutum, jjarts of venter and

tarsus 3 ...... . 265

268. Ixodes loricatus ^, dorsum and venter ...... 267

269. „ „ 5 ) capitulum in dorsal aspect .... 267

270. Ixodes loricatus var. spinosus ^ , capitulum, scutum, jjarts of

venter, etc. 269

271. Ixodes coxaefurcatus (^ , dorsum, venter, sj^iracle, tarsus 4 . . 270

272. Ixodes vespertilionis ^, venter ........ 273

273. „ „ (^, capitulum in dorsal aspect .... 273

274. „ „ (J, digit and hypostome 273

275. „ „ c?, leg 1 273

276. „ „ ^, foot 273

277. ,, „ ? , capitulum and scutum, parts of venter,

spiracle and tarsus 4 . . . 274

278. „ „ $ , hypostome and digit 274

279. „ ,, $ , capitulum in dorsal and ventral aspects . 274

280. „ „ o, capitulum and scutum, parts of venter,

spiracle and tarsus 4 . . . . 275

281. „ „ L, capitulum in dorsal and ventral aspects,

with scutum, coxae and tarsus 3 . 275

282. „ „ L, digit 275

283. Hypostome of Ixodes ricinus $, taken whilst copulating . . 305
284-287. Process of oviposition in Ixodes ricinus 309

288. Ornithodorus mouhata $, with empty spermatophore attached to

vulva 319

289. A series of eggs of different species of Ixodoidea to show their

relative sizes 330

290-299. Hypostomes of Argasidae compared 332

300-307. „ „ Ixodidae „ 343

105

SECTION I
THE CLASSIFICATION OF TICKS.

I. Historical Review of the Subject ^

Although the classification of ticks may be considered as dating
from Linnaeus (1746, p. 479), who included them under Acari in the
large genus Acarus, scientific nomenclature did not actually commence
until the time of Latreille. This author (1795, p. 15) called the
Acari " tiques " and divided them into 11 genera of which two were
Ai^gas and Ixodes. Later (1804, p. 46) he included these two genera
with five others {Sarcoptes, Uropoda, etc.) under Riciniae. Hermann
(1804, p. 63) included Rhynchoprion (Argas) and Gynorhaestes (Ixodes)
under Acarina, but kept them far apart, not recognizing their close
relationship. Leach (1815, p. 387) called the Acarina Monomerosornata.,
and divided them into 11 families, one of which, Ixodides, included
Argas, Ixodes and Europoda. He was succeeded by von Heyden
(1826, p. 608), who upset previous classifications and again separated
Argas from Ixodes. Sundevall (1833) divided the Acari into six
families, one of which, Ixodides, included ticks only, viz. Argas and
Ixodes. DuGfes (1834, p. 5) divided the group into seven families, one
of which, Ixodei, included Ixodes, and Gamasei included Argas.

C. L. Koch (1844, p. 220 ; 1847, p. 13) ranked ticks in a special order
distinct from the Acari, naming the order Ricini, which included three
families: 1. Argasiden (Argasides, in English, with two genera: Argas
and Ornithodoros) ; 2. Ixodiden (Ixodide.*^, in English, with four genera :
Hyalomma, Haemalastor, Amblyomma and Ixodes) ; and 3. Rhipi-
stomiden (with four genera: Dermacentor, Haemaphysalis, Rhipistoma
and Rliipicephalus) ; families 2 and 3 were distinguished from each
other by the possession of long and short palps respectively. Koch's
classification was generally adopted by subsequent authors, and has

^ A number of classical authors refer to ticks. See Bibliography under Aristotle,
Cato, Varro, Columella, Pliny, Sammonius, Aldi'ovandi.

N. I, 8

106 Olassijication

stood the test of time, with some modifications, better than any other
which has been advanced. Gervais (in Walckenaer and Gervais, 1844,
p. 229), however, at the same period, divided Acarina simply into seven
genera, of which one genus, Ixodes, included all the ticks. Nicolet
(1855, p, 381) divided aerial and terrestrial (in contrast to aquatic)
mites into five groups, one of which was Ixodides. FiJRSTENBERG
(1861, p. 208) took over the classification of ticks from C. L. Koch,
without making any change. Donnadieu (1875) included Argas under
Ixodides. Mi^gnin (1876 a, p. 293; 1877, p. 86; 1880, p. 117; and
1892, p. 25) gave the Ixodides as one of eleven divisions of Acarina.
Murray (1877, p. 185) divided mites into eight families, of which one,
family 5, was the Ixodidae. Kramer (1877, p. 215) likewise gave
ticks family rank under the name Ixodidae, and Michael (1883, p. 50)
includes Argas in the family Ixodidae.

Canestrini and Fanzago (1877, p. 110) divided the family Ixodini
into four genera : Ixodes, Hyalomma, Haemaphysalis and Rhipicephalus.
Karsch (1879, p. 96) established the genus Margaropus. Canestrini
(1890, p. 491) subsequently divided the family Ixodidae (or Ixodinae)
into three groups: I. Poliopli, with almost the whole venter "'' coraz-
zato," i.e. covered by chitinous plates, literally " cuirassed " (one genus :
Ixodes), II. Tetraopli, with four adanal shields (two genera: Hyalomma
and Rhipicephalus), III. Anopli, with naked venter (two genera :
Dermacentor and Haemaphysalis). Ordered according to the structure
of the palps, he divided the Ixodidae into two sections : A. Cultri-
palpi (Genera Ixodes and Hyalomma), and B. Conipalpi (Genera
Phaulixodes^ [Berlese], Rhipicephalus, Dermacentoi' , Haemaphysalis
and Herpetobia^ [Canestrini]). He confined his classification to the
Italian ticks, of which he had a personal knowledge. Canestrini
(1892, p. 563) places ticks under Acaroidea as an order Metastigmata
comprising two families: 1. Ixodidae, 2. Argasidae. Marx (1892a,
p. 233) follows, in a measure, the classification of C. L. Koch, but
substitutes Cynorhaestea for Ricini (Ricinus being preoccupied), and
ranks Cynorhaestea as a suborder instead of an order; he divided
the suborder into two tribes, or groups : I. Catastomata, comprising
two families : 1. Argasidae (with two genera : Argas and Ornithodoros),
and 2. Eschatocephalidae (provisionally forming a "connecting link
between the two tribes ") ; and 11. Antistomata, comprising three
families: 1. Haemalastoridae (with two genera: Haemalastor and

' Since condemned, recognized as nymphal forms : Phaulixodes = Rhipicephalus,
Herpetobia = Haemaphysalis.

Classification 107

Sai'conyssus), 2. Ixodidae (with three genera: Ixodes, Amblyomnia and
Hyalomma), 3. Rhipistomidae (with five genera: Boophilus, Rhipi-
cephalus, Dermacentor, Rhipistonui and Haemaphysalis). Soon after-
wards, Banks (1894, p. 209) established the superfatnily Ixodoidea,
which we retain in this work.

Neumann (1896, p. 2) placed ticks (Ixodides) as a family in the
order Acari, and divided them into two subfamilies : I. Argasinae
(with two genera: Argas and Ornithodoros), and II. Ixodinae ; he sub-
sequently (1897, p. 325 and 1899, p. 107) recognized three tribes under
Ixodinae : Tribe 1. Ixodae (= Cultripalpi of Canestrini, including
four genera: Ixodes, Hyalomma, Amblyomma and Aponomma); Tribe 2,
Rhipicephalae (= Conipalpi of Canestrini, including three genera :
Rhipicephalus, Dei^macentor and Haemaphysalis); and Tribe 3 to
include Haemalastor. By Haemalastor he meant to refer to forms
like Ixodes vespertilionis, of which there are many bad species ; he,
however (1899, p. 166), recognised its close affinity to Ixodae, except
for the structure of the j/" capitulum, and gives Haemalastor only
generic rank after Ixodes. He founded the genus Aponomma (1899,
p. 180) to include Amblyomraa-like forms, differing from Amblyomma
in not possessing eyes, and in having broader bodies, besides occurring
almost exclusively on Ophidia and Saiiria. Neumann (1901, p. 318)
briefly reviews the subject of classification, and notes the fact that
C. L. Koch and himself were the only authors who based their classi-
fications upon an extended study of actual specimens. He (p. 321)
condemns Phaidixodes Berlese and Herpetobia Canestrini ; Haemalastor
Koch is referred to Hyalomma ; Sarconyssus Kolenati is referred to
Eschatocephalus Frauenfeld; (p. 276) Boophilus Curtice is united to
Rhipicephalus; Rhipistoma Koch and Opisthodon Canestrini, Gonixodes
Duges are included under Haemaphysalis ; Caris Latreille, Crotonus
Dumeril, Cynorhaestes Hermann are referred to as " des synonymes plus
ou moins precis d' Eschatocephalus et di' Ixodes"; Pseudixodes ^a\\ev is
referred to Dermacentor; Ophiodes Murray falls into synonymy {= Apo-
nomma), Xiphiastor Murray, and Adenopleura Macalister are condemned
as undeterminable, being possibly Amblyomma spp. ; Margaropus Karsch
is stated to be nothing more " qu'un individu anormal de Rhipicephalus
annidatus (Say) var. microplus." After thus clearing the ground of a
good deal of lumber, he (p. 322) reiterates that he is in agreement with
most zoologists in regarding the Ixodidae as merely a family of the
order Acarina, and he divides the family into ten genera which he
groups into two subfamilies : I. Ixodinae divided into two tribes :

8-2

108 Classification

1. Ixodae (Genera Ixodes, Eschatocephalus, Aponomma, Amhlyomma
aud Hyalonima), 2. Rhipicephalae (Genera Haemaphy sails, Rhipicephcdus
a.nd Dej'macentor); and II. Argasinae (Genera J.?'^as and Ornithodoros).

Salmon and Stiles (1901, p. 384), whilst following Neumann with
regard to the grouping of the various genera, raise the ticks to the
rank of a superfaraily, as had been done by Banks. They simply raise
Neumann's subfamilies and tribes to the rank of families and sub-
families respectively. Their classification is as follows : Snperfamily
Ixodoidea comprising two families : I. Argasidae (Genera Argas
and Ornithodoros), and II. Ixodidae, including two subfamilies :
1. Rhipicephalinae^ (Genera Rhipicephalus, Boophilus, Haemaphy salis
and Dermacentor), and 2. Ixodinae (Genera Ixodes, Eschatocephalus,
Aponomma, Amhlyomma and Hyalomma).

Neumann next (1902, p. 115) established the genus Geratixodes.
In 1904, p. 444, he created the subgenus Euixodes, to include all
Ixodes other than Geratixodes and Eschatocephalus, both of which were
degraded to subgenera. (We include all of these in the one genus
Ixodes in this work, see pp. 133-135, and do not see a valid reason
for retaining the subgenera.)

Neumann (1904, p. 444) attempted, further, to order the genera of
his subfamily Ixodinae according to their real affinities, a task, as we
have seen, previously attempted by Canestrini (1890), and Marx (1892).
Nevertheless, the original order adopted by C. L. Koch (1844), depending
upon the relative length of the palps, had hitherto been generally
accepted, and was until now retained by Neumann.

As pointed out by Neumann (1904, p. 445), the relative length of
a single structure like the capitulum does not give a safe basis for
classification, since, in the absence of •type specimens for purposes of
comparison, it leaves too much to the personal judgment of the
naturalist in defining what is "long" and what is "short." Although
of undoubted use, the division of Ixodinae into the two tribes mentioned
is unnatural and misleading, since it leads to misconceptions regarding
the real relationships existing between the genera. Thus Ixodes and
Hyalomma, included in Tribe 1, are not nearly so closely related as
Hyalomma and Rhipicephalus, the latter serving as the type of Tribe 2.
Moreover, the presence or absence of eyes, whilst of generic value, is of
no value as a means of ordering the genera according to their affinities,

^ Lahille (1905, p. 12) remarks that the subfamily should be styled Dermacentorinae
aud not Rhipicephalinae, if the rules of nomenclature are to be strictly adhered to, for
Dermacentor (Koch, 1844, p. 235) has priority over Rhipicephalus (Koch, 1844, p. 238).

Classification 109

for it brings Ixodes, Aponomma and Haemaphysalis together on the
one hand, and Amhlyomma, Hyalomma, Rhipicephalus and Dermacentor
on the other. Neumann considers it essential to take as a basis of
chxssification some external feature which is determined by a difference
in anatomical structure. Such a feature is to be found in the grooves
on the ventral surface. This would lead us back to the principle
adopted by Canestrini {vide supra), who was struck by the affinities
existing between Hyalomma and Rhipicephalus. Canestrini did not
include Amhlyomma and Aponomma under his Anopli, because he
confined himself to genera occurring in Italy, Neumann agrees that
Canestrini's groups Poliopli and Tetraopli are homogeneous, but that
this does not hold for the group Anopli, which includes Amhlyomma
{Aponomma'^), Dermacentor and Haemaphysalis. {Amhlyomma and
Aponomma are obviously allied, but Dermacentor and Haemaphysalis
remain disconnected.) Neumann attempted to represent the supposed
affinities between the different genera by the following plan, wherein
the position and length of the connecting lines indicate approximately
the degrees of affinity :

Ixodes Haemaphysalis

Hyalomma Amblyomma Aponomraa

I I

Rhipicephalus — ■ — Dermacentor

Neumann adopts Canestrini's groups as sections in the subfamily
Ixodinae, substituting names derived from type genera for the cor-
responding names given by Canestrini, thus :

Section I. Ixodeae (for Poliopli).

„ 2. Rhipicephaleae (for Tetraopli).
„ 3. Amhlyommeae (for Anopli).
Neumann characterizes the sections as fullows :

1. Ixodeae : ^ venter entirely covered by shields. Anal groove
contouring the anus in front and independent of the genital grooves.
Eyes absent. Capitulum elongate.

2. Rhipicephaleae : (/ venter bearing two adanal shields, usually
acciimpanied by accessory shields. Anal groove contouring the anus
behind and usually joining the genital grooves in front. Eyes present.
Capitulum long or short.

3. Amblyommeae : ^ venter devoid of shields. Anal groove as
under 2. Eyes often present. Capitulum as under 2.

^ We regard Aponomma as simply a subgenus oi Amhlyomma.

110 Classification

Neumann (1907 a, p. 26), in his latest paper in which he deals with
classification, includes the following genera under the various sections
above mentioned : I. Ixodeae (Ixodes), II. Rhipicephaleae (Rhipicepha-
lus, Margaropus^, Hyalomma), and III. Amblyommeae {Amhlyomyna,
Dermacentor, Aponomma, Haemaphysalis). Lahille (1905, pp. 11
et seq.) places ticks in a suborder, Arpagostoma", comprising two
families : I. Argasidae, and II. Ixodidae ; he divides the latter into
three groups according to the c^s: Group 1. Anopli (without ventral
plates) including four genera: Aponomma and Neumanniella (eyeless
= anommata) and Amblyomma and Dermacentor (ommata); Group 2.
Artiopli (with an even number of ventral plates) including three genera :
Hyalomma, Rliipicephalus and Boophilus (all possessing eyes) ; and,
finally. Group 8. Perissopli (with five ventral plates), comprising three
genera : Ixodes, Eschatocephalus and Ceratixodes (anommata). Lahille
(p. 16) founded the genus Neumanniella, of which more presently, to
include Ixodes transversalis Lucas, 184-4 ; 1845, p. 61, PI. I, Fig. 3
= Aponomma transversale (Lucas) Neumann, 1899, p. 188, Fig. 42.

Warburton (vii. 1907, p. 90) proposed the following classification:

IXODOIDEA

Argasidae Ixodidae

I , . _j

Prostriata Metastriata

. I 1

Brevirostrata Longirostrata

I 1 I 1

Group 1 Group 2 Group 1 Group 2

Argas Ixodes Haemaphysalis Dermacentor Hyalomma Amblyomma

Oruithodoros^ Rhipicentor* Aponomma

Rhipicephalus

Margaropus

Boophilus

Among the Ixodidae he separated widely Ixodes from the other
genera as Prostriata, which refers to the anal grooves surrounding the
anus in front, whereas in all the other genera the anal groove curves
about the anus posteriorly. The latter are grouped under Metastriata,
and are divided into two sections : Brevirostrata and Longirostrata,
according as they possess a short or long capitulum ; and each section
naturally falls into two groups, as indicated above. Banks (vili. 1907,

^ Meaning Boophihis Curtice, and including Margaropus Karsch.

- Should be Harpagostoiita (from barpagon = a grappling iron).

3 We have not hitherto adopted the latinized transcription Ornitliodorus but shall do
so henceforth in accordance with the rules laid down in The International Code of
Zoological Nomenclature. * Since added.

Classification 111

p. G2, as stated in Banks, 1908, pp. 13, 14) advanced another classifica-
tion. He divides Ixodidae into two subfamilies : I. Ixodinae {Ixodes
and Ceratixodes^), having the anal groove in front of the anus, and,
II. Amblyomminae (including the other genera of ticks) having the
anal groove posterior to the anus. He divides the Amblyomminae
into four tribes as follows :

1. Rhipicephalini (including lihipicephcdus and Margaropus).

2. Haemaphysalini {Haemaphysalis).

3. DermaceiUorini {Dermacentor).

4. Amblyommiiii {Amhlyomma, Aponomma and Hyalomma).

DONITZ (1907 a, pp. 1-8, reprint) re-established the genus Margaropus
Karsch, 1879, which Neumann (1901, p. 281) had regarded as a
monstrosity of " Rhipicephalus amudatus var. micropla." Neumann
has never recognized the genus Boophilus Curtice ; he continued to
include Boophilus under Rhipicephalus until he became convinced of
the validity of the genus Margaropus, when he concluded that it had
priority over Boophilus. Since then, he has referred species of Boophilus
to Margaropus, but as we shall see, both genera are perfectly distinct
and valid.

NUTTALL and Warburton (1907, p. 398) founded the genus
Rhipicentor.

DoNiTZ (1910, p. 400) passes under review the recent attempts at
classification by Neumann, Lahille, Warburton and Banks. He agrees
with us in condemning Eschatocephalus and Geratixodes and referring
them to Ixodes, and he also recognizes the genera Boophilus and
Margaropus as distinct. He condemns Neumanniella Lahille, 1905,
until there is better reason for recognizing such a genus. It will be
remembered that Neumann (1899, p. 188) stated that Aponomma
transversale (Lucas) Neumann possessed no anal groove, and might,
consequently, be referred to another genus if another species resembling
it were found. Lahille, however, founded the genus Neumanniella
without ever having seen the types. Donitz denies that we know
enough of the relationship existing between the genera (apart from
Ixodes) grouped under Ixodidae to reach any final conclusions regarding
their arrangement in respect to each other; it is possible that differences
of internal structure may prove of help in the future. He prefers, in a
measure, the older classification of Neumann which arose out of that
originally advanced by Koch. Koch, as we have seen, divided ticks

1 Made a subgenus by Neumann, 1904, and now suppressed by us and by Donitz, 1910.

11*2 Classification

into three families : Argasidae, Ixodidae (with long palps) and Rhipi-
stomidae (with short palps). The relative length of these structures
is certainly convenient in the practical work of determination. Banks
(1908, p. 12) objects to too much weight being laid on the relative
length of the palps, partly on the ground that it hardly permits us to
"distinguish some species of Amblyoinma from Dermacentor." DoNlTZ
does not approve of Neumann's subsequent adoption (1904) of the
plan of Canestrini (1890), the latter's Poliopli, Tetraopli and
Anopli corresponding to Ixodeae, Rhipicephaleae and Amblyomnieae
of Neumann. As Doriitz points out, Ambli/omma are not Anopli ; a
number possess small chitinous plates^ (analogous to the plates in
Ixodes) situated posteriorly upon the ventral surface. Again, Mar-
garopus has a ventral plate corresponding to the median plate in
Ixodes, but Margaropus, by its general structure, is allied to BoophiliLS
and Rhqncephalus, i.e. Tetraopli (Rhipicephaleae), and not to Poliopli
(Ixodeae). Banks (1908, p. 13) very properly criticizes the classification
of Lahille (190o), which is based on the characteristics of the (f, and
thus separates Hyalomnia widely from Amhlyomma, most authors
agreeing that they are allied, the $s being at times difficult to
distinguish. Banks fails to see "the importance of secondary sexual
characters " which lead to Lahille's classification, and considers that
they " are certainly not of sufficient value to characterize groups higher
than genera and are better employed for groups of lesser rank." DoNiTZ
(1910, p. 401) likewise objects to Lahille's classification on the ground
already stated that Amhlyomma are not Anopli, and that Margaropus
would be classed as Perissopli, whereas they are closely related to
Artiopli. He agrees with Warburton and with Banks in separating
the genus Ixodes widely from the other genera, because of the structure
of the anal groove, apart from other important differences. Donitz
(p. 403) divides the Ixodidae into three divisions ; I. Ixodeae (one
genus, Ixodes), II. Amblyommeae (=Longirostrata of Warburton, with
three genera : Amhlyomma, Aponomma, Hyalomma), and III. Rhipice-
phaleae (=Brevirostrata of Warburton, with six genera: Rhipicephalus,
Boophilus, Margaropus, Dermacentor, Rhipicentor and Haemaphysalis).
Banks (1908, pp. 13, 14) does not agree with the classification of
Warburton in so far as the latter places Dermacentor with Rhipicephalus.
Banks considers this ill-advised, and places Dermacentor close to Am-
hlyomma. The genus Rhipicentor Nuttall and Warburton, 1907, however,
directly links Rhipiceplialus to Dermucentor, as is indicated by the
1 " Plaques" in our desciiptions, see Fig. 135a.

Ixodoidea 113

name, and the discovery of this intermediate type confirms the view
advanced by Warburton. Banks agrees with us in not placing Hijalomma
near Rhipicephalus, as has been done by Neumann (1907 a, p. 26).

Summar]/.

The perusal of the foregoing account of the views held by
different authors who have written upon the classification of ticks,
shows that a great deal of confusion has undoubtedly existed. At
a fairly early date, the Argasid and Ixodid ticks were recognized as
possessing very distinctive features. The confusion that has arisen is
mainly due to efforts in classifying the Ixodidae. The proposal to
separate the latter into two distinct sections (Warburton, Banks) is
based on anatomical grounds, i.e. the structure of the anal groove in
the Prostriata in conti^ast to the Metastriata. In some ticks (Boophilus,
Margaropus) the anal groove may be obsolete ; they are, however,
obviously related to other metastriate forms in general structure. It
appears advisable, only as a matter of convenience, to give these sections
the new names (suggested by Warburton), in view of the great confusion
of subfamily, tribal, group, and other names, all of which have been
based on the original generic names. The division of the Metastriata
into Brevirostrata and Longirostrata is also one of convenience, and
their subdivision into two groups each follows naturally when the
characters of the genera are taken as a whole.

With regard to the genus Neumanniella Lahille, 1905, to which
reference has been made in the preceding pages, we would state that
we have examined the types of Ixodes transversalis Lucas and agree
witii Neumann in retaining the species under Aponomma. On very
careful examination we have detected traces of an anal groove posterior
to the anus. Consequently the genus Neumanniella falls into synonymy.

Superfamily IXODOIDEA Banks, 1894.
Synonymy and Literature :

Genus Acarus (in part) Linnaeus, 1746, p. 479.

Family Riciniae (in part) Latreille, 1804, p. 46 ; 1806, p. 151 ; 1829, p. 286.

Risso, 1826, p. 180. Oudemans, 1896, p. 191.
Family Ixodides Leach, 1815, p. 396 (includes Europoda). Sundevall, 1833.

Fuller, 1896, p. 763.
Family Ixodea Burmeister, 1837, p. 579. Grube, 1859, p. 455. Gerstaecker,

1860, p. 464.

114 Ixodoidea

Order Ricini Koch, 1844 ; also 1847, p. 5. Fiirstenberg, 1861, p. 208. Canestrini,
1890, p. 482.

Genus Ixodes Gervais, in Walckenaer and Gervais, 1844, p. 229.

Family Ixodida Kiichenmeister, 1855, p. 421. Kolenati, 1857, p. 19.

Group Ixodides Nicolet, 1855 (cited by Michael, 1883-1887).

Family Ixodidae Leach, in Gerstaecker, 1863, p. 343 ; 1873, p. 464. Murray,
1877, p. 185. Conil, 1877, p. 28. Kramer, 1877. Glaus, 1880, p. 652.
Michael, 1883, p. 50. Berlese, 1885, p. 131. Ludwig, 1886, p. 612. Riley,
1887, p. 744. Marx, 1892, p. 232. Trouessart, 1892, p. 28. Neumann, 1892 b,
p. 94. Railliet, 1893, p. 631. Braun, 1895, p. 257. Osborn, 1896, p. 255.
Ward, 1900 (a), p. 193, and {b\ p. 430. Neumann, 1901, p. 322. Stiles and
Hassall, 1901, p. 1. Neumann, 1904, p. 444. Lahille, 1905, p. 16. Wheler,
1906, p. 400. Pocock, 1907, p. 190. Donitz, 1907, p. 1. Neumann, 1907 a,
p. 26. Newstead, Dutton and Todd, 1907, p. 99. Bonnet, 1908, p. 247.
Blanchard, 1909, p. 80.

Family Ixodides Donnadieu, 1875. Megnin, 1876 a, p. 293; 1877, p. 86; 1880,
p. 117 ; 1892, p. 25. Railliet, 1886, p. 495 ; 1895, p. 703. Neumann, 1888,
p. 82 ; 1892 a, p. 90 ; 1896, p. 1 ; 1897, p. 324; 1899, p. 107 ; and subsequently
to 1910. Blanchard, 1890, p. 322 ; 1909, p. 5. Brumpt, 1910, p. 510.

Order Metastigmata Canestrini, 1892, p. 563.

Suborder Cynorhaestea Marx, 1892, p. 233.

Suborder Arpagostoma Lahille, 1905, p. 11. (Should be Harpagostoma.)

Superfamily Ixodoidea Banks, 1894, p. 209. Salmon and Stiles, 1901, p. 383.
Warburton, 1907, p. 90. Banks, 1907, p. 62, and 1908, p. 7. Nuttall,
Warburton, Cooper and Robinson, 1908, p. 1 (Part I of this work). Castellani,
1910, p. 460.

Superfamily Characters. Acari of the suborder Metastigmata
{the breathing apertures being somewhat posteriorly situated) charac-
terized by the possession of a movable false head, or capitidum, of a
special structure. It consist?, of a basal portion {basis capituli), a pair
of palps, protrusible chelicei'ae with digits serrate externally, and a rigid
hypostome almost always toothed on its ventral surface. All are blood-
sucking mites, parasitic on animals.

N.B. The Acarina, or Mites, arc divided into the suborders Vermiformia,
Astigmata, Metastigmata, Helerostigmata, Prostigmata, Notostigmata.

Besides the Ixodoidea, the Metastigmata include the Onhatidae and the Gamasidae.
The only mites in the least likely to be confounded with the Ixodoidea are the
remarkable group of Gekobiidae, parasitic on lizards. Their mouth-parts strongly
resemble those of the ticks, but they belong to the suborder Prostigmata, having
their breathing orifices near the palps.

The Ixodoidea are divided into two families, I. the Argasidae (see
Part 1) and II. the Ixodidae.

Ixodidae 115

II. Family IXODIDAE Murray, 1877.

Synonymy and Literature:

Family Ixodei Diigfes, 1834 n, p. 15; 1834 c, p. 33.

Family Ixodiden Koch, 1844, p. 220; 1847, p. 13. Furstenberg, 1861, p. 208.

Family Ixodidds Gervais and van Beneden, 1859, p. 460.

Family Ixodidae Murray, 1877, p. 185. Pavesi, 1884, p. 483. Canestrini, 1890,
p. 530 ; 1892, pp. 5G3, 581 ; 1897, p. 468. Trouessart, 1892, p. 22. Supino,
1897 a, p. 241. Pocock, 1900 a, p. 48. Salmon and Stiles, 1901, p. 384. Banks,
1905, p. 42. Lahille, 1905, pp. 11 ei! seq. Banks, 1907, p. 62. Hunter and
Hooker, 1907, p. 46. Manson, 1907, p. 205. Warburton, 1907, p. 89. Banks,
1908, pp. 12 et seq. Nuttall, Warburton, Cooper and Robinson, 1908, p. 1.
Donitz, 1910, p. 400. Stiles, 1910, p. 12.

Family Ixodini Canestrini and Fanzago, 1877, p. 110. Canestrini, 1890, pp. 475,
491 ; 1892, p. 558.

Family Ixodinae Karsch, 1880, p. 41.

Tribe Ixodid^S Megnin, 1880, p. 120.

Subfamily Ixodidae Berlese, 1885, p. 131.

Tribe Ixodin^S Railliet, 1886, p. 495. Neumann, 1888, p. 89; 1892 a, p. 93.

Subfamily Ixodinae Trouessart, 1892, p. 38. Railliet, 1895, p. 704. Neumann,
1896, p. 2 ; 1827, p. 325 ; 1899, p. 107. Ward, 1900 (a), p. 196 ; {b) p. 435.
Neumann, 1904, p. 444; 1907 a, p. 26. Howard, 1908, p. 89. Blanchard,
1910, pp. 55-77. Brurapt, 1910, p. 510.

Tribe Ixodinae Neumann, 1892 b, p. 96.

Group or Tribe Antistomata Marx, 1892 a, p. 233,

Family Characters \ Scutate ticks luith terminal capitulum and
spiracles posterior to coxae IV. Sexual dimorphism marked, the males
being almost entirely covered by the scutum, and incapable of great
distention ; while the scutum of the distended female appears as a small
shield behind the capitulum. Porose areas present on the $ capitulum.
Where eyes are present they ar'e situated laterally upon the scutum. 'The
fourth article of the palp is reduced to form a tactile papilla. Type
genus : Ixodes.

1 In this connection refer to Explanation of Terms and Signs used in the descriptions,
p. 127.

116 Classification of Ixodidae

CLASSIFICATION OF THE IXODIDAE.

There are nine genera: Ixodes, Haemapliysalis, Dermacentor,
Rhijncentor, Rhipicephalus, Margaropus, Boophilus, Hyalomma and
Amhlyomma (including the sub-genus Aponomma). They present
varying degrees of affinity, which may be indicated by arranging them
into groups according to the following scheme :

Ixodidae

I ^ — 1

Prostriata Metastriata

, I . ,

Brevirostrata Longirostrata

I — ^ n r ^ n

Group 1 Group 2 Group 1 Group 2

I.I I '

1. Ixodes 2. Haemaphysalis 3. Dermacentor 8. Hyalomma 9. Amblyomma

4. Rhipicentor (Aponomma)

5. Rhipicephalus

6. Margaropus

7. Boophihis

Ixodes is clearly marked off from the other genera by a number of
characteristics, of which the most striking are the anal groove sur-
rounding the anus in front (Prostriata) and the absence of festoons.
The remaining genera fall naturally into two divisions: the one
characterized by a comparatively short, and the other by a comparatively
long capitulum. Occasionally, forms are encountered which are aberrant
in this respect.

SECTION I. PROSTRIATA.

With anal grooves surrounding the anus in front.

Genus (1) Ixodes: inornate, without eyes and without festoons;
spiracles round or oval ; palps and basis capituli of variable form ; coxae
either unarmed, trenchant, spurred or bifid ; tarsi without spurs. Sexual
dimorphism pronounced, especially with regard to the capitulum ; iii the
(/ the venter is covered by non-salient plates : one pregenital, one median,
one anal, two adanal and two epimeral plates. Figs. 115-119.

Type species : Ixodes ricinus (Linnaeus).

Ixodidae: Genus Ixodes

117

IXODES

-^u/ur-U-r,

Fig. 115. I. ricinns (Linn.), c? : in dorsal and ventral aspects (from Nuttall, 1908,

G. H.F.N, del.).

^cuZu^i'n.

Fig. 116. I. hexagomis Leach, 1815, ? (replete) : in dorsal and ventral aspects

(Original, G. H. F. N. del.)

118

Ixodidae: Greniis Ixodes

cii-acZ

alCcc/k- 3

Fig. 117. 7. cavipalpus Nuttall and Warbnrton, 1908, c7 : capituhim in dorsal and
ventral aspects (from Nutt. and Warb., G. H. F. N. del.).

"y'^^jS^nr^

Fig. 118. I. cavipalpus, Nutt. and Warb., 1908, ? : capitulum in dorsal and ventral
aspects (from Nutt. and Warb., G. H. F. N. del.).

Fig. 119. /. bicomis Neumann, 1906, ? : capitulum in ventral aspect to show auricula.

(Neumann del.)

Ixodidae : Gciuih Haeinap}uiHal\>^

IH)

SECTION II. METASTRIATA.

With anal groove contouring the anus behind.

Note : In two genera, Boopliilus and Margaropns, clearly attributable
to this section by their general structure, the anal groove is faint or
obsolete. Any tick in which the anal groove cannot be made out may
with safety be referred to the Metastriata.

(a) Brevirost7'ata.
Group 1.

Genus (2) Haemaphysalis : inornate, without eyes hut with
festoons; with usually short conical jmlps whose second articles pro-
ject laterally beyond the basis capituli, which is rectangular dorsally.
With dorsal process on first trochanter. Usually of small size and
but slightly chitinized. Sexual dimorphism slight. The (/ shows no
ventral plates or shields. Spiracles in </ usually ovoid or comma-
shaped; in %, rounded or ovoid. Fig. 120.

Type species : Haemaphysalis concinna C. L. Koch.

HAEMAPHYSALIS

f^aJ/ii^"'^^

^o^ij'-^''^-^-'^

**tx *^'<x.^«- /uist -a^^xji^ q^a

Fig. 120. //. welUiifjtoiii Nuttall aud Warburton, 1908, j in dorsal and ventral aspects
and ¥ in dorsal aspect (from Nutt. aud Warb., E.W. del.).

120

Ixodidae: Genus Dermacentor

Group 2.

Genus (3) Dermacentor: usually ornate, with eyes and festoons;
with short, broad or moderate palps and basis capituli rectangular
dorsally. In some species coxae I to IV of the ^ increase pro-
gressively in size; in all species coxa IV is much the largest; the (^,
moreover, shows no ventral plates or shields. Coxa I bifid in both sexes.
Spiracles sub-oval or comma-shaped. Figs. 121, 122.
Type species : Dermacentor reticidatus (Fabricius).

DERMACENTOR

jije/yuZaZ

Fig. 121. D. varipfjatus var. kamshadalus Neumann, 1908, j : in dorsal and ventral
aspects (from Neumann, 1908).

Fig. 122. D. variabilis (Say, 1821), ? : capitulum and scutum (sketch from Salmon and
Stiles, 1901; G. H. F. N. del.).

Ixodidae: Genus Rhqncentor

121

Genus (4) Rhipicentor: inornate, with eyes and festoons ] with short
palps, with basis capituli hexagonal dorsally and having very prominent
lateral angles. Coxa I bifid in both sexes. The </ resembles lihipi-
cephalus dorsally, Dermacentor ventrally ; coxa IV is much the largest;
no ventral plates or shields. Spiracles sub-triangular ( $ ) or comma-
shaped ((/•). Figs. 123, 124.

Type species : Rhipicentor bicornis Nuttall and Warburton.

^a-^^

RHIPICENTOR

'^e^f-^tiro'T'^-^

Fig. 123. E. bicornis Nuttall and Warburton, 1908, c? and ? : dorsum (from Nutt. and

Warb., E.W. del.).

Fig. 124. R. bicornis, <j : venter, etc., as under Fig. 123 (G. H. F. N. del.).

N. I. 9

122

Ixodidae: Genus Rliipiceplialus

Genus (5) Rhipicephalus : usually inornate, with eyes and festoons,
with short palps and basis capituli usually hexagonal dorsally. (The ^
of one species, Rhipicephalus pulchellus, has a Dermacentor-\ike capitu-
lum, and both sexes are ornate.) Coxa I bifid. The j/* possesses a pair
of adanal shields and usually a pair of accessory adanal shields ; some
j/'s, when replete, show a caudal protrusion. Spiracles bluntly or
elongate comma-shaped. Figs. 125-127.

Type species : Rhipicephalus sanguineus (Latreille).

RHIPICEPHALUS

-2

Fig. 125. U/i. Z^itrsa Canestrini and Fanzago, 1877, i : dorsum. (Original, L. E.R. del.

■S/tji/iMAtjL

ciir^.jaJ- AZ

Fig. 126. Fig. 127.

Fig. 126. Eh. sanguiiieus (Latreille, 1804), d : venter (from Warb. and Nutt., 1909,

G. H.F.N, del.).
Fig. 127. Rh. simpsoni Nuttall, 1910, ? : capitulum and scutum (from Nuttall,

F.M.H. del.).

IxocUdae: Genus Margaropus

123

With anal grooves obsolete.

Genus (6) Margaropus^: inornate, with eyes, hut without festoons,
with short palps and capituluni intermediate between that of Rhijnce-
phalus and Boophilus ; highly chitinized ; the unfed adults of large size.
The $ with very small scutum. Coxae conical, unarmed but for a
small spine posteriorly on coxa I. The ^ with a median plate prolonged
in tiuo long spines projecting beyond and to either side of the anus ; with
coxae similar to those of $ ; legs increasing progressively in size from
pair I to IV, the articles especially of leg-pair IV greatly swollen. When
replete, the cf shows a caudal protrusion. Anal groove obsolete.
Spiracles rounded or short-oval in both sexes. Figs. 128, 129.

Type species : Margaropus winthemi Karsch.

MARGAROPUS

j,ey*u/:ac ff^fcez

Fig. 128.

■^LCL-C^ OA^CCO!,

Fig. 129.

Fig. 128. il/. winthemi Karsch, 1879, i : venter (from Neumann, 1907, Nu. del.).
Fig. 129. M. winthemi Karsch, 1879, ? : capitulum and scutum. (Original, sketch
by C. W.)

1 This name has become established. It should read Margaritopiis, signifying beady-

9—2

124

Ixodidae: Genus Boophilus

Genus (7) Boophilus: inornate with eyes, hut without festoons ; with
very short compressed palps ridged dorsally and laterally ; basis capituli
hexagonal dorsally ; slightly chitinized\ the unfed adults of small size.
Coxa I bifid. Anal groove obsolete in %, faintly indicated in $ . The
$ ivith a small scutum ; the </ with adanal and accessory adanal shields.
Spiracles rounded or oval in both sexes. Figs. 130, 131.

Type species : Boophilus annulatus (Say).

BOOPHILUS

-/laJ/La/ ^2^1^ /-

tt^cAc^-Ai .toA^i^

Fig. 130. B. decoloratus (Koch, 1844), <f : dorsum and venter (from Warb. and Nutt.,

1909, G. H.F.N, del.).

Fig. 131. B. annulatus (Say, 1821), ? : capitulum and scutum (sketch from Salmon and
Stiles, 1901, G. H.F.N, del.).

Txodidae: Genus Hyalomma

125

(6) Longirostrata.
Group 1.

Genus (8) Hyalomma : ornamentation absent or present, at times
confined to the legs ; with eyes, with or luithout festoons, with long palps
(shorter in Hyalomma monstrosiim (/*) and basis capituli sabtriangidar
dorsally. The % approaching Amblyomma. The </ ^uith a pair of
aclanal shields, and with or withoid accessory adanal shields and two
posterior abdominal protrusions capped by chitinized points. Coxa I
bifid. Spiracles comma-shaped. Figs. 132, 133.

Type species: Hyalomma aegyptiam (Linnaeus).

HYALOMMA

•6-o(U^ ■^,A^rCi

^■ey*tx^ci£ q,/i

Fig. 132. Kijal. acgtiptium (Linn.), i : dorsum and venter (from Warb. and Nutt. 1909,

G. H.F.N. andE.W. del.).

1^^' 7'(

Fig. 133. Hyal. aegijptitim (hinn.), ^ : capitulum in dorsal and ventral aspects. (Original,

G. H.F.N, del.)

126

Ixodidae: Genus Amhlyomma

Group 2.

Genus (9) Amblyomma : generally ornate, with eyes and with
festoons. With long palps, of which article 2 is specially long ; basis
capituli of variable form. The ^ without adanal shields, but small
ventral plaques are occasionally present close to the festoons. Spiracles
subtriangular or comma-shaped. Figs. 134, 135 a and b.

Type species : Amblyomma cajennense (Fabricius).

Subgenus: Ai)onoTDQ.iD.a, = Amblyommxi, which are devoid of eyes, or
in which the eyes are poorly developed. The body is frequently very
broad. They occur almost exclusively on Reptilia.

AMBLYOMMA

i^iiA^e- ony'&i^iM^

a/^Lcu/ ^^<riri^

-fje.<f^to-rryi^

Fig. 134. A. cooperi Nuttall and Warburton, 1908, <? : dorsum and venter (from Nutt.

and Warb., E.W. del.).

-/i^Znn^

Fig. 135 a.

Fig. 135 h.

Fig. 135 a. A. variegaUtm (Fabricius, 1794), 3 : posterior part of venter, showing plaques.

(Original, G. H. F. N. del.)
Fig. 135 6. A. cooperi, ? : dorsum, schematized (from Ticks, Part I, G. H. F. N. del.).

Terminology 127

Explanation of Terms and Signs.

Refer to Figures 115-138.

For the sake of brevity and precision, we have adopted certain signs
and terms in our descriptions of Ixodidae, as follows : —

cJ, 9, o, L, signify male, female, nymph and larva respectively ; c^s, ^sandos
for plural.

? after the names of parts, like Hypostome, Chelicera, Tarsus, etc., means that
these parts are missing in the type.

L. = the extreme length of the tick, excluding the capitulum and legs. We prefer
to exclude the capitulum when measuring the length (from the scapulae to
the posterior border) because the capitulum is frequently injured or tilted
so as to render measurement difficult.

ir. = its width at the widest part. Where the measurements are given as .t'X ?/,
measurement x stands always for the length and y for the breadth, [l. and
w. refer to length and width of various structures. )
Capitulum = the " rostrum," " head " or " false head " of various authors.

Btms capituli, or, shortly, base = t\\Q basal ring, "Kragen" of some authors. It
often shows a transverse elevated dorsal ridge with edge directed backward.
The ridge may have protruding angles ; the latter we term cornua.
There may also be a ventral ridge. The auricula signifies a protruding
retrograde process at the lateral angles of the ventral ridge posterior to
the insertion of the palps. When the length of the capitulum is given it
is measured from the tip of the hypostome to the dorsal ridge in the
median line. Where the dorsal ridge is absent it is similarly measured
to the ventral ridge. When the measurement is made ventrally this is
stated, the dorsal measurement being usually employed. The clear space
between the porose areas (only present on the 5 capitulum) is spoken of
as the interval.
Hypostome = the "maxilla," "radula," "labium," or " Unterkiefer " of various
authors ; a median ventral structure rising from the basis capitiili, and
bearing recurved teeth. The dentition is indicated by figures on either
side of a vertical line. Thus 3 | 3 means three longitudinal tiles of teeth
on each half of the hypostome. The hypostome may be pointed, rounded,
or emarginated distally. It may be armed from tip to base with teeth or
only bear teeth along a part of its length. An unarmed, protruding median
ridge, which broadens basally, may run down the length of the hypostome,
starting near the tip. When a hypostome is described as having a corona,
the ti[) beai's a number of very minute denticles.

128

Terminology

Chelicerae = t\ie "mandibles" or " pseudo-chelicerae " of some authors, paired
structures lying dorsally to the hypostome. The terminology usually
applied to the portions of the diyit of the chelicerae appears to us wrong
and confusing, and we prefer to recognize (1) an "internal article," the
latter bearing a " dorsal process " which is a portion of it (not articulated),
and (2) an " external article " which articulates with the internal article
upon its outer side :

1. Internal a?-^ic/e = "median apophysis" or "immovable finger" of some authors.

Dorsal process = '■'■ internal apophysis " of some authors.

2. External amcZe = " external apophysis" or "free" or "movable finger" of
some authors.

We attach little importance to the description of the chelicerae in routine
systematic work.

DORSAL PB0CBS3

EXTBKNAL ARTICLE;'-.,.

--FORE.
--ARTICULATION

DORSAL PROCESS

PLEXOR TENDON
EXTENSOR TENDON

EXTERNAL ARTICLE

v.. .-PORE.

INTERNAL ARTICLE.

Fig. 136.4.

Fig. 136 5.

THE DIGIT.

Fig. 136 A. Distal extremity of chelicera in dorsal aspect, showing articulation of digit
and the two tendons (black) which move it laterally. B. Digit with articles
separated. Illustrating the structures referred to in the descriptions. Haemaphysalis
punctata ? . (Nuttall, Cooper and Kobinson, 1908, Figs. 7 and 8.)

Palps : articulating antero-laterally upon the basis capituli. Frequently cultri-
form so that when apposed they protect the hypostome and chelicerae.
In ticks possessing long palps four palpal articles can usually be made
out very distinctly. These are numbered 1 to 4, beginning with the basal
article. In some ticks some of the articles, especiallj' articles 2 and 3,

Terminology 129

may be fused. In all Ixodidae the 4tli article is reduced to a small hair-
crowned papilla lying in a cup-like hollow of article 3. The articles which

are of importance in classification are Nos. 1, 2 and 3.
Dorsum = the whole dorsal surface of the body.

Sc\Uum = i\ie "dorsal shield," " Kopfschild," etc., of some authors. The size is

stated by giving the length first, then the greatest width (including the

eyes, if protruding), thus "9 x "7 mm.
Psendoscutu')n = ihiit portion of the $ scutum which corresponds in shape and

position with the $ scutum, and is sometimes outlined by a ridge, puncta-

tions or hairs, without being a definite structure.
Emarginate means hollowed out for the reception of the capitulum. This

portion of the scutum is styled the emargination in the figures.
Scapidae = ih.e anterior angles or "shoulders" of the scutum projecting on

either side of the emargination and included in measurements of the length.
Cervical grooves = t\iQ term commonly applied to the pair of grooves running

backward from the inner angles of the scapulae.
Lateral grooves = grooves running along the sides of the scutum in both sexes.

In the ^ they may be prolonged backward and include one or more

festoons ; they are spoken of as " complete " if they are continuous along

the anterior border of the festoons. -

Marginal grooves run along the sides of the body in the $ , starting at the

postero-lateral scutal border (they correspond to the ^ lateral grooves

which are prolonged backward). The marginal grooves may include several

festoons, or they may be " complete."
Median and postero-lateral depressions, grooves or furrows are more or less

evident in some (^ s and most $ s. They correspond to lines of attachment

of the dorso-ventral bands of muscles.
Median and lateral Jields = 'parts of the scutum lying (1) between the cervical

grooves, and (2) external to the latter.
Festoons are the uniform rectangular areas into which the posterior margin of

the body is divided up in most of the Ixodidae. They are most distinct

in unfed specimens, but disappear more or less in distended females.

The dividing grooves are sometimes referred to as the " intervals " of the

festoons. The festoons frequently have distinct chitinous plates on their

ventral surface. (Figs. 120, 121, etc.)
Foveae = ihe "submedian dorsal porose areas" of Salmon and Stiles, two small

spots often observable near the middle of the scutum in the (J ; posterior

to the scutum in the J . (Figs. 121, 123, 130.)
Marginal fold = 3. raised fold of the body, external to the scutum, as seen in

Ixodes (see Fig. 115).
Venter = the whole ventral surface of the body.

ilenitai orifice : situated anteriorly, posterior to tlie capitulum, and in the

median line. Called vulva for brevity's sake in the 9 •
Api-on : a delicate chitinous flap arising in front of the sexual orifice which it

covei's.
Genital grooves : starting at either side of the genital orifice and running

130 Tetmmology

backward between the coxae, then externally to the anal grooves toward
the posterior body-margin which they frequently attain.

A1UIS situated in the median line posterior to the coxae ; the term applied for
convenience' sake to the external anal apparatus consisting of a more or
less evident ring within which are two laterally-moving valves.

Anal grooves in Ixodes (Prostnata) curve around the anus in front and run
backward toward the posterior margin. Whereas, in most cases, the
grooves fuse in a semicircle in front of the anus ; in other cases they
form an ogive, they are then termed " ogival " (like a gothic arch) ; in some
species the grooves are not continuous anteriorly.

In Ifetastnaia the anal grooves round the anus behind, and in most cases
they run forward and outward toward the genital grooves, which they
may attain. In some cases they are continuous with a postero-median
groove from which they fork anteriorly.

Plates are large dense armour-like chitinous structm-es occurring in Ixodes ^,
not rising above the surface of the body (as do the shields, for instance,
in lihipicephalus, etc.) ; the plates are bounded by the A'entral grooves
above named, or by soft portions of integument. We distinguish 1 pre-
genital, 1 median, 1 anal plate along the median line of the body ; 2 adanal
plates to either side of the anal plate ; and 2 epimeral plates with indistinct
external border extending forward outside the genital groove to near
coxa IV.

Plaques are small chitinized plates occurring, for instance, in some species of
Amhlyomma, and situated veutrally in front of the festoons. (Fig. 135.)

Shields (adanal and accessory) are salient chitinous structures occurring in the
males of Rkipicephalus, Boophihis and Hyalomma on either side of the
anus. A pair, close to the anus, are always present and are termed adanal.
There is frequently a second pair external to these, and known as accessory ;
they are well developed in Boophilus.

Spiracle = the "peritreme," or "stigmal plate," etc., of various authors. A respira-
tory organ situated ventro-laterally posterior to coxa IV. It may be circular,
oval or comma-shaped, and shows a more or less central structure which we
term the macula — which may or may not represent a pore. The punctate
surface may be enclosed by a more or less broad " frame " of chitin which
is incomplete postero-dorsally in comma-shaped forms. The structure of
the spiracle is of use in classification. (Fig. 137.)
Legs : Coxae are the immovable portions upon which the movable articles are
articulated ; of the latter especially the first {trochanter) and last (tarsus)
articles are of importance in classification. All of these may bear spurs,
spines, or teeth in various situations noted in the descriptions. When
coxae are described as bijid, they bear two spurs and are deeply incised ;
when described as trenchant they have a knife-like margin. It is of
importance to note the manner in which the tarsi terminate : if they
taper, are humped, bear ventral spurs, etc. The length of the claws in
relation to the pads, or suckers {pidvilhim), should also be noted. (Fig. 138.)
Punctations are circular depressions dotting the integument, and frequently

Terminology

131

bearing hairs. The parts, scutum, capituhim, etc., on which punctations
occur are referred to as finely or coarsely punctate, etc.
Ornamentation refers to enamel-like coloration which may be present on the
scutum, capitulum and legs, etc. Ticks on which such ornamentation
occurs are spoken of as ornate ; for instance, Amhlyomma and Dermacentor
arc usually ornate.

0OO

Fig. 137, illustrating the spiracles of 15 species of Ixodidae to show the diiJerent forms
they may assume. (Original, G. H, F. N. del.)

All the figures are orientated so as to show their positions on the tick's body : A pointing
anteriorly, V and D ventrally and dorsally respectively.

Spiracle 1.
2.
3.
4.

5.

6.
7—15.

7.
8.
9.

10.

-15.
11.
12.
13.
14.
15.

11-

long axis of spiracle transverse
to body axis.

long axis normal, i.e.
directed forward.

Kound as in /. ricinus ? .

Bluntly oval as in I. boliviensis ? )

Elongate sub-oval as in I. tasmani ? i

,, oval as in I. angustus ?

Ovoid, flattened posteriorly (or sub-triangular) as

in I. aiKjustus i

With slight postero-dorsal protrusion as in Haetnaphysalis hystricis <? .
With distinct darkly chitinized marginal frame "complete" in 7, incom-
plete postero-dorsally in the remaining spiracles, which are comma-
shaped ; or (10) sub-triangular, with rounded angles.
Bluntly ovoid as in Boophibis annulatits ? .
Short comma-shaped as in Dermacentor variabilis ? .

,, ,, ,, Amhlyomma longirostrum ? , with frame broad-

ened dorsally.
Sub-triangular, with rounded angles and with frame broadening much

dorsally as in Amhlyomma geoemydae ?.
More or less elongate comma-shaped spiracles :
As in Rhipicentor nuttalli 3 .

,, Dermacentor reticulatus s , and Rhipicentor hicornis s .
,, Rliipicephalus sanguineus t? , and Amhlyomma versicolor <J .
,, Dermacentor andersoni s .
,, Hyalomma aegyptium i , and Rhipicephalus sp. i.

The "macula" is central in (1), eccentric in the others, being situated ventrally and
anteriorly, and possessing a variable form.

132

Termmologif

Fig. 138, illustrating the tarsi of 10 species of Ixodidae to show different forms of Tarsus
4. (Original, G. H. F. N. del.)

The transverse line indicates the pseudo-articulation.

Tarsus 1.
2.
3.
4.
5.
6.
7.
8.
9.

Tapering gradually as in I. ricinus.

,, ,, 1. -f- signatus.

,, obliquely ,, I. ornithorhynchi.

Humped prior to tapering as in I. hexagonus.
Tapering abruptly as in I. cordifer.

,, to one spur as in Haemaphysalis cornigera.

Ending bluntly with two spurs as in Rhipicephalus masseyi.
Tapering to a long spur as in Margaropus whithemi.

,, abruptly aud bearing two spurs as in Amhlyomma cooperi.

I

133

SECTION II
Genus I. IXODES Latreille, 1795.

Synonymy and Literature.

"Ricinus caninus" Ray, 1710, p. 10 (?/. ricinus).

Acarus Linnaeus, 1746, p. 479, pro parte ; also subsequent authors (dc Geer, 1778 ;
Fabricius, 1794 ; Panzer, 1795). See further under list of si^ecies of Acarus.

Ixodes 1796. Latreille, p. 179.— 1804. Latreille, p. 46.— 1805. Fabricius, p. 351.—
1806. Latreille, p. 155.-1807. Chabrier, pp. 366 et seq. (oviposition).— 1815.
Leach, p. 397.— 1826. von Heyden, p. 610; Risso, p. 182.-1829. Latreille,
p. 287. — 1834. Dug5s, (a), p. 15 (classification). — 1837. Burmeister, p. 579. —
1844. Koch, p. 231 ; Gervais, p. 234 ; Sangalli, p. 831 (effects of bite) ; Gene,
p. 751 (anatomy and biology). — 1847. Koch, p. 20 ; Gervais, p. 351. — 1849.
Gene (translation of Gene, 1844).— 1857. Kolenati, p. 24.-1858. Heller, p. 58
(sexual organs). — 1859. Gervais and van Beneden, p. 460 ; Grube, p. 455. —
1861. Fiirstenberg, p. 208 ; Moquin-Tandon, p. 304 ; Pagenstecher, pp. 1-40
(anatomy, etc.). — 1863. Gerstaecker, p. 344. — 1866. Lucas, p. Ivii ; VerrilU,
p. 116. — 1877. Canestrini and Fanzago, pp. 110, 178 et seq. (brief description) ;
Murray, pp. 187 et seq. ; Conil, p. 25.— 1878. Conil, p. 99.-1880. Megnin, (a),
pp. 121, 320; (6) p. 603 ; Taschenberg, p. 150 (mouth-parts, 0 2).— 1881. Haller,
(a), p. 165 ; (5), p. 380 (mouth-parts).— 1882. Haller, p. 309.— 1883. Braun,
p. 211.-1885. Berlese, p. 131 ; Raymondaud, p. 129.— 1886. Ludwig, p. 612.
—1888. Railliet, p. 496 ; Neumann, p. 89.— 1889. Berlese, fasc. Iv, n. 6 (classi-
fication).— 1890. Canestrini, pp. 475 et seq., p. 492 ; Blanchard, p. 323. — 1891.
Batelli, pp. 100 et seq. (respiration, mouth-parts, etc.) ; Trouessart, p. 290. —
1892. Neumann, (a), p. 92 ; (6), p. 96 ; Canestrini, p. 581 ; Efisio', p. 256 ;
Trouessart, pp. 20-47 ; Lewis, p. 449 ; Marx, (6), p. 232 (mouth-parts, classi-
fication) ; Bernard, p. 289 (resjiiration) ; Michael, pp. 204, 447 (mouth-parts).
—1893. Railliet, p. 706.— 1894. Brandes, p. 405 (anatomy).— 1895. Megnin,
p. 354 ; Braun, p. 257.-1896. Oudemans, p. 191 ; Osborn, p. 262.-1897.
Neumann, pp. 283, 325, 360, 384 ; Supino, (a), p. 241 (classification).— 1899.
Neumann, pp. 108 et seq. ; Morgan i, p. 138 ; Wheler; pp. 5 et seq. (classification,
etc.) ; Nuttall, p. 402 (effects of bite).— 1900. Ward, (a), p. 196 ; (6), p. 436 ;
Brucker, p. 423 (mouth-parts). — 1901. Salmon and Stiles, p. 459 (classification) ;
Jourdain, \>. 142 (mouth-parts). — 1902. Neumann, p. 115 (genus Eschatocephalv^

^ Inaccessible, cite^ in bibliography by Salmon and Stiles, 1901.
'^ 0 denotes that the paper contains nothing worth noting.

134 Genus Ixodes

maintained^ and that of Ceratixodes created). — 1904. Neumann, p. 444 {Ixodes
to include 3 subgenera : Eidxod.es, Ceratixodes and Eschatocephahis, vide
infra). — 1905. Lahille, pp. 12, 28 (classification). — 1906. Neumann, p. 195
(coitus and sexual dimorphism). — 1907. Donitz, p. 88 (useful) ; Pocock, p. 190
(classification) ; Himter and Hooker, p. 54. — 1908. Nuttall, pp. 385 et seq.
(structure, biology, oviposition) ; Bonnet, p. 253 (descriptions condensed from
Neumann ; figures original, but poor) : Banks, p. 55 ; Howard, p. 92. — 1909.
Rohr, p. 135 (quotes Neumann, also with regard to species enumerated);
Blanchard, pp. 80 et seq. (brief descriptions and lists of species, largely drawn
from Neumann and other authors since). — 1910. Donitz, pp. 400, 432 (classifi-
cation discussed) ; Stiles, pp. 12-17 (classification) ; Brumpt, Fig. 353 (copulation).

Cynorhaestes Hermann, 1804, p. 66 {pro parte) and subsequent authors. — 1826,
Risso, p. 183 ; Heydeu, p. 610. — 1831, Treviranus, p. 188, after which the name
fell into synonymy. (Referred to as Cynorrhaestes Hermann by Fabricius,
1805, p. 351, and Dug5s, 1834 c, p. 33; as Cynoraesthes by Latreille, 1829, p. 287 ;
as Cynochaestes by Meguin, 1880, p. 120 ; as Cynorhoestes by Trouessart, 1891,
p. 290.)

Crotonus Dumeril, 1822, p. 56. — 1829, p. 401 (C. ricinus = I. ricinus (Linn.)).

Haemalastor Koch, 1844, p. 223 ; 1847, p. 49 (type, H. longirostris). Used partly
as equivalent to Esckatocephalus {vide infra) by Kolenati, 1857, p. 437 ; 1860,
p. 578 ; Karsch, 1880, pp. 141 et seq. ; Megnin, 1880, p. 120 {Haemalostor) [sic\;
and Neumann, 1899, p. 178.

Note : Haemalastor was established by C. L. Koch to include one species
{longirostris), of which he only knew the 9- Neumann, 1901, p. 290, dis-
covered, however, upon examination of the type, that it possessed eyes, in
consequence of which he referred it to the genus Hyalomma. Consequently,
the genus Haemalastor disappears, together with the single species upon
which it was founded. For list of species referred to Haemalastor by various
authors, see List of condemned species.

Eschatocephalus Frauenfeld, 1853, p. 55. Recognized by L. Koch, 1877, p. 150;
Joseph, 1882, p. 16 ; Neumann, 1899, p. 179, and 1901, p. 290 (name revived),
and all subsequent authors except Donitz, 1910. For a list of species referred
to Esckatocephalus by various authors, see List of condemned species.

Note: Esckatocephalus was retained as a genus by Neumann (1902, p. 116)
until 1904, p. 445, when he made it a subgenus oi Ixodes, because he considered
that the generic characters were insufficient to maintain the generic rank, the
differences "being only specific." He states that "the ^, by virtue of its
claviform palps, which are circular on cross section, differs sufficiently from
other Ixodes to place this species (meaning /. vespertilionis) in a subgenus."
Still retained as a genus by Blanchard, 1909, p. 96. We are of the same
opinion as Donitz, 1910, p. 400, that there is no valid reason for retaining the
genus, and we have condemned Esckatocephalus as a subgenus.

Dermanyssus Kolenati, 1857, p. 10 = Ixodes {vide List of condemned species).

Sarconyssus Kolenati, 1857, p. 21 (various " species " = /x'oo?es vespertilionis. See
List of condemned species).

Hyalomma Pickard-Cambridge, 1878, p. 222, Hyalomma puta = Ixodes putus q.v.

Ceratixodes Neumann, 1902, p. 115. The main points whereby the genus can be

Key to Species 135

distinguished were originally stated to be : the convex inner surface of the
palp, and the pointed extremity of the third palpal article in the ^ ; the
slightly hollow internal surface and swollen ends of the palps in the $ ; anal
grooves of $ as in Ixodes, but absent in the $ ; anal and adanal shields
in the ^ . Neumann, 1904, p. 444, subsequently recognized that the 9 (-^•
piitus q.v.) differed so slightly from other $ Ixodes, that the genus could not
be retained ; the $ was found to possess an anal groove and the ^ only difters
from other ^ Ixodes in respect to its palps. He therefore degraded Ceratixodes
to a subgenus. Still retained as a genus by Blanchard, 1909, p. 97.
Euixodes Neumann, 1904, p. 444. A subgenus founded to include all Ixodes other
than Cei'atixodes and Eschatocephalus (the latter genera reduced to subgenera).
Since we see no reason for retaining the latter subgenera, Euixodes falls into
the synonymy of Ixodes.

Generic Characters. Anal grooves embracing the anus anteriorly,
and usually uniting in a pointed arch or the arc of a circle. Scutum
always inornate, and without eyes. No festoons. Spiracles round or
oval. Tarsi without spurs {except in Ixodes putus). The male scutum
is aliuays surrounded by a prominent marghial body-fold, and the male
venter presents seven non-salient chitinous plates, namely, a 2iregenital,
a median, an anal, two adanal and tivo epimeral plates. The genital
aperture is comparatively posterior in both sexes, and in the distended
female the legs occupy a very anterior position. There is marked sexual
dimorphism as regards the hypostome. Figs. 115-119.

N.B. In most species the second article of the palp is narrowed at
the base, leaving a space between it and the hypostome, and this gives
the ticks a peculiar facies easily recognized by the naked eye. There
are, however, exceptions.

Type species : Ixodes ricinus (Linnaeus) Latreille, 1804.

Key for the determination of species of Ixodes.
Males.

There are 21 species of males recorded of which one {pratti, p. 174)
has not been adequately described.

PAGE

[Body with 5 posterior brushes of hairs . , . putus 256

I „ without such hairs . . . . .1

Legs inordinately long. ..... vespertilionis 272

,, moderately „ . . . . .2

[Basis capituli with dorsal punctate excavation . coxaefurcatus 270
I „ „ without ,, „ .3

136

3.
4.
5.
6.

8.

9.

10.

11.

12.

13.

14.

15.

16.

17.

18.

Genus Ixodes

Dorsum showing two lateral folds
„ „ one „ fold

(Anal grooves divergent
( „ ,, convergent

jAnal grooves convergent

\ „ „ parallel or divergent

[Anal grooves approaching but not uniting behind
( „ „ uniting behind, . . . .

[Anal plate oval, basis capituli with median point
I ventrally .......

1 Anal plate racket-shaped, basis capituli tricuspid
[ ventrally .......

[Coxa IV with long internal spur .
I ,, „ without long internal spur

[Tarsus 4 humped some distance from end .
I „ „ tapering

[A single distinct spur on coxa I .

(Two spurs or none .....

[Median ventral plate about as broad as long .
[ „ „ „ much longer than broad

Coxa I with two spurs ....

„ „ without spurs ; at most slightly pointed

[Both spurs very long and close together

I „ „ otherwise .....

[Both spurs short, subequal ....
(Inner spur long and strong, outer short.

Strong cornua ; anal grooves short, divergent
No cornua ; anal grooves long, parallel

rTarsi humped ....
\ „ tapering ....

Coxae trenchant .....
„ not trenchant ....

[Punctations very fine, equal

( „ unequal, nearly obsolete

4

PAGE

5

loricatus

266

holocyclus

235

6

10

pilosus

schillingsi

cordifer
9

rasus
ugandanus

11
12

hexagonus
ricinus

13
16

gigas
14

15

boliviensis

minor
angustus

canisuga
17

tenuii'ostris
18

cavipalpus
ruhicundus

221

238
233

228
230

178
147

203

166

164
195

210

246

193
204

Kcif to Species

137

Females.

1.

2.

8.
9.
10.
11.

12.

There are 48 species known by their females.

[Legs inordinately long (on bats) .

(Legs moderate ......

[Anal grooves horseshoe shaped or closed behind
I „ „ sub-parallel or divergent

[Anal grooves horseshoe shaped .

I „ „ closed behind ....

(Coxa I with two long sharp spurs ; well-marked
auriculae (Madagascar) ....
Coxa I otherwise .....

'Coxa I with a long internal spine ; auriculae (on
monkeys, Africa) .....

Coxa I with no internal spine ; no auriculae (on
cattle, S. Africa) .....

JAnal grooves a circle (Africa)

I „ „ an oval, pointed behind .

Coxa I with sharp internal spur .
,, „ without internal spur

Scutum broader than long, palps short, with
article 1 much enlarged ....

Scutum longer than broad, palps long, normal
(Australia) .......

[Coxae practically unarmed ....
[Some of the coxae with distinct spurs .

Scutum broadest quite in front (on marine birds)
„ ,, near middle or posteriorly

fScutum broadest behind the middle

( ,, „ in middle or a little anteriorly

fScutura broader than long ....
I „ longer than broad ....

(Palps with article 1 greatly enlarged on ventral
aspect .......

Palps with article 1 small ....

vespertilionis 274
1

2

8

3

iimatus
4

226

schillingsi 239

pilosus

222

7risus 229

6

ugandanus 232

7

australiensis 250

holocyclus

236

9

20

piitus

258

10

11

14

12

13

tasmani

244

ornithorhynchi

242

10

138

Genus Ixodes

13.

14.
15.
16.
17.

18.

19.
20.
21.

22.

23.

24.

25.
26.

{Anal grooves short, strongly divergent (Europe,
on small mammals) ......
Anal grooves long, sub-parallel (Australia) .

[Article 1 of palps enlarged, cup-like (Australia) .

1 „ „ „ normal

[Anal grooves short, divergent ....
I „ „ long, sub-parallel ....

[Scutum sub-circular; slight cornua
I „ elongate ; no cornua (on bats)

(Sharp spurs on trochanters (Africa)

(No trochantal spurs ......

/Coxae trenchant; scutum smooth, with fine punc-
I tations (Africa) ......

Coxal borders rounded ; scutum rugose, with
large punctations ......

{Scutum slightly rugose ; widest rather in front of
middle (Europe)
Scutum very rugose ; widest in middle (America)

(No internal spur, but a blunt external spur on
coxa I ....... .
Coxa I with internal spur .....

{Spurs on trochanters (? on birds, Nightingale
Island) ........
No spurs on trochanters .....

/Porose areas large, confluent, occupying most of

basis capituli (on marine birds)
Porose areas tranversely elongate, separate (on

marine birds) .......

^Internal coxal spur long .....

J „ „ „ short; a slight external spur

I, or tooth ........

[A strong external spur .....

(External spur short or absent ....

(Both spurs long, parallel, sub-equal (India)
(Internal spur much the longer ....

[Anal grooves short, anus very postei'ior (Mexico).
\ „ „ long (Brazil)

tenuirostris

247

fecialis

248

vestitiis

252

15

16

17

japonensis
simplex

208
207

cavipalpus
18

194

ruhicundus

206

19

camsuga
texanus

212
214

21

23

percavatus
22

220

unicavatus

264

signatus

261

24

33

25

27

acutitarsus

202

26

hicornis

186

fuscipes

169

27.
28.

29.

30.

31.

32.

33.
34.

35.
36.

37.

38.
39.
40.

41.

Key to Species

Cornua well marked, sharp . . . . .
„ faint or absent . . . . .

jPorose areas large, near together (N. America) .
I „ „ small, far apart (America)

(Hypostome very long and pointed ; anus very
posterior (Italy)
Hypostome normal

{Punctations on scutum large and few; coxal spine
very long (Sumatra)
Punctations small and numerous
[Anal grooves short ; auriculae strong
( „ „ long; no auriculae .

{Porose areas transversely elongate (Madeira)
(possibly identical with ricinus)
Porose areas less elongate .
J Scutum nearly as broad as long .
I „ elongate, distinctly longer than broad
[Anal grooves pointed in front ; tarsi humped
I „ ,, rounded in front ....

{Scutum oval; auriculae, but no cornua (America)
„ cordiform ; cornua, but no auriculae
(America) .......

(Article 1 of palp with anteriorly directed spur (oi
birds, Brazil) .......
No such spur .......

(Basis capituli winged, pointed laterally (S.
America) ........
Basis capituli normal ......

[Tarsus 4 humped, or narrowing abruptly

I „ „ tapering more or less gently

[Anal grooves rounded in front ....

I „ ,, pointed „„....

[Basis capituli elongate .....

\ „ „ broader than long ....

''Slight sharp cornua, porose areas oval, longer than

broad (Chili) .
No cornua, porose areas much broader than long
(N.Zealand)

139

PAGE

28

29

sculptus

171

diversifossus

163

acuminatus

161

30

spinicoxalis

171

31

holiviensis

167

32

nigricans

159

ricinus

1.50

34

36

hexagonus

179

3.5

dentatus

162

pratti

174

auritulus

187

37

loricatus

267

38

39

43

40

42

fossulatus

201

41

stilesi

216

neumanni

218

10—2

140

Genus Ixodes

42.

43.

44.

45.

46.

'Scapulae long and sharp ; basis capitiili broadest
anteriorly (Mexico) ......

Scapulae short and rounded ; basis capituli tri-
angular (America) ......

(Scutum nearly twice as long as broad (1"3 x "7 mm.)
with rounded sides (S. America)
Scutum less elongate, with angular sides

/Anal grooves not reaching posterior border ;
J strong white hairs on scutum (on birds)
I Anal grooves reaching posterior border ; scutal .
V hairs short or absent .....

Basis capituli broader than long ; porose areas

almost confluent . , .
Basis capituli longer than broad ; porose areas

separate .......

Sharp auriculae ; anal grooves rounded in front
(on mouse, Christmas Island) ....

No auriculae ; anal grooves pointed in front
(America) .......

ruhidus

175

marxi

173

■minor

165

44

hrunneus

190

45

caledonicus

198

46.

nitens

185

angustus

196

Nymphs.

N.B. Of the 22 nymphal forms recorded there are 19 species
described sufficiently to be included in the following key. Not included
are fuscipes, p. 170 ; hrunneus, p. 190 ; siniplex, p. 208.

2.

5.

With legs inordinately long
„ „ otherwise

(Scutum without cervical grooves
1 ;, with

[Scutum with lateral grooves

1 „ without lateral grooves .

(Coxae unarmed ....
I „ armed ....

(Palp with article 1 large and pointing forward
I „ „ „ „ otherwise

(With anal grooves fused to a point behind .
( ;, „ ., otherwise . ,

vespertilionis 27.:

tenuirostris

2

3
9

vestitus
4

angustus
5

holocyclus
6

248

254

197

237

Key to Species

141

6.

7.

8.

9.

10.

11.

12.

13.
14.
15.
10.
17.

[With anal grooves in the form of a horseshoe
I „ „ „ parallel or divergent

[With long palps, concave externally .

I „ palps otherwise ....

[With tarsus 4 tapering

1 ,. „ ,) humped

[With coxae unarmed .

\ „ „ armed

[With tarsus 4 humped

I „ „ ,, othervi^ise

[With very long scutum

(With short, very broad scut

{Coxae with small blunt protuberance postero
externally
Coxae with marked spurs postero-externally

[Without auriculae ....

[Otherwise ......

[Trochanters with postero-external spur
( „ without postero-external spur

With anal grooves horseshoe-like

„ „ „ otherwise

With scutum longer than broad .

,, „ as long as broad

With scutum about as broad as long

„ „ narrow, elongate

17

7

holiviensis

169

8

ricinus

151

ruhidus

176

10

12

canisuga

213

11

putus

259

ornithorhynchi 242

hexagonus

181

13

unicavatus

265

14

15

neumanni

219

schillingsi

241

16

signatus

263

caledonicus

199

pilosus

224

loricatus

268

Larvae.

N.B. There are 14 species whose larvae have been recorded, three
of which (fuscipes, p. 170; branneus, p. 190; loricatus, p. 268) have not
been adequately described nor figured. The following key, therefore,
relates to but 11 species.

1.

[With legs inordinately long

( „ „ otherwise ....

[Scutum with cervical grooves absent .
I „ » „ „ present .

vespertilionis 275
1

tenuirostris 248

2

142 Genus Ixodes

2,

3.
4.
5.
6.

jScutum with lateral grooves .... vestitus 255

\ „ without lateral grooves . ... 3

(Coxae unarmed ....... 4

( „ with spurs ...... 7

[Palp with article 1 large and pointing forward . angustus 197

] „ otherwise ....... 5

[Capitulum with dorsal ridge pointing out laterally pilosus 224

( „ otherwise . . . . . .6

("Basis capituli with latero- ventral protuberances . canisuga 213

( „ „ „ ventral ridge . . . hexagonus 181

w (Basis capituli without auriculae .... unicavatus 265, 266

( „ „ with auriculae .... 8

{Scutum with cervical grooves diverging widely

behind 9

Scutum with cervical grooves otherwise . . signatus 263

Coxa IV with two spurs caledonicus 200

,, „ without two spurs .... ricinus 152

143

SPECIFIC DESCRIPTIONS

OF VALID SPECIES OP IXODES

AND OF THEIR

VARIETIES AND SUBSPECIES

1. IXODES RICINUS^ (Linnaeus, 1746 and 1758).

Pis. IV and VII, Text-figs. 115, 139-148.

Synonymy: not Reduvius Charleton, 1668, p. 49.

IRicimis caninus Ray, 1710, p. 10 (Synon. in Walckenaer and Gervais, 1844,

p. 236). ?.
not Acarus reduvius Linnaeus, 1746, p. 479. >S>/st. naiur. 1758, 10th edit.,

p. 616; 1788, 13th edit., p. 2925.
Acarus ricinus Linnaeus, 1746, p. 480 ; 1758, p. 615 ; 1788, p. 2925.

Eegarding the synonymy of this species, Neumann (1901, pp. 281-282) writes
as follows :

"Linnaeus, in his Fauna suecica (1746), describes under No. 1192 an Acarus
ovinus, which he makes synonymous with Reduvncs Charleton {Onomasticon zooicum,
1668) and Pediculus ovinus Rajus (Historia insectorum, 1710). In his Systema
naturae (10th edit., 1758, p. 615), Linnaeus changes his Acarus ovinus to Acarus
reduvius, and gives it under this new name as No. 1966 in Fauna suecica (2nd edit.,
1761), retainiiig his synonymies. On referring to the text and the figures of the
writers cited by Linnaeus, it is clearly evident that the writers mentioned were
dealing with Melophagus ovinus and not one of the Acari. It is consequently due
to an error that Linnaeus placed Reduvius or Pedicidus ovinus in his genus Acarus.
Whatever may have been the form to which he desired to attach the name Acarus
reduvius, this name loses all taxonomic value, and it is necessary to revive the
name of Ixodes ricinus {Acarus ricinus Linnaeus), which applies to the same species
of tick and does not lead to confusion."

Acarus ricinoides de Geer, 1778, vii, p. 98 ; p. 101 {reduvius).
Ixodes ricinus Latreille, 1804, i, p. 156 (?).
Ixodes reduvius Latreille, 1804, p. 51.

^ According to Dubreuilh, 1900, p. 450, the word Ricinus, used to signify a tick by
Pliny, gave its name to the castor-oil plant, as stated by Dioscorides, iv, 161, because the
seeds of the latter resemble a tick {i.e. the gorged females).

144 Genus Ixodes

Cynorhaestes reduvms Hermann, 1804, p. 65.

Ci/norhaestes ricinus Hermann, 1804, p. 66.

Ixodes megathyreus Leach, 181.5, xi, p. 398.

Ixodes hipunctatus Risso, 1826, v, p. 183.

Cynorhaestes hermanni Risso, 1826, v, p. 183.

Cynorhaestes megathyreus (Leach) Risso, 1826.

Crotonus ricinus Dumeril, 1829, liv, p. 401.

Ixodes traheatus Audouin, 1832, xxv, p. 420 ( 9 )•

Ixodes plumheus Duges, 1834, i (2), p. 20.

not Ixodes plumheus Dugfes, 1834 c, PI. VII, Figs. 7-12 { = Rhipicephahis s^.).

Ixodes reduvius Hahn, 1834, ii, p. 62.

Ixodes fvscus Koch, 1835, Heft xxxvii.

Ixodes lacertae Koch, 1835, Heft xxxix ; 1844, p. 234, larvae and nymphs, fide

Canestrini, 1890, p. 497.
Ixodes rufus Koch f I^^^^^' 1^35, Heft v, No. 11, Heft xxxvii. No. 8 ; 1844,
Ixodes sidcatus Koch -;' P" ^32 ; 1847, p. 21. The types examined by Neumann
Ixodes sciuri Koch | (^^^l' PP" 281-289) proved to be 7. ricinus nymphs, all

V collected in Germany.
Ixodes reduvius Linu., in Koch, 1835, Heft v, No. 11.
Ixodes pustularum Lucas, 1866, vi ; Bidl., p. Ivii.
Ixodes fodiens Murray, 1877, p. 191. As synonym in Neumann, 1899, p. 122.

Neumann, 1904, p. 45, rightly states that this name is Murray's, not

Megnin's, as stated by Murray, 1877, and quoted from the latter by

Salmon and Stiles, 1901, p. 465.
Ixodes ovatus Neumann, 1899, p. 116; in part, see Neumann, 1904, p. 452,

where he states the lot of ovatus was subsequently found to comprise

ricinus. — Donitz, 1905, p. 132.
Ixodes ovatus Neumann, 1899, as modified by Neumann, 1904, p. 452 = Zro(/es

ricinus var. ovatus (Neumann) Nuttall and Warburton, 1911.
Ixodes reduvius (Linnaeus) in Railliet, 1895, p. 706 ; Neumann, 1899, p. 112,

and many subsequent writers.

In N. Tyne Valley, where they occur commonly on sheep, they are

locally known as "face ticks," since they are found in this situation on

sheep (Wheler, 1899).

Styled the "Castor-bean tick" by Salmon and Stiles, 1901, p. 463, and

several subsequent authors. The castor-bean is however much more like a

gorged Amblyomma $. (See footnote, p. 143.)

The synonymy of this species might be extended almost indefinitely. We
confine ourselves, however, to the names concerning which little doubt can be
entertained.

Iconography: De Geer, 1778, PI. V {Ac. ricinoides), Figs. 16-19, ? dorsum, part
of capitulum, tarsus ; PL VI {Ac. reduvius), Figs, 1-8, $ dorsum, spiracle
(recognized as such), capitulum, tarsus, ^ and $ in copula (recognized as
coitus), (^ dorsum, capitulum. Excellent, considering the date ; better than
some figures printed a hundred years later. — Audouin, 1832, PI. XIV, Figs.
3rt-?i {traheatus).— Dngbs, 1834, PI. VII, Figs. 7-12 {plumheus).— Rahn, 1834,

TICKS

PLATE IV

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

Fig. 6.

Fig. 7.

Figs. 1-7. Ixodes ricinus.

Fig. 1. cf dorsum, x 12. (Wheler, 1906, Fig. 16.)

Fig. 2. (J venter (mounted specimen). (Original, E. G. Wheler, phot.)

Fig. 3. ? dorsum, x 12. (Wheler, 1906, Fig. 17.)

Fig. 4. ¥ venter, x 9. (Wheler, 1906, Fig. 11.)

Fig. 5. o dorsum, x 12. (Wheler, 1906, Fig. 18.)

Fig. 6. L dorsum, x 12. (Wheler, 1906, Fig. 19.)

Fig. 7. ? with one ,? coupled and a second tf "waiting his turn." (Wheler, 1906, Fig. 36.)

Reprinted (Fig. 2 excepted) from the original blocks illustrating the paper by
E. G. Wheler, Journ. Aijric. (Cdnibridge), Vol. i.

/. riclnns 145

II, PI. LXVI, Fig. 152 (c^).— Koch, 1835-44, Heft 5, Fig. 11 (reduvtus ?);
Heft 37, Figs. 5-7 {ricinus S, $s), Fig. 8 (scmri) ; Heft 39, Fig. 11
(lacertae), Figs. 3, 4 (fuscus S^ 9), Fig- " {ruftcs, 9 )•— Pageiistecher, 1861, ii,
PL I, Figs. 1-10 ; PI. II, Figs. 1-22. Really excellent figures of anatomy. —
Megnin, 1867, i, PI. IV. — Murray, 1877, Fig. 2 (fodieiis; figure useless).—
Haller, 1882, PI. V, Fig. 4 (rici)ms).—BrAun, 1883, Figs. 55, 56 (ricinus).—
Railliet, 1886, p. 348 (ricinus). — Aurivillius, 1886, Figs, a and 6 of $ ; very
poor. — Berlese, 1888, fasc. XLix, PI. VI {reduvius, $ dorsum, venter, capitulum
and scutum, etc. ; poor) ; 1889, fasc. LV, No. 16, giving generic characters
of Ixodes : (^ venter and spiracle ; 9 capitulum in ventral aspect, capitulum,
scutum, spiracle ; larva, ventral aspect ; a digit. Outline figures, recognizable
as ricinus. — Neumann, 1888, Fig. 47. — Canestrini, 1890, PI. XLIII, Figs. 1-1 ^r
(/. reduvius) : ^ venter, hypostome ; 9 dorsum (fed and unfed), part of
capitulum, digit, etc. (very poor). Figs. 3-3 & '■^ Phauli.vodes rufus (Koch)"
dorsum, digit, hypostome ; scarcely recognizable. — Blanchard, 1890, Figs. 592-
595 (ricmMs).— Neumann, 1892, Figs. 53-55.— Megnin, 1892, Fig. 4 ; outline
figures of egg, larva, nymph and female, with details all very inaccurate. —
Mosler and Peiper, 1894, Fig. 110 (ricinus). — Neumann, in Railliet, 1895, Fig.
481 ; i-eproduced V)y Salmon and Stiles, 1901, Fig. 222, and by other authors ;
(^ venter, capitulum, tarsus 4 and foot. Somewhat schematic (reduvius). —
Gene, in Railliet, 1895, Fig. 480 ( $ ovipositing, original figure by Gene). —
Braun, 1895, Figs. 127, 128 (ricinus). — Megnin, 1895, figs, worthless, of different
stages.— Osborn, 1896, Fig. 155; poor.— NilesS 1898, PI. IV, Figs. 3-6 (ricinus).
-Morgan!, 1899, Pis. VII, VIII, Figs. 1-5 (ricm«s).— Neumann, 1899, Fig. 1,
^ 9 digits (reduvius) ; Figs. 2, 3, hypostome and digit of ^ and 9 (ovatus), here
reproduced. — Wheler, 1899, pp. 38, 39 (reduvius), various stages, ^, 9i o and
larva, from photographs of mounted and unmounted specimens ; somewhat
too freely retouched. — Lewis, 1900, figures illustrating process of copulation. —
Salmon and Stiles, 1901, coloured Plate XCIII, Figs. 223, 224, 9 (gorged),
much enlarged, dorsum and venter ; PI. XCIV, Figs. 225, 226, 9 capitulum in
ventral aspect, capitulum and scutum, much enlai'ged (they omit lateral grooves
on scutum). — Kossel, Schiitz, Weber and Miessner, 1903, Pis. I-III, 18
excellent photomicrographs of all stages, with details of structure. — Mohler,
1905, PI. I, Figs. 2, 2a, 26 (9 sketchy, coloured).— Nordeuskiold, 1905, text-
figures, illustrating anatomy and histology. — Wheler, 1900, Pis. VII, VIII,
reproductions of photographs of (a) mounted specimens (^ and 9 )> poor ; and
(b) of unmounted ( (^ , 9 dorsum and venter, o and larva ; here reproduced) ;
Pi. X, Fig. 38, shows an abnormality : a 9 minus capitulum ; Fig. 36 ( (J and
9 in copula) ; other figures illustrate gorged 9 > ^^^^ ? ovipositing. — Donitz,
1907, PI. VI, Figs. 34-36, 38, ^ and 9 capitula, ^ venter ; good.— Banks,
'1908, PI. II, Figs. 9, 10, 13 ; PI. IX, Fig. 5, brief description of 9 only; figures
relate to 9 '^^d ^^'^ poor.— Bonnet, 1908, p. 254, Figs. 15, 16; poor. —
Nordeuskiold, 1908, Pis. XXVI-XXVIII, Text-figs, a and 6; histology, im-
portant.— Nuttall, 1908, Figs 5-7, o, larva, ^, reproduced here. — Blanchard,
1909, p. 91, Figs. 102-105 (after Neumann and Bonnet).— Samson, 1909 a,
Pis. IX-XII, 18 text-figs, (anatomy, histology and biology); 1909b, PI. IX,

1 Inacces.sible, cited by Salmon and Stiles, 1901.

146 Genus Ixodes

7 text-figs, (spermatogenesis). — Braun and Liihe, 1910, Figs. 89, 90. Original,
^ venter and 5 capitulum in ventral aspect. Good representations after
drawings by A. Dampf.
Literature : 1710. Ray, p. 10 {Ricinus caninus). — 1746. Linnaeus, p. 480 {Acarus
ricmus).— 1758. Linnaeus, p. 615 {Acarus ricinus, not Ac. reduvius; latter
confused with Melophagus ovinus). — 1778. de Geer, pp. 98, 101 {Acams
ricinoides, Ac. reduvius ; mechanism of bite described, coitus, etc.). — 1804.
Hermann, pp. 65, 67 {Cynorhoestes reduvius and C. ricinus). Latreille, p. 51
(/. reduvius) ; p. 156 (/. ricinus)^. — 1805. Fabricius, p. .351 (/. ricinus). —
1807. Chabrier, p. .366 ; Illiger % p. .370.-1815. Leach, p. .398 (/. megathyreus
on hedgehogs and dogs); p. 397 (/. ricinus on dogs). — 1817. Miiller^, p. 278
(7. ricinus). — 1826. von Heyden, p. 610 ; Risso, p. 18.3 (/. hipunctatus,
Cyn. hermanni, Cyn. megathyreus (Leach)). — 1829. Dumeril, p. 401 (Crotonus
ricinus) ; von Heyden, p. 288. — 1831. Treviranus, p. 191. — 1832. Audouin,
p. 420 (/. traheatus). — 1834. Hahn, p. 62, 0^; Dugfes, p. 20 (/. plumbeus). —
1835-44. Koch, Heft 37, No. 5 (S); H. 5, No. 11 ; H. 37, No. 8 (/. sciuri);
No. 3 (7. fuscus on deer) ; No. 7 (/. rufus, 9 , on deer) ; No. 1 1 (/. lacertae,
o on Lacerta agilis, Munich). — 1837. Burmeister, p. 579 (/. marginatiis). —
1844. Walckenaer and Gervais, p. 236 ; Gene, pp. 1 et seq., a very valuable
contribution to the biology and structure of ricinus ; the first author to describe
oviposition in ticks. — 1847. Koch, p. 21 . (/. sciuri and /. fuscus) ; p. 20 (/.
ricimis) ; p. 22 (/. lacertae). — 1850. von Siebold, p. 546, passages quoted by
Leydig, 1855, p. 468 and Heller, 1858, p. 311 re spermatozoa. Also cited by
Samson, 1909, p. 216. — 1855. Klichenmeister, p. 422 (mere cited description
of $ /. ricinus).— ISbl . Kolenati, p. 24.— 1858. Milner^, pp. 6, 13.-1859.
Gervais and van Beneden, p. 411. — 1861. Moquin-Tandon, p. 302. Pagen-
stecher, pp. 1-38, important regarding structure and biology ; figs, often
reproduced since, thus : Claus, 1887, p. 470, anatomy of body, and Braun,
1906, p. 364, etc.— 1863. Gerstaecker 2, p. 344.-1864. Cox, p. 82 (on dogs
and ferrets, but may have been hexagonus). — 1866. Lucas, p. Ivii (/.
'pustidarum).—l8&J. Megnin, p. 107.-1869. Packard^, p. 613.-1870. Verrill2,
p. 118.-1876. Wagner, p. 129, 0.-1877. Murray, p. 191 (/. fodiens) ;
Canestrini and Fanzago, pp. 110, 180 ; Conil, p. 26. — 1877. Murray, pp. 190,
193 (/. fodiens, vide synonymy); quotes Lucas, 1866, and Megnin, 1867. —
1878. ConiP, p. 100.— 1879. Cobbold, pp. 267, 350.-1880. Haller, p. 38 ;
Megnin, p. 127 ; Oudemans, p. xvii ; Taschenberg, p. 151, 0. — 1881. Bertkau,
p. 145 (copulation and oviposition) ; Haller, (a), p. 165. — 1882. Haller, p. 309.
■ — 1883. Braun, pp. 211 et seq.; van Beneden, p. 142, 1 ricinus, penetrating
beneath the skin in man ; thinks ticks do so normally ! — 1885. Johannessen,
p. 401 (ettects of bite on man).- — 1886. Railliet, p. 497. Aurivillius, {a), p. 105,
3 figures of ? (poor); (6), p. 139, same paper as preceding ; tick beneath the
.skin of a fox. Ludwig, p. 612. — 1888. Neumann, p. 90 ; Berlese, fasc. xlix,
n. 6; Winkler, p. 113 (anatomy). — 1889. Berlese, figures. — 1890. Canestrini,
pp. 474 et seq. ; Blanchard, pp. 324 et seq. — 1891. Blanchard, p. 689 (penetration

1 Most authors after this date refer to the species as either ricinus or reduvius.

- Cited in bibliography by Salmon and Stiles, 1901, unimportant.

3 The sign 0 denotes that the publication referred to contains nothing worth noting.

/. ricinus 147

beneath human skin, excision of a $ ricinus) ; BatelH, pp. 78 et seq. (physiology
of digestion). — 1892. Neumann, p. 95 ; Megnin, p. 32 ; Efi.sio', p. 256 ; Janson
and Tokisliige', p. 349.-1894, iNIosler and Peiper, p. 320 ; Dolly i, p. 1000.—
1895. Braun, p. 257 ; Neumami, in Railliet, p. 707 (figs.) ; Railliet, p. 706 ;
Megnin, p. 364 ; poor description and worse figures. — 1896. Osborn, p. 262,
brief mention; Oudemans, p. 191; Schneidemiihl, p. 339; Dubreuilh and
Beille, p. G9, O.— 1897. Wood and Fitzi, p. 346.— 1898. Nilesi, pp. 29, 45 ;
Weich.selbaum, \). 325. — 1899. Morgan, p. 129, 0 ; Neumann, p. 116 (/. ovatas
in part), p. 112 (/. reduvius) ; Wheler, pp. 38, 49 (biology and structiu-e) ;
MossoS p. 20; Nuttall, 1899, p. 42, cites Dubreuilh, 1838, Despri;s, 1867,
and others re effects of bite. — 1900. Kossel and Weber, relation to redwater
in Finland. Ward, (a), p. 203 ; (6), p. 436. Lewis, p. 383 (copulation observed
and figured'. — 1901. Salmon and Stiles, p. 262 (translation of Neumann) ;
Neumann, p. 281 ; Lewis, p. 154 (mouth-parts) ; Perroncito, p. 562 (effects of
bite). — 1902. Kossel, pp. 853 et seq., relation to redwater in Finland and
Germany ; Wheler, pjj. 1-20, biology and structure. — 1903. Kossel, Schiitz,
Weber and Miessner, pp. 39-50, relation to redwater, structure and biology. —
1904. Megnin, p. 569. — 1905. Nordenskiold, p. 485, figs, (excellent work upon
the histology) ; Louping 111 and Braxy Committee's Report, pp. 21 et seq., 0.
Mohler, 1905, p. 14, 0.— 1906. Wheler, p. 411 ; Vassal, p. 285, also Martin,
p. 105 (experiments with trypanosome infection : negative) ; Braun, p. 364. —
1907. Donitz, p. 90 ; Pocock, p. 201 (cites Wheler, 1902).— 1908. Nuttall
(figures illustrating a lecture, reproduced here) ; Banks, p. 24, 0 ; Bonnet,
p. 254, 0; Galli- Valerio, p. 611, biology; Nordenskiold, p. 637, histology,
important ; Stockman (reprint, relation to redwater). — 1909. Samson, (a),
pp. 185-236 ; (6), pp. 486-499 (see under Iconography, Nordenskiold, (a),
p. 511 (spermatogenesis) ; (6), p. 30 (development) ; (c), p. 449 (anatomy
and histology, important)). Blanchard, p. 91. Ashworth, p. 133. — 1910. Braun
and Liihe, p. 178, 0 ; Samson, pp. 1 et seq., dissertation, anatomy and biology.

Male (PI. IV, Text-figs. 115, 139-142) : average L. 245, W. 1-33 mm.^
Body dark red-brown, narrow in front, broadly rounded behind, very

' Cited in bibliography by Salmon and Stiles, 1901, unimportant.

2 Note regarding measurements : The size of different specimens may vary, thus 6 d' s,
comprised in two lots collected iu England (N. 933, 934), measured in mm.—

Scutum

Length, including Body ^ ' n

capltulum width length width

f 2-6 V- 1-3 1-85 1-1

Lot 933 2-5 1-4 20 1-2

(2-3 1-35 1-75 1-1

('2-5 1-4 1-9 1-1

Lot 934 \ 2-4 1-3 1-8 I'O

( 2-2 1-2 1-6 0-95

Wheler (xii. 1899), who has made numerous measurements, states that the length
varies from 2-35 to 2*8 mm. Neumann (1899, p. 113) gives the size at 2-5 x 1-5 mm.

The size of the unfed ? is given at 2-85 x 3*25 mm., and that of the gorged s at

148

Genus Ixodes

convex when gorged. Marginal fold slight, generally of lighter colour
than the scutum. Scutum (average 1'8 x 1"1 mm.) glossy with very
numerous fine punctations ; cervical grooves very superficial ; lateral
grooves absent, or their commencement faintly indicated. Numerous
fairly long scattered white hairs. Emargi nation moderate, scapulae
blunt. Venter: genital orifice large, between coxae III; pregenital
plate nearly twice as long as broad ; median plate fairly broad with
rounded sides, aual plate with lateral borders divergent, adanal plates
narrowing posteriorly. Spiracles large, oval, with long axis directed

Fig. 139. I. ricinm <? ; venter and dorsum, x 30. Specimen from Coilessan, Scotland.
(N. 933. Nuttall, 1908, Fig. 7, G. H. F. N. del.)

10x6*4 mm., by Wheler. Neumann states that the unfed ? measures 4x3 mm., the
gorged ? from 10 x 6 to 11 x 7 mm. Two of the most replete specimens in our collection
measure 10 x 7'5 and 10-5 x 65 mm. respectively. Nym2)hs, unfed, measure 1*5 x 1-68 mm.,
when gorged they attain 3-3 mm. in length (Wheler, 1899, p. 49).

/. n'cliius

140

forward and macula anterior. Gapituluin oUO fx long ; base trapezoid,
broader in front, slightly convex behind, without cornua. Hypostome
with six or eight marginal teeth, increasing in strength and con-
nected by crenulations, two large basal teeth being directed ventrally.
Palps broad, article 2 as broad as long, impressed dorsally, article
3 slightly longer than 2. Legs: long and strong; coxa I with internal
spine overlapping coxa II, and slight external tooth. A single external
conical tooth on coxae II-IV, the postero-internal angles of coxae II
and III are sub-dentate. Tarsi long, tapering gradually.

Fig. 140. I. ricinus i : capitulum in dorsal and ventral aspects, x 70. Specimen from
Coilessan, Scotland. (N. 933. Original, G. H. F. N. del.)

Fig. 141. I. ricinus <? : tarsi 1 and 4 seen in profile, x 33. (N. 930, drawn from balsam-
mounted specimen. Original, G. H. F. N. del.)

Fig. 142. I. ricinus : (C) digit x 155, and (H) hypostome x 100 ; (C) digit x 220, and
(H) Hypostome x 75 of s and $ respectively. (Neumann, 1899, Figs. 2, 3,
illustrating the structures in I. ovatus. We find them identical in I. ricinus.)

150

Grenus Ixodes

Female (Pis. IV and VII, Text-figs. 142, 143, 144): Body oval,
with numerous short, white hairs; when replete may attain 11 x 7mm.
Scutum (1"8 X 1*15 or 1*4 x 1"05 mm.) dark brown, sub-pentagonal,
with rounded angles ; emargiuation slight ; scapulae rather pointed ;
cervical grooves superficial, barely reaching the posterior border ; lateral
grooves fairly distinct for about half the length of the scutum ;
numerous very fine punctations, especially on the posterior portion.

Fig. 143. I. ricinus ? (partly replete) : dorsum, part of venter, tarsus 4, hypostome
and spiracle. Specimen from Coilessan, Scotland, D. MacKenzie coll., iv. 1905.
(N. 934. Original, G. H.F.N, del.)

Fig. 144. I, ricinug ? : capitulum in dorsal and ventral aspects, x 70. Specimen from
Coilessan, Scotland. (N. 933. Original, G. H. F, N. del.)

/. riciutis

151

Venter: vulva between coxae IV; spiracles circular, genital grooves
only slightly divergent, anal grooves rounded in front, parallel behind
anus. Gapiiidum long (800 fi). Base pentagonal with parallel sides
and concave posterior border; no cornua ; slight auricular ridges on
its ventral surface. Porose areas piriform, the broader end internal,
situated near the posterior border, the interval equal to their lesser
diameter (see Fig. 143, exaggerated in Fig. 144). Palps long, article 2
nearly twice as long as broad, impressed dorsal ly, article 3 distinctly
shorter than 2, rounded anteriorly. Hypostome long, with sides nearly
parallel, dentition 3 j 3, about ten sharp teeth per file, the external
the stronger. Legs : slender ; coxae as in the </ ; tarsi long, tapering
gradually.

Nymph (PI. IV, Fig. 4, Text-figs. 14.5, 146) : with the charac-
teristics of the $ , except that the lateral angles and the grooves of the
scutum are more pronounced, and the postero-lateral angles of the basis
capituli more pointed. The internal spine on coxa I is short, but
distinct.

Fig. 145, I.ricinuso: dorsum and venter, x 30. (Nuttall, 1908, Fig. 5,
G. H.F.N, del.)

152

Genus Ixodes

Fig. 146. I. ricinus o : capitulum in dorsal and ventral aspects, x 70. (Original,

G. H.F.N, del.).

Larva (PI. IV, Fig. 5, Text-figs. 147, 148): with the essential
characters of the O.

See further under Notes on Biology (p. 294). Our description is
based upon the examination of a large material, collected in different
parts of the world.

Geographical Distribution.

Our collection contains specimens from the following places and
hosts [observations by other authors are added in brackets] :

EUROPE : Scotland : From Blackaddie, Sanquhar, v. 1908 (N.i
400, 401, 417), (/"s $s, from dog, cattle and sheep (R. Bramwell); from
Shelfhill Farm, Hawick, v. 1908 (N. 405), ^s, $s, from sheep
(J. Murray) ; Mr W. F. Cooper's collection also contains $ s from sheep
at Hawick, IV. 1906 (C. Grieve); from Auchenbrack, Tynron (N. 423),
$s, OS and larvae from sheep, V. 1908 (J. McMonnies) ; from Argyll-
shire (N. 1054), cT from sheep, v. 1910 (Captain Leschallus); from
Thornliill, Dumfriesshire (N, 415, 416), from cattle and lambs, v. 1908
(J. R. Wallace); from Coilessan (N. 933), ^s, $s, from sheep, IV. 1905
(D. MacKenzie) ; from Inverness-shire (N. 355), from wild cat, iv. 1907.
In Dr J. H. Ashworth's collection, Edinburgh, we found specimens from
dog, Obney Farm, Bankfort, viil. 1904; from deer, near Crawford,
X. 1908; from wild cat, no localit}^ given; and from host?, Perth.

1 Numbers following the letter N, refer to specimens as catalogued in our collection in
Cambridge.

/. ricinus

153

Mr A. E. Shipley has brought us o s and larvae for identification which
were found on grouse'^. [Mr W. Evans (i. 1907, p. 35) records this tick
on c&ivs, near Callander, mostly attached to udders and adjacent parts
of legs (sexes in copula, 21 September); os found on head of I'ed-deer

Fig. 147. I, ricinus larva : venter and dorsum, x 65. Specimen from Co. Cavan.
(N. 929. Nuttall, 1908, Fig. 6, G. H.F.N, del.).

Fig. 148. I. ricinus larva : capitulum in dorsal and ventral aspects, x 200. Specimen
from Co, Cavan. (N. 929. Original, G. H. F. N. del.).

^ Neumann, 1901, p. 282, states that ricinus larvae have been found on the martin, but
does not state the locality.

N. I. 11

154 Genus Ixodes

from Argyllshire (August) ; $ s found under rocks at Oban and Aberfoyle
(April); cT near Killin (Det. by Neumann)]. England and Wales:
From Northumberland (N. 1089) (/s, $s, os, on sheep, v. 1905
(J. Hedley); from Huntingdonshire (N, 1037) ?s, on cat, ill. 1906
(E. C. Burleigh); from Grantchester, Cambs. (N. 123) o, 1906 (N. D. F.
Pearce) ; from Longner Hall, Shrewsbury (N. 1036) $ , on young rat,
IV. 1906 (R. F. L. Burton) ; from Lyndhurst, Hants (N. 656), larva,
on Mustela erminea, vii. 1900 (G. Tate; ex N. C. Rothschild coll.);
from Tavistock, Devonshire (N. 963) jf, $s, on cattle affected with
piroplasmosis (redwater), i. 1910, and (N. 1041) iv. 1910 (G. H. Gibbings).
[E. G. Wheler, 1899, p. 6, records the species on sheep, at Gowanburn
Farm, N. Tyne Valley, $ s in March, adults and o s being more numerous
in April ; specimens only occasionally found on deer and dogs ; Kossel
and Weber, 1900, found this tick associated with redwater in cattle in
N. Wales.] Ireland : From Sligo (N. 1035) ? s on collie dog, v. 1905
{M. F. Nuttall); from Timmer (N. 630) os on Sciurus vulgaris, iv.
1901 (N. C. Rothschild); from Dallyhaise, Co. Cavan (N. 929), unfed
larvae, vil. 1905 (J. W. Steen) ; (N. 1038) ?s, on cattle affected
with piroplasmosis, v. 1905 (A. E. Mettam) ; (N. 1185) j/*, on dog,
Maam Cross, Co. Galway, 1. x. 1910 (G. H. F. Nuttall); Mr W. F.
Cooper's coll. contains (/s and ?s from Roscommon, viii. 1906. We
determined specimens belonging to the Dublin National Museum as
follows : from Glengariff, V. 1891 ; Clonbrock, ix. 1897 ; on cattle
Lochrea, vi. 1909; on cattle, Wexford; on cattle. Dingle, vi. 1909
Limerick, 1898 ; Cuppamore, Co. Limerick, ix. 1895 ; on horse, Stepaside
Co. Dublin, VI. 1909; on cattle, Cookstown, vi. 1909; Kenmure, 1898
on dog, Lugan, v. 1907 ; Achill, 1898 ; Glandore, 1895 ; Trillick, Co
Tyrone ; Crookbaveu, 1895 ; Roundstone, Ardagh, Castletown, vi
1893; Kenmore, vii. 1898; Lough Corrib, 1905. France: [Fairly
common in France, according to Neumann, 1899, p. 115, who records it
from the badger and genet, at Nimes (M. Mingaud)]. We have examined
numerous specimens in the collection of Dr H. Brumpt: </*, ?s and os,
taken from hedgehog and deer, Chantilly, ix. and xi. 1909 ; and from
the roe-deer, Halles (Paris), x. 1908. Belgium: (N. 631) ? from
Erinaceus europaeiis (E. A. Boulenger, ex N. C. Rothschild coll.).
Holland : [larvae found on Mus decumamis (Oudemans)]. Germany :
From Kittendorf and Schwandt, Mecklenburg-Schwerin, on various
hosts, as follows: (N. 57) c/", on man, ix. 1905; (N. 769, 770) (/"s, ?s,
on Cervus elaphus, viii. 1909; (N. 771) (/•, ? , o, on Capreolus capraea,

/. ricmus 155

VIII. 1909 ; also (N. 58) in ix. 1905, all collected by G. H. F. Nuttall;
and (N. 1182), in ix. 1910 (Count v. Schlieffen's gamekeeper). From
the same place (N. 1169), ?s, on dogs, vi-vii. 1910, and (N. 1170) on
fox, VI. 1910 (G. V. Oertzen's gamekeeper); and from the neighbouring
places: (N. 1183) %, from horse, at Vossfeld, vii. 1910, and (N. 1184)
?,0, from a cZo^r, at Rosenow, vii. 1910 (Count v. Schlieffen's game-
keeper). (N. 1124, 1181 and 1186 a) ?s, from Godesberg on Rhine,
on dogs, vi. vii. and ix. 1910 (Baron v. Rigal's gamekeeper; G. H. F.
Nuttall). [/. lacertae Koch was found on lizards in Germany. Kossel,
Schtitz, Weber and Miessner, 1903, pp. 39-50, observed all stages on
cattle, and demonstrated by experiment that ricinus conveys redwater
in Germany.] Heligoland: [Neumann, 1899, p. 116, states it was
found on Regidus ignicapillus and on Strix hrachyotus, and os were
taken from Tringa pugnax, according to G. Haller (Poppe coll.)]. Italy :
[Canestrini, 1890, p. 497, obtained specimens on cattle, dogs, goats and
roe-deer in the Trentino, and on goats and roe-deer in Veneto. He
states that Richiardi found them on RhinolopJtus ferrum equinum at
Pisa, and Canestrini found a young $ on Rh. euryale at Pisa. He
found young stages in Trentino, Istria, Veneto and Toscana on Lacerta
muralis and L. viridis. Canestrini and Fanzago, 1877, p. 112, found
many on hunting dogs, and on the goat, roe-deer and wild cat (p. 113,
/. rufus on Cervus dama and p. 114 /. lacertae on lizards)]. Spain:
[Neumann, 1901, p. 282, records its occurrence on lizards of different
species]. Galicia : [Neumann, 1899, p. 116, records its occurrence on
dog, sheep and Myoxus avellanariiis (Bureau of Animal Industry coll.,
Washington, D.C.)J. Albania and Rhodes: [Neumann, 1901, p. 282,
states that cfs, $s and os have been found on lizards: Lacerta agilis,
L. arenicola and L. vivipera)]. Russia: [The Hamburg Museum
contains specimens (^ and ?) collected by Dickmann in Amur, ac-
cording to Neumann, 1904, p. 452. He alsc records, 1899, p. 116, a o
from Caucasia in the M^gnin collection (now at the Laboratoire de
Parasitologic, Paris)]. Finland : [The prevalence of this species was
established by Kossel and Weber, 1900, in connection with their studies
on redwater in cattle. See also Kossel, 1902, p. 853; Kossel, Schiitz,
Weber and Miessner, 1903, p. 39 ; Nordenskiold, 1905, p. 485].

To the foregoing list of European hosts we add, on the authority of
Neumann, 1899, p. 115, the cormorant, polecat and mole, no locality
being given.

AFRICA: From Algeria: (N. 998) ^, $, on dog, Azazea, ill. 1910

11—2

156 Genus Ixodes

(G. H. F. Nuttall); (N. 1053 d) %, on sheep, Hammam-Meskoutine,
V. 1910 (P. Deshabert); (N, 1178) $s, from cattle, Algiers, summer
of 1910 (M. Serve); (in W. F. Cooper's coll.) %, on cattle, Maison
Carree, ii. 1908. [Neumann, 1901, p. 282, reports os and larvae, from
Lacerta ocellata and Tropidosaurus algirus. Tunisia (Paris Mus.) :
Neumann, 1899, p. 116, records specimens from Algeria, collected by
Simon (Paris Mus.).]

ASIA : Transcaucasia : (N. 799) ? s, from Surnabad, viii. 1903,
collected by Dr E. Dschunkovvsky, from whose collection we determined
specimens taken in the same place from fox, hare and sheep, x. 1903.
Arabia: [Occurrence on lizards; Neumann, 1901, p. 282]. Japan:
[Occurrence on horse at Akita, and hare at Saga. Neumann, 1899,
p.* 116, and 1904, p. 452].

AMERICA: From California (N. 272) on Felis concolor, Xli. 1894
(Gilroy), and (N. 291) on mouse, Palo Alto, iii. 1893 (both presented
by Prof. V. L. Kellogg) ; (in C. G. Hewitt's coll., Ottawa, determined by
us) %, on man, Vancouver, British Columbia, 1907 (G. W. Boggs).
[Osborn (1896, p. 262) states this species is common on ground squirrels
in the Mississippi Valley (determined by Marx). Banks (1908, p. 25)
states that the Marx collection in Washington contains specimens
found on sheep in Kansas, and on cattle in Texas. He thinks the
species may have been imported on the hosts, and adds : " Practically
all of the previous records of this species in this country apply to
/. scapularis or to /•. cookei" (species which we degrade to mere
varieties of /. ricinas). Neumann, 1899, p. 116, states that ricinus
occurs in many parts of the United States (Bureau of Animal Industry
coll., Washington, D.C., some perhaps are varieties of ricinus, q.v.) and
mentions New Berne, Baltimore, Pennsylvania, Carolina, Florida, Cali-
fornia, Kansas, Texas, the hosts being Lepus sylvaticus, Felis pardalis,
Bos taurus, opossum, grey fox, panther, and ivild cat.']

Ixodes ricinus var. scapularis (Say, 1821).

Fig. 149 (original).

Lit. and Synon.: Ixodes scapularis Say, 1821, pp. 59-82; 1859, pp. 19-22.
Ixodes affiants Neumann, 1899, pp. 120, 121, labelled /. communis by G. Marx
(Bureau of Animal Industry coll., Washington, D.C. ; also Neumann coll.),
fide Banks, 1908, p. 26.

/. ricinus var. scapularis

157

Ixodes scapularis Say, in Banks, 1904, p. 144 (occurrence in Florida); 1908,
pp. 25, 26, PI. IX, Figs. 1, 2 ; PI. II, Fig. 15 ( $ unfed, inaccurate ; sexes
in copula ( i^ attached too far po.steriorly), ^ venter and capitulum, details
fairly accurate (Fig. 15 very sketchy). Hooker, 1908 a, ob-served copulation.

Male: (according to Banks, 1908, p. 25) " L. 2 mm. Dark coloured
as in female; the palpi are very short, second and third joints no longer
than broad ; shield with subparallel sides, densely punctate and very
hairy ; legs and coxae as in the female ; the stigmal plate large and
elongate."

Female (Fig. 149) : Scutum rather more elongate-oval than in the
type, with grooves very faintly marked. Capitulum : base rather
narrower than in the type, with more distinct cornua ; porose areas
rounder and nearer together ; palps somewhat more pointed anteriorly.
Legs : tarsus 4 tapering rather more abruptly.

Apparently common in the United States (Florida, Texas, Virginia,
Iowa, N. Carolina, Maryland, Indiana), where it occurs on man, dog,
cattle, and various wild mammals, Neumann's affinis was found by
C. Curtice on Felis pardalis in Costa Rica.

Fig. 149. I. ricinus var. scajmlaris 9 : capitulum and scutum, capitulum in ventral
aspect, coxae, tarsus 4, spiracle, genital and anal grooves (less magnified). From
specimen (N. 718) from Miami, Florida, U.S.A. (ex N. Banks coll. Original,
G. H.F.N, del.).

158 Uenus Ixodes

We possess specimens (N. 626), j/*, $, from Gariacus virginianus,
Craven County, N. Carolina, xi. 1897 (ex N. C. Rothschild coll.), and
(N. 718), $, from Miami, Florida (ex N. Banks coll., presented by
the U. S. Dept. of Agriculture). The collection of C. G. Hewitt,
Ottawa, contains % specimens (determined by us), taken in Canada
from man, and found by M. Mahafty, vi. 1904, at Bracebridge, Ontario.

Neumann, in describing /. affinis, remarks on its similarity to
/. ricinus. Banks gives it specific rank, and points out certain
respects in which it differs from /. ricinus, in some of which we are
unable quite to agree with him. We have very carefully examined
four % kindly given to us by the U. S. Dept. of Agriculture,
Washington, D.C. and other specimens taken in Craven County,
N, Carolina (Rothschild coll.). The form is certainly very near /.
ricinus, and if found in Europe would be attributed to that species
with little hesitation considering the variation revealed by the review
of a very large number of typical /. ricinus. It seems to us impossible
to accord it higher rank than that of a variety of /. ricinus.

Ixodes ricinus var. ovatus (Neumann, 1899).

Lit. etc: Ixodes ovatus Neumann, 1899, p. 112, Figs. 2, .3. It was subsequently

found that the lot desci'ibed contained typical ricinus. The figs, of the ^

and $ hypostomes and digits as in ricinus.
Ixodes ovatus Neumann, 1904, p. 452, species maintained (description quoted

below).
I.vodes ovatus Neumann, in Donitz, 1905, p. 132. Donitz considers the species

probably but a variety of Hcimcs.

Neumann (1904, p. 452) states that ovatus differs from ricinus in
the following points :

Male : unknown.

Female : Scutum about as long as broad, cervical grooves very
shallow and long, lateral grooves indicated by a slightly prominent ridge,
parallel to the borders ; porose areas oval, slightly longer than broad,
converging anteriorly ; coxa I with a very short internal spine, II
unarmed, sharp on its posterior border, III and IV with a small
external tuberosity ; tarsi shorter than in /. ricinus.

Described from two $ found on a horse and a dog at Saga, Japan,
by Yamaguchi. The differences noted only appear to us to be varietal.

/. nif/rlcans 159

Ixodes ricinus var. californicus (Banks, 1904).

Lit. etc. ; Ixodes californicus Banks, 1904, \i. 309, PI. XLI, Fig. 57 (scutum and
coxae)*; 1908, p. 24, PI. II, Fig. 12 (9 capitulum and scutum).

We have not seen this tick, which appears, from its description, to
be specifically indistinguishable from /. ricinus. Banks states that
the capitalum of californicus is much narrower, and that the two
forms are separable by the different porose areas, but his description
of the porose areas of californicus precisely applies to our specimens
of ricinus. Moreover, the figures given by Banks do not appear to
us to agree very well with his description. It is undoubtedly very
near to ricinus, and it appears to be only a variety.

Male: (2 -mm. I.) similar to /. scapularis ^, but scutum with more nearly
parallel sides. Spiracle nearly circular ; palps very short ; coxa I with long sharp
posterior spine.

Female : Colo^ir -. Scutum yellowish-brown, legs darker brown, abdomen
yellowish-grey. Scutum 1*2 mm. I., but little longer than broad, somewhat
trapezoidal, lateral " carinae " not distinct, but traceable, many fine punctations.
Abdomen punctate, hairy. Capitulum not nearly as broad as in /. ricinus,
posterior angles acute, porose areas subtriangular, broader than long, separated by
about their length, the inner edge oblique ; palps moderate, article 2 slightly longer
than 3, the latter not twice as long as broad. Spiracle nearly circular, rather small,
finely granular. Legs : coxa I with long sharp basal spine, coxae I-IV with minute
tooth at apex behind ; legs rather slender, very hairy ventrally, tarsus 1 fully one
and one-half times the length of the metatarsus, tapering distally ; tarsus 4 but
little longer than metatarsus, tapering gradually.

Origin : California (Claremont, Santa Clara Co., Santa Cruz Mts., Redwood
Creek, Humboldt Co.). Hosts : grey fox and black-tail deer.

The above description is that given by Banks, rearranged, as far as
possible, in accordance with the method adopted in this work.

2. IXODES NIGRICANS Neumann, 1908.

Fig. 150 (original).

Lit. and Synon, : Ixodes obscurus Neumann, 1899, p. 121 (no figure).

Ixodes nigricans Neumann, vii, 1908, p. 75. Name given for the reason that

ohscinnis was preoccupied (/. ohscxirus Fabricius, 1805, p. 55, suppressed).
Male ; unknown.

1 The earlier description by Banks (1904) applies, apparently, to a ? and o, when con-
sidered in conjunction with the figure. He states that coxa I has a " black hump " at the
postero-external border, but no [sic) basal spine ; coxae II and III with prominent spine, and
coxa IV with small protuberance at the postero-external border. Found on Toxostoma
crissalis, at Claremont, California (Baker). This description does not agree with the later
one.

160

Grenus Ixodes

Female (Fig. 150) : Body short oval, 7x5 mm. Dark brown all over. Scutiim
oval, borders rounded, 1-3 x Tl mm., glossy, dark, with a few hairs ; cervical grooves
long, barely visible; many very fine equal punctations. Dorsum bearing many
punctiform prominences, some still surmounted by a hair. Venter bearing similar
granulations and hairs ; vulva facing the last intercoxal space ; anal grooves slightly
divergent ; spiracles rounded, small, reddish. Capihdum 1 mm. long, digit ^
hypostome thick, tip slightly tapering, 4 | 4 files of 10-11 teeth, diminishing from
outside inward, the external stout, pointed ; the internal blunt, distant from their
mates ; palps flat, the external border straight, the internal border convex, article 3
as long as 2. Legs of medium length, slender, bearing long hairs on the ventral
border ; tarsi long, tapering gradually ; pad almost equals the length of the claws.

Description (Neumann, 1889, p. 121) based on one gorged % from
Funchal, Madeira, collected by K. Kraepelin (Hamburg Museum).

Fig. 150. I. nigricans ? : capituhim and scutum ; parts of venter, spiracle and tarsus 4.
Drawn from the type. (Hamburg Miis. Original, G. H. F.N. del.)

Note: Through the courtesy of Professor Kraepelin we have been
able to examine and figure the type (1 ?). /. nigricans appai'ently
differs from /. ricinus only in (a) the scutum, which is oval (without
lateral angularity) and faintly grooved, and (h) the somewhat smaller
spiracle. A review of a series of undoubted ricinus from a single batch
reveals scutal variations which, in our opinion, render it quite possible

1 Digit 175 /i long; dorsal process longitudinally crescen tic ; external article with five
successive teeth, the last stout.

/. aciiminatus

161

for nigricans to fall within that species, but the opinion of so high an
authority as Professor Neumann leads us for the present to refrain from
referring it to /. ricinus. (From I. rasus, to which Neumann states
that it is allied, it is immediately distinguishable by the anal grooves.)

3. IXODES ACUMINATUS Neumann, 1901.
Figs. 1.51 and 152.
Lit. and Icon. : Neumann, 1901, pp. 287, 288, Figs. 7, 8 (here reproduced).

Male : unknown.

Female (Fig. 151), young : Body elongate oval, L. 2 mm., at least
twice as broad (1*2 mm.) toward the posterior third as it is in front,
yellowish red. Scutum longer than broad (TS x 0"9 mm.), lozenge-
shaped oval, antero-lateral borders subrectilinear, divergent, the portion
bounded by the posterior rounded border forming | of its length ;
emargination slight : surface glossy, without cervical grooves, the latter
fusing with the origins of the lateral rectilinear grooves, which are
parallel to the borders and end at their posterior curved portion ; punc-
tations numerous, very fine ; some scattered hairs. Dorsum covered by
long hairs, especially in the posterior portion ; deep marginal grooves

Fig. 151. Fig. 152.

Fig. 151. I. aciDuinatus ? : venter. (Neumann, 1901, Fig. 7.)

Fig. 152. I. acuminatum ? : H, hypostome, C, digit, 120 ix I. (Neumann, 1901, Fig. 8.)

162 Genus Ixodes

ending at the posterior curve of the border ; Venter bearing similar
hairs; vulva facing coxae IV ; sexual and anal grooves but slightly
divergent. Spiracles large, whitish, circular. Capitulum 0'8 mm. long;
dorsal base almost as broad as long ; lateral borders parallel, posterior
border rectilinear, with slight cornua ; porose areas circular, widely
separated, near to the posterior border ; ventral surface longer than
broad, auricula short, flat, rounded. (Digit, see Fig. 152 C.) Hypostome
(Fig. 152 H) lanceolate, very pointed, with 3 j 3 then 2 j 2 rows of sharp
teeth ; palps long, slender, narrow. Legs long, slender ; coxae broad,
flat, all bearing a short spur at the postero-external angle ; coxa I bears
besides a long slender spine internally. Tarsi long, slender, tapering
gradually ; pads as long as claws. Gorged female 4 mm. long, 2 mm.
broad, red-brown, the coxae situated close together in the anterior
fourth of the body.

Neumann's description based on 2 $ s from Mus agrarius collected
at Genoa by C. Parona (Neumann coll., Toulouse).

4. IXODES DENTATUS Marx, 1899.

Figs. 153, 154.

Lit. and Icon. : Neumann, 1899, p. 119, Figs. 4 and 5 (reproduced).
Ward (x. 1900, brief mention only on p. 204).

Banks, 1908, p. 28 ; PI. IV, Fig. 6 ( 9 capitulum, scutum, coxae), PI. IX, Fig. 3
( $ gorged, capitulum and scutum) ; inaccurate, since the two sets of figures
completely disagree.

Male : unknown.

Female: Body oval, L. 62 mm., W. 3'5 mm. [? replete], broad,
blackish brown. Scutum oval, scarcely longer (1 mm.) than broad, with
sides rounded, posterior angle broad ; cervical and lateral grooves well
defined, the latter straight, almost united behind to the cervical grooves ;
many large punctations, especially abundant behind. Dorsum shagreened,
with numerous punctations, each of which may bear a very short hair ;
grooves normal. Venter shagreened like the dorsum. Vulva facing
coxae IV ; sexual grooves very divergent. Anus toward the posterior
third ; anal grooves long, divergent. Spiracles small, rounded, in front
of one-half the body length. Capitulum (Fig. 153) with dorsal base
rectangular, slightly broader than long ; porose areas small, far apart ;
auricula forming a retrograde tooth. Chelicera ? Hypostome lanceo-
late, 4 j 4. Palps with article 2 slightly longer than 3. Legs short,
maroon-brown. Coxae (Fig. 154) close together, not occupying the

/. diversifossus

163

anterior quarter of the body length ; coxa I bears a long retrograde spur
at the internal angle ; all the coxae bear a spiniform process at the
postero-external angle. Tarsi medium, tapering gradually ; pads almost
equal to the claws in length.

The above description, from Neumann, is based on drawings by
Marx, and a $ . labelled by him Ixodes dentatus, found on the rabbit
(Smithsonian Institution, Washington, D.C.).

Fig. 153. Fig. 154.

Figs. 15.3, 154. I. dfniatus ? : capitulura in ventral aspect, coxae and trochanters I-IV
(Neumann, 1899, Figs. 4 and 5, G. Marx del.).

5. IXODES DIVERSIFOSSUS Neumann, 1899.

Lit. : Ixodes diversifossus Neumann, 1899, pp. 13(i, 137 (no figure).

Banks, 1908, p. 27, PI. Ill, Figs. 12, 14 ($ capitulum, scutum, spiracle, coxae I
and II, tarsus 1); not reproduced, as we do not trust their accuracy.

Male : unknown.

Female : Body oval, reddish yellow, o x 2*5 mm. [? partly gorged].
Scutum oval, 1*3 x Tl mm., with rounded borders, glossy, maroon-
brown ; cervical grooves long, superficial ; lateral grooves well marked,
straight, attaining the posterior border ; many punctations, large on
the lateral and posterior borders, very fine elsewhere. Dorsum
glabrous or nearly so, traversed by a marginal groove, very distant
from the lateral and posterior borders, where it leaves a margin
almost equal to one-quarter of the whole width. Veiiter : vulva
facing the last intercoxal space. Anal grooves parallel, of medium
length, reunited in an arc of a circle in front. Spiracles circular,
large, whitish. Capituluui with base broader than long, with pointed
cornua; porose areas rounded, small, far apart; auricula forming a

164

Genus Ixodes

retrograde spine. Chelicera ? Hypostome 2 | 2 or 3 ] 3, the marginal
teeth stout. Palps long ; article 1 with a short retrograde point ven-
trally, article 2 scarcely longer than 3. Legs long, brown ; spurs on
coxa I, the internal very long ; a simple pointed prominence at the
postero-external angle of coxae II-IV. Tarsi long, tapering gradually.
Description, b}^ Neumann, based on 2 $s (mutilated) collected by
Hassall from Procyonlotor, from New Mexico (Smithsonian Institution).

6. IXODES MINOR Neumann, 1902.

Figs. 155-159 (Figs. 156-159 original).
Lit. and Icon.: Neumann, 1902, pp. 109-111, Fig. 1 (reproduced).

Male (Figs. 155-158): Body : L. 145, W. 08 mm. at the posterior
third, maroon-brown. Scutum convex, glossy, marginal fold very narrow
with deep marginal groove ; cervical grooves narrow, superficial,
diverging and extending to about the middle of its length ; punctations
numerous, deep, uniform ; hairs whitish, lying flat, scattered. Venter :
sexual orifice broad, between coxae III ; median plate longer than wide,
pentagonal, with numerous large punctations ; anal plate smooth, narrow
anteriorly, broad posteriorly, with sides diverging considerably ; adanal
shields trapezoid, slightly broader anteriorly, with fine punctations; long

Fig. 155. Fig. 156.

Fig. 155. I. minor <? : venter. (Neumann, 1902, Fig. 1.)

Fig. 156. I. minor <? : venter (L. 1*73 mm.), from the type mounted in balsam.
(Neumann coll. Original, G. H. F. N. del.)

/. minor

165

hairs fairly abundant over the whole ventral surface. Spiracles large,
slightly oval. Capitulimi (Fig. 157) long (0*45 nun.), with base about as
broad as long, subrectangular, sides rounded, cornua very prominent.
Hypostome (Fig. 158) long, slightly lanceolate, on each border two rows
of 8 stout teeth, the internal connected ventrally by sinuous ridges.
Palps relatively long, articles 2 and 3 of the same length. Legs relatively
long ; coxa I bearing a short pointed spur at the postero-internal angle,
a blunter, shorter spur at the postero-external angle ; coxae II-IV
with a short spine toward the middle of the posterior border. Tarsi long,
tapering gradually.

Fig. 157.

Fig. 158.

Fig. 157. I. minor cj : base of capitulum, showing cornua. Drawn from the same
specimen as Fig. 156. (Scale = 0-25 mm. Original, G. H. F, N. del.)

Fig. 158. I. minor c? : hypostome, drawn from the same specimen as Fig. 156. (Scale =
0-2 mm. Original, G. H. F. N. del. )

Female (gorged, Fig. 159) : Body elongated oval, L, o, W. 2*5 mm.,
broadest at the posterior third, yellowish brown. Scutum oval, much
longer than broad (1'3' x 0"7 mm.), with curved contours, emargination
slight ; cervical grooves superficial, almost obliterated anteriorly, very
divergent posteriorly and extending to about the posterior third ; lateral
grooves faintly indicated by the marginal prominence ; punctations
numerous, uniform towards the posterior third and at the sides, finer and
rarer in the middle ; surface glossy, smooth, dark brown. Dorsum with
scattered hairs, numerous punctations; three posterior superficial grooves.
Venter with similar punctations and hairs; vulva small, opposite
coxae IV ; anal grooves diverging, ogival in front of the anus. Spiracles

1 Neumann gives the length of the ? scutum as 1-10 mm., which is probably a
misprint. He has kindly lent us the types, now mounted as microscopic preparations,
and, as far as their present condition permits, we have corroborated his description.

166

Genus Ixodes

small, circular, brown. Gapitulum long (0'8 mm.) ; dorsal base subtriau-
gular, with very prominent cornua ; porose areas large, circular, separated
by a space equal to their diameter ; auricula forming a prominent
horn. (Digits) Hypostome (absent, but probably sharply lanceolate as
in I. acuminatus). Palps long, slender, narrow ; the first article slightly
prominent externally on the ventral surface, the second slightly longer
than the third. Legs close together anteriorly, similar to those of j/*,
relatively slenderer.

Neumann's description is based on a </ and a $ found copulating
upon Hesperomys sp. ? (Muridae) in Guatemala by J. Rodriguez.

Fig. 159. I. minor ? : capitulum in dorsal and ventral aspects, spiracle, tarsus 4, scutum
and coxae. N.B. The dotted lines on the scutum indicate the lateral and cervical
grooves as described by Neumann ; they are invisible in the mounted specimen of the
type. (Neumann coll. Original, G. H. F. N. del.)

7. IXODES BOLIVIENSIS Neumann, 1904.

Fig. 160 (original).

Lit. : Neumann, 1904, pp. 457, 458 (no figure).

Male (Fig. 160); Body oval, broader posteriorly, L. 1*6, W. I'l mm.
Scutum convex, reddish brown, marginal fold narrow ; cervical grooves
almost obsolete ; lateral grooves absent ; two groups of deep punctations :
one group tracing a pseudo-scutum, the other group on a median circle
towards the posterior third, elsewhere almost obsolete ; a few rather long
sparse hairs. Venter reddish brown, with long hairs, especially lateral

1 Digit, slender, 100 /a long ; dorsal process crescentic, longitudinal and with retrograde
points, external article with five teeth, larger and more separated proceeding toward the
base (as in /. acuminatus).

/. holiviensis

167

to the genital grooves ; median shield with deep punctations, anal shield
a truncated oval, longer than wide, with sides curved and diverging;
adanal shield narrower posteriorly than anteriorly. Spiracles large, short
oval, whitish. Capitidum 525 fi long; the base slightly broader than
long and broader anteriorly on the dorsal surface ; its posterior border
rectilinear ; auriculae forming slight prominences. Hypostome long and
broad, with lateral teeth united by transverse serrated ridges. Palps
broad, flat, articles II and III of the same length. Legs long. Coxa I
with two parallel spurs, the internal very long, the external short ; two
flat spurs, short and separated, on coxae II and III ; a single external
conical spur on coxa IV. Tarsi long, tapering gradually.

Fig. 160. /. holiviensis j (type) : dorsum and venter, hypostome, spiracle and tarsus 4.

(Original, G. H. F. N. del.)

Female (Fig. 161) : Body oval, sides rounded, L. 2*6, W. 1'9 mm.
[partly fed], yellowish brown, the capitulum, scutum and legs maroon-
brown. Scutum oval, sides convex, longer than broad (I'o x r2 mm.);
cervical grooves almost obsolete ; lateral grooves represented by a
slightly visible depression ; fine punctations, mostly peripheral, a few
hairs mostly on the sides; scapulae sharp and prominent. Dorsum
with marginal grooves and some hairs. Venter with hairs scarcely more
abundant ; vulva between coxae IV ; anal grooves slightly diverging,
rounded in front. Spiracles large, circular, whitish. Capitulum 0"7 mm.
long; base broader than long, with parallel sides, with slight coruua;

168

Genus Ixodes

porose areas small, slightly broader than long, interval equal to their
long diameter; auriculae prominent, slightly diverging, uncate. Hypo-
stome (?). Palps long, broad, flat, the articles 2 and 3, almost equal.

Fig. 161. I. holiviemiii ? (type): capitulum and scutum; part of venter; spiracle.
(Original, G. H.F.N, del.)

Fig. 162. I. boliviensis o (type, unfed) : dorsum, part of venter, tarsus 4. Same
origin as cf and ? in Figs. 160, 161. (Original, G. H. F. N. del.)

/. fiiscipes

169

Legs : Coxa I with two parallel spurs, the internal very long, the ex-
ternal short ; a small external tuberosity on coxae II-IV. Tarsi ?

Nymph (Fig. 162) : Body oval, 1*1 mm. long. Capitulum very long
(0'63 mm.) ; with narrow palps, cultriform ; hypostome long, narrow,
lanceolate, dentition 2 | 2. Legs long ; coxa I with a stout internal spur,
coxae II-IV as in $ ; tarsi long, narrow, tapering gradually.

Described from a jf, a mutilated $ and a o, found on Icticyon
(Speothos) venaticus (Canidae), at Charuplaya, Bolivia, 27. vi. 1901
(N. C. Rothschild coll., now in Neumann coll.. No. 1251).

8. IXODES FUSCIPES Koch, 1844.

Figs. 163 and 164 (original).

Lit. etc. : Lvodes fuscipes Koch, C. L., 1844, p. 233 ; 1847, p. 106, PI. XXI,
Fig. 80, $.
Ixodes spinosus Neumann, 1899, pp. 146, 147 (no figure). Neumann, 1901,
pp. 288, 289, identified /. spinosus with /. fuscipes after examining the
type.

Male : unknown.

Female (Figs. 163, 164) : Body short oval, broadly rounded behind,
4*8 X 2*4 mm., yellowish or reddish yellow. Scutum rounded, at least as
broad (1*4 mm.) as long ; glossy, dark brown in front and on the sides.

Fig. 163. Fig. 164.

Fig. 163. 7. fuscipes ? : capitulum and scutum ; capitulum in ventral aspect, coxae,

spiracle and tarsus 4. Found on Dasyprocta aguti. (Neumann coll. 750. Original,

G.H. F.N. del.)
Fig. 164. J. fuscipes ? : capitulum (mutilated) and scutum, coxae and part of capitulum,

showing auricula. (Sketch from specimen in British Mus. Original, 0. W. del.)

N. I. 12

170 Genus Ixodes

brownish yellow between the cervical grooves ; the latter shallow,
attaining the posterior quarter ; lateral grooves indicated by straight,
nearly parallel raised margius which limit external declivities ; puncta-
tions deep, equal, numerous, absent on the cervical grooves, and anterior
border. Dorsum almost glabrous, showing a trace of a marginal groove.
Venter with more numerous hairs. Vulva between coxae IV ; sexual
grooves but slightly curved, diverging little. Anus rather near the
posterior border ; anal grooves parallel. Spiracles large, rounded,
whitish, situated about midway along the body-length. Capitulum
long (0*9 mm.) ; base almost as broad as long, pentagonal ; slight
cornua, the anterior border narrowed, porose areas small, far apart ;
auricula forming a horn-like protrusion. Hypostome narrow, pointed in
front, bearing 3 1 3 files of 13-14 teeth, decreasing inward in each
transverse row, the internal files much separated from each other,
especially behind.^ Palps long, narrow, article 1 prolonged into a ventral
retrograde spine ; article 2 longer than 3. Legs long. Coxae bear
some long hairs ; a moderate protuberance at the postero-external
angle ; a long spine at the postero-internal angle of coxae I. Tarsi
long on pairs I and IV, of ordinary size on pairs II and III, with
almost parallel borders ; the dorsal border abruptly tapering near the
extremity ; pads large, as long as the claws.

Nymph: Body short oval, L. 1'8, W. 1*5 mm., whitish. Scutum of
the same form as in $ ; cervical and lateral grooves clearly defined.
Capitulum 0*7 mm. long, similar to that of ? ; base broader in front
than behind ; no ventral spine on the first palpal article. Legs similar ;
the spine on coxa I shorter.

Larva : short oval, L. 1 mm. Scutum broader than long. Capitulum
similar to that of o, hypostome less pointed ; spine on coxa I shorter.

Neumann described this species as Ixodes spinosus from 4 $s, 7 os
and 3 larvae collected by Goeldi in Brazil from Dasyprocta aguti.
Subsequent examination of Koch's type oi I.fuscipes (1 % from Brazil)
convinced him that the two species were identical. He has kindly lent
us a $ from Goeldi's collection, and the above description has been
altered in a few unimportant respects from that of Neumann of
the $ of /. spinosus as the result of our study of this specimen.
(Fig. 163). We have also examined an older $ in the British Museum,
attributed by Neumann, probably correctly, to the same species, but it
has a distinctly longer scutum (Fig. 164). This specimen was found on
Felis pardalis, at Panama (Watson coll.).

1 The dentition of the hypostome shown in Fig. 163 differs from that here described ;
the structure is doubtless variable.

/. smdptus and spinicoxalis 171

9. IXODES SCULPTUS Neumann, 1904.

Lit.: Neumann, 1904, p. 462 (no tigure).

Banks, 1908, p. 30, PI. IV, Figs. 7 and 9 ( 9 capitulum, scutum, coxae) ; not
reproduced.

Male : unknown.

Female : Body oval, narrower anteriorly, brownish, L. 2, W. 1'3 mm.
[? unfed]. Scutum as long as broad (1 mm.), subcircular ; cervical grooves
superficial throughout almost their whole length, deep at their anterior
origin ; whence the lateral grooves start, these latter deep, limited
externally by a prominent ridge, which extends almost to the posterior
border; numerous punctatious, uniform, medium. Dorsum shows a
deep complete marginal groove and numerous hairs. Venter with
similar hairs ; vulva narrow, between coxae III ; genital grooves
diverging, anal grooves ogival anteriorly; with sides almost parallel;
spiracles brownish, circular. Capitulum medium (600 /a), with mitre-
shaped base, with prominent cornua ; porose areas large, deep, oval,
scarcely longer than wide, close together. Hypostome narrow, lanceolate,
dentition 2 j 2. Palps with second article almost twice as long as the
third. Legs stout and strong. Coxa I with internal spur, long, stout,
covering a part of coxa II ; a short spur at the postero-external angle
of all the coxae ; tarsi short, broad, without dorsal protuberance, but
taper ng abruptly.

Description (from Neumann) from a specimen found with a %
I. ricinus (L.), from the Santa Cruz Mountains, California (G. Marx
coll., Smithsonian Inst, Washington, D.C.). Banks, 1908, p. 31, states
that he has seen % specimens, differing but slightly from the type,
found on the rock squirrel at Del Rio, Texas (Bishopp).

10. IXODES SPINICOXALIS Neumann, 1899.

Figs. 165 (original) and 166 (Nn.).
Lit. and Icon. : Neumann, 1899, pp. 123, 124, Fig. 6 (reproduced).

Male : unknown.

Female (Fig. 165): Body oval, capitulum, scutum and legs
maroon-brown. Scutum oval, with rounded borders, 1*7 x 1"5 mm.,
glu.ssy, dark on the sides, lighter in the median field ; cervical grooves
superficial, almost attaining the posterior border; punctations deep,
subequal, distant, more crowded on the borders, where some whitish
hairs may occur. Dorsum : grooves but slightly pronounced ; whitish
scattered hairs, mostly on the sides. Venter: many hairs; vulva on

12—2

172

Grenus Ixodes

a line with the posterior border of coxae IV. Sexual grooves uniting
in front of the vulva, straiofht and divergent. Anus not far from the
posterior border ; anal grooves almost parallel. Spiracles large, circular,
whitish, equidistant from the vulva and anus. Capitulum long (1 mm.),
with dark brown base, the appendages brown-red. Base about as
broad as long, subtriangular ; porose areas large ; auricula forming
a flattened retrograde tuberosity. (Digits) Hypostome (Fig. 166)
narrow, lanceolate, long, 4 | 4 then 3 | 3 files of 9-10 teeth, the external
very stout, the median less so; the internal narrow, long and blunt.
Palps long, broad, flat ; article 3 almost as long as 2. Legs large, stout,
the two last articles of pair 4 extending beyond the posterior border of
the body, brown, except the distal extremity of the intermediary articles,
which is white ; short hairs on all the articles. Coxae large, contiguous ;
coxa I bears a long spine at the postero-internal angle; a very short
spiniform prominence at each of the posterior angles of all the coxae.
The movable articles are slightly larger distally. Tarsus 1 long, almost
as long as articles 4 and 5 combined, tapering abruptly at the distal
pseudo-articulation ; tarsus 4 shorter, tapering abruptly ; pad two-
thirds the length of the claws.

Description based on 4 ?s found on Mustela flavigula in Sumatra by
H. 0. Forbes (British Museum).

Fig. 165. Fig- 166.

Fig. 165. I. spinicoxalis ? : capitulum (mutilated) and scutum, coxae and part of basis

capituli. (Sketch from the type in the British Mus. Original, C. W. del. )
Fig. 166. I. spinicoxalis ? : hypostome x 115. (Neumann, 1899, Fig. 6.)

1 Digit 160 fi long; dorsal process elongate, parallel to digit, crescentic, with two
retrograde points; external article large, longer than half the digit, with six or seven
teeth, the 3-4 anterior ones small, equal, the three others increasing in size, the basal
very stout.

/. marxi

173

11. IXODES MARXI Banks, 1908.

Fig. 1G7 (original).

Lit. etc. : Banks, 1908, p. 32, PI. IX, Fig. 8 ( 9 dorsum, G. Marx del.), PI. Ill,
Fig. 10 (9 capituluin, scutum and coxae; Banks del).

Male : unknown.

Female (Fig. 167): Scutum (1-2 x 0-9 mm.), longer than broad,
broadest rather in front of the middle (but not so anteriorly as in
7. canisuga); cervical grooves broad and very shallow, only visible for
about half the scutal length; no lateral grooves; scapulae rather
rugose ; scapular angles sharp ; emargination moderate ; a few fine,
shallow punctations on the median area of the scutum. Venter: vulva
between coxae III ; anal grooves very ogival in front, the sides long,
straight, slightly diverging, much nearer together than in /. canisuga ;
spiracle very small, circular. Gapitulum: base subtriangular, concave
posteriorly, with slight cornua, porose areas ill-defined, subcircular, far
apart, and far from lateral border ; slight laterally projecting auriculae,
ventral surface of basis capituli flat ; palps of medium length, rather
narrow, article 2 rather longer than article 3, article 3 somewhat pointed
anteriorly ; hypostome short, narrow, lanceolate, 2 1 2, the outer teeth
much the stronger, about 6 teeth per file. Legs like those of /. canisuga,
but all the articles shorter; coxa I with slight internal spur, coxae
II-IV unarmed ; tarsi tapering abruptly, almost humped.

CC3^

Fig. 167. /. marxi i : capitulum and scutum, ventral aspect of capitulum with coxae,
tarsus 4, spiracle, genital and anal grooves. From specimen kindly loaned by Dr
L. 0. Howard, Government Entomologist, Washington, D.C., U.S.A. (Original,
G. H.F.N, del.)

174

Genus Ixodes

Described from a $ ^ found ou a squirrel, District of Columbia,
U.S.A., kindly loaned by the U.S. Department of Agriculture, being
one of the specimens doubtfully attributed by Neumann to /. hexagonus
var. inchoatus (=/. canisuga). It is somewhat allied to I. canisuga,
but can easily be distinguished from it by the diflferent shape of the
basis capituli, the more posterior position of the lateral angles of the
scutum, the very small spiracles, and other minor characteristics.

This species occurs in Canada and the United States, having been
found, according to Banks, on the red squirrel in Guelph, Ontario,
Canada ; Salineville and Waseon, Ohio ; Portland, Michigan ; Ithaca,
New York ; District of Columbia ; and on the fox in Denver, Colorado.

12. IXODES PRATTI Banks, 1908.

Fig. 168 (original).
Lit. and Icon. : Banks, N., 1908, pp. 27, 28, PI. IV, Figs. 1, 3-5.

Male: (according to Banks, 1908, p. 28) "Z. 2 mm. Body very
slender, more than twice as long as broad, dark brown, and very hairy ;
a deeply impressed lateral and posterior groove ; surface densely
punctate, Capitulum not broader behind than long in middle, sides
parallel, posterior angles not prolonged ; palpi short, but longer than
width of capitulum, the third joint a little longer than broad, legs
rather short, coxae I with a very long, slender spine behind, II with

Fig. 168, I. pratti ? : capitulum and scutum ; capitulum in ventral aspect, tarsus 4
(claws missing in specimen), spiracle, coxae. Specimen from Death Valley, California,
determined by N. Banks. Eeceived from U.S. Dept. of Agricult. (N. 716. Original,
G. H.F.N, del,)

^ The tube containing the specimen also contained three labels bearing the inscriptions :
(1) No. 47-3, Ixodes americanus Marx, Squirrel, D.C. (Marx, coll.) ; (2) Ixodes hexagonus
var. inchoata Neumann (Nn. det.) ; (3) Ixodes viarxi Banks.

/. pratti and 7'ubidus 175

two humps behind, and III with one hump; venter punctate ; stigmal
plate twice as long as broad, finely granulate."

Female (Fig. 168): Scutum 1-2 x 11 mm., but slightly longer
than broad, somewhat narrowly rounded behind, cervical grooves shallow,
extending rather more than half the length of the scutum, lateral
grooves bounded externally by a prominent dark ridge with a few fine
punctations ; the area be ween this ridge and the cervical groove is
strongly pitted with rather large, more or less confluent punctations ;
fine, discrete punctations over the rest of the scutum. Capitulum :
base broader than long, with slightly convex sides and fairly strong
cornua slightly curving inwards ; porose areas piriform, the broader end
internal, interval moderate, no auriculae ^ Palps rather short and
broad, article 1 strongly prominent ventrally. Hypostorae strong, well
covered with 2 j 2 files of strong teeth, with no median unarmed area.
Venter : vulva facing the second intercoxal space ; anal grooves rounded
in front, diverging behind ; spiracle rather small, nearly circular. Legs :
coxa I with rather strong internal spur and a slight external spur ; a
faint indication of an external spur on coxae II-IV. Tarsus 4 tapering
abruptly.

We have not seen the ^ of this species, and therefore quote the
description given by Banks. The description of the % is from a
specimen (N. 716) presented to us by the U.S. Department of Agri-
culture, Washington, D.C., and taken in Death Valley, California
(? host). Banks records it from Kerrville, Texas ; Sherwood, Texas
(on a prairie dog) ; and from Thomomys (a rodent) in Death Valley,
California.

13. IXODES RUBIDUS Neumann, 1901.

Figs. 169 and 170 (original).
Lit. : Neumann, 1901, p. 282 (no figure).
Male : unknown.

Female (Fig, 169; unfed): Scutum 1*4 x 1*3 mm., ovate, broadest
at anterior third, glossy, hairless, convex, with lateral borders folded
down, closely punctate and irregularly pitted ; the punctations larger
and more confluent in front, discrete and uniform behind. Cervical
grooves long, reaching the posterior border, gradually diverging ; no
lateral grooves. Scapulae prominent. Dorsum with deep marginal
grooves, ending where the genital grooves turn over the posterior border
of the body ; well-marked furrows in continuation of cervical grooves.
1 The presence of large auriculae is indicated in Banks' figure, which is very inaccurate.
We have not included the i in our key, Banks' description being insufficient.

17(3

Genus Ixodes

Venter : vulva between coxae III ; genital grooves parallel at first,
curving outwards behind coxae IV, then nearly parallel but slightly
divergent to posterior border. Anal grooves ogival in front of anus.
Spiracle large, nearly circular, with eccentric macula. Gapitulum:
(0*8 mm. I.), base pentagonal, about as broad as long ; cornua indistinct,
posterior border concave ; porose areas medium, slightly broader than
long, near posterior border, interval almost equal to their length.
Palps medium, article 2 half as long again as article 3, their convex
inner border somewhat beaded. Hypostome 2 1 2, about 6 teeth per
file. Legs long, coxa I with long slender internal spine and short
external spur ; a small external spur on coxae II-IV. Tarsi strongly
humped.

Fig. 169. Fig. 170.

Fig. 169. I. rubidus ? (type, unfed) : dorsum, part of venter, spiracle and tarsus 4.

(Neumann, coll. 713. Original, G. H. F. N. del.)
Fig. 170. I. rubidus o (type, partly fed) : capitulum and scutum ; capitulum in ventral

aspect with coxae, spiracle and tarsus 4. Same origin as ? previously figured.

(Original, G. H.F.N, del.)

Nymph (Fig. 170): found in company with this ?, agrees with it
fairly well except that the scutum (0"6 x 0*5 mm.) differs in shape ; the
internal spur on coxa I is nearly absent and the basis capituli bears
auriculae.

Our description is based upon the examination of one ? and of
one of three o s collected by A. Duges off Bassaris astuta at Guanajuato,
Mexico (Types, No. 713 in Neumann coll.).

/. hexagomis 177

14. IXODES HEXAGONUS Leach, 1815.

Figs. 171-177 (original).

Synonymy: l. Ixodes hexagomis Leach, 1815, xi, p. 397, on hedgehog.

2. Ixodes autumnalis Leach, 1815, xi, p. 398, on pointer dogi.

3. Ixodes erinacei Andouin, 18.32, xxv, p. 415 ($)*'^.

4. Ixodes reduoius Audouin, 1832, xxv, p. 422; ne<; Linn^ {$y.

5. Ixodes cremdatus Koch, 1835-44, Heft 39 1.

6. Ixodes vuljjis Pagenstecher, 1861, ii, p. 40 1'^.

7. Ixodes erinaceiis Murray, 1877, p. 190 ^

8. Ixodes ridnus Megnin, 1880, p. 129 ^

9. Ixodes sexpunctatus Koch, 1847, p. 22 (nymphs found in Germany), types

examined by Neumann, 1901, p. 283.

10. Ixodes cremdatus Koch, in Berlese, 1889, fasc. 55, PI. IV, $ dorsum and

venter.

11. Euixodes hexagonus (Leach) in Bonnet, 1908, p. 255. (/. canisuga Johnston,

1849 ; /. cookei Packard, 1869 ; /. crueiarius Fitch, 1872, are listed by
Blanchard, 1909, p. 86, as synonyms ; see our List of condemned species.
We recognize canisttga.)
Iconography : Audouin, 1832, xxv, PI. XIV, Figs. 2, 4.— Koch, 1835-44, Heft xxxix,
Figs. 5, 6 ( 5 /. sexpunctatus) ; Figs. 8, 9 ( (^ $ /. crenulatits). — Pagenstecher,
1861, II, PI. I, Figs. 12, 13.— Berlese, 1889, fasc. 55, No. 4 (/. cremdatus
V. Synon.) ; 1892, fasc. 61, No. 10 ; ? dorsum, venter, capitulum and scutum,
spiracle and tarsus 4.— Canestrini, 1890, iv, PL XL 1 1 1, Fig. 2 : ? tarsus 4
and foot 4, poor.— Railliet, 1895, Fig. 484: ? (gorged), dorsum and venter;
Fig. 478, capitulum (after Delafond) poor (reproduced by Ward, 1900, Fig. 256).
—Neumann, 189.3, in Railliet, 2nd edit., fasc. 1, Fig. 485 (tarsus 4), same
figure reproduced by Ward, 1900, Fig. 257, and Salmon and Stiles, 1901,
Fig. 229. — Neumann, 1899, Fig. 10 {(^ venter) not reproduced, but replaced
by our original figure. — Salmon and Stiles, 1901, Fig. 229, reproduce figures by
Neumann, 1893 {v. supra) and 1899 ; also by Berlese (y. supra). — Wheler, 1899,
various stages figured, reproductions of photographs of balsam-mounted speci-
mens and details thereof ; much retouched, and of little value ( ? , o and
larva).— Mohler, 1905, PI. II, Figs. 4, 4o, 4 6 (sketchy, coloured).— Wheler,
1906, PI. VIII, Fig. 22, $, rough sketch after Neumann, 1899, Fig. 10;
PI. VIII, Fig. 20: ? dorsum, good photograph of opaque object (reproduced). —
Banks, 1908, PI. Ill, Figs. 11, 13 ( $ coxae, capitulum and scutum), very
inaccurate.— Bonnet, 1908, Figs. 14, 17-20 original, but bad.— Blanchard, 1909,
Figs. 91-93 (from Neumann and Bonnet).
Literature : Some of the authors cited below doubtless refer to /. canisuga, but
their descriptions are too vague to be certain. In most cases /. hexagomis is
undoubtedly described, and this is rendered certain by reference to the Icono-

^ Also included in Synonymy by Neumann, 1899, p. 129.
2 Also included in Synonymy by Canestrini, 1890, p. 500.

178 Genus Ixodes

graphy above given. The small numbers iu brackets (2-11) accompanying the
following citations refer to the names under which the authors described the
species in the order given in the Synonymy ; where no number is given the name
I. hexagonus was used.

1815. Leach, p. 397 ; p. 398 (2).— 1832. Audouin, p. 415 (3) ; p. 422 (4).—
1835-44. Koch, Heft 39. (5, 9).— 1844. Koch, p. 2.34; p. 233 (5, 9);
Walckenaer and Gervais, p. 240 ; p. 241 (2) ; p. 242 (3).— 1847. Koch, p. 23 ;
p. 22 (5, 9).— -1859. Gervais and van Beneden, p. 461 (2), — 1861. Pagenstecher,
p. 40 (6).— 1877. Canestrini and Fanzago, p. 184 (5) ; Conil, p. 28 (6) ; Murray,
p. 190 (3).— 1880. Megnin, p. 129 (8).— 1882. Haller, p. 310 (3).— 1889. Berlese,
fasc. 55 ; Wheler.— 1890. Canestrini, p. 481 ; p. 500 (9).— 1891. Batelli, p. 78;
p. 81 (physiology of digestion) p. 98 (3). — 1892. Berlese, fasc. Lxi. n. 10, descrip-
tion.—1893. Railliet, p. 709.— 1899. Neumann, {a) p. 129; (6) p. 464.-1900.
Ward, {a) p. 204 ; {b) p. 436 (but a few lines); Dubreuilh (occurrence on man). —
1901. Salmon and Stiles, p. 467.— 1904. Neveu-Lemaire, p. 152.— 1905. Mohler,
p. 15 (brief inaccurate description). — 1906. Wheler, p. 412. — 1907. Hunter and
Hooker, p. 55. — 1908. Banks, p. 31; Bonnet, 255 (11); Levendain (cited by
Blanchard).— 1909. Blanchard, p. 86.

Male^ (Fig. 171): L. 3-5 to 4 mm. (capitulum incl.), W. 2 to
2'5 mm. (Nn.). Body oval, nearly as broad in front as behind. Sctitum
long oval, marginal fold moderate. Faint, strongly divergent cervical
grooves. No lateral grooves. Three longitudinal posterior furrows (or,
at least, areas devoid of punctations). A pseudo-scutum faintly indicated.
Very numerous punctations. Venter finely punctate. Pregenital plate

Fig. 171. I. hexagonus i : dorsum, venter, spiracle and tarsus 4. Specimen collected
in France by E. Simon, 1896. (Neumann coll. 719. Original, G. H. F. N. del.)

1 The male is extremely rare : although we possess hundreds of females we have only
recently come into the possession of a male through the courtesy of Professor Neumann.
The specimen is from the same lot as that shown in Fig. 171, and was taken in the Foret
de la Londe, Seine Inf^rieure. We have recently (7. iii. 1911) captured one male in a
hedgehog's nest near Cambridge. There are no males in the collections of the Museums
in London, Paris and Berlin.

/. hexagonus

179

almost a regular hexagou, preceded by two small plaques behind
coxae I ; median plate very broad ; aual plate ogival ; adanal plates
with nearly parallel sides, but rather broader in front. Spiracle rather
large, rounded. Capitulum comparatively very small. Base pentagonal,
narrowing posteriorly, slightly broader than long. Palps medium,
articles 2 and 3 nearly equal in length. Hypostome of the same type
as that of canisuga (Fig. 204), but not emarginate, it is bluntly rounded
in front and the crenulations are more distinct and nearly meet in the
middle line. Legs : coxa I with a strong, sharp spine on its internal
border, and a very slight external spur; coxae II-IV with slight
external and internal spur. Tarsi humped; pad small.

Female (Figs, 172-175) : when gorged may attain 8x5 mm.
Scutum (1'2 X l"^, or 1'5 x 1'4 mm.) a regular hexagon, except that the
posterior border is convex, broadest in the middle, scapulae sharp and
prominent ; cervical grooves faint, wavy, reaching posterior border ;
lateral grooves indicated by a slight ridge on the antero-lateral borders ;
numerous moderate punctations, often confluent in the anterior region;
smaller and discrete in the posterior region ; often a fairly distinct
subcircular depression on either side of the middle of the scutum.
Dorsum : punctations scarcely visible ; numerous short, white hairs.
Venter : vulva between coxae III ; genital grooves nearly straight,
slightly diverging ; anal grooves ogival, with sides parallel. Spiracle
large, subcircular. Capitulum of medium length ; base rectangular,
nearly twice as broad as long; porose areas pear-shaped, transverse, the
broader end internal, or sometimes oval, interval equal to or less than
their smaller diameter ; postero-lateral angles slightly salient (in some

Fig. 172. I. hexagonus ? : dorsum, venter, spiracle and tarsus 4. Specimen off
Mustella vulgaris, Tring ; F. J. Cox coll., 1908. (N. 608. Original, G. H.F.N, del.)

180

Genus Ixodes

specimens amounting to slight cornua). Palps medium, article 2 slightly
longer than 3 ; article 1 prominent ventrally. Hypostome large, with
strong, equal teeth, 3 1 3 to 2 | 2. Legs as in the </, except that the
internal spur on coxa I is shorter and directed somewhat more
outwardly.

Fig. 173. I. hexagonus ? : capitulum and scutum, ventral aspect of capitulum with coxae,
tarsus 4, spiracle and anal groove. Specimen from Myopotamm coijptt, S. America.
(N. 826. Original, F. M. H. del.)

Fig. 174. I. hexagonus ? : capitulum in dorsal, ventral and lateral aspects. (Original,

G. H. F. N. del.)

Fig. 175. I. hexagonus ? : tarsi 1 and 4, highly magnified, from mounted specimen.
(British origin. Original, G. H. F. N. del.)

/. hexagonus

181

Nymph (Fig. 176): bears a close resemblance to the ?, but the
inner spur on coxa I is less marked, and the hypostome bears 2 j 2 files
of 7-8 teeth.

Larva (Fig. 177): resembles the o, but the coxae are unarmed,
coxae I and II show a slight tuberosity at the postero-internal angle ;
tarsus 4 is less humped. Hypostome 2 2, with fewer teeth per file.

Fig. 176. I. hexagonus o : capitulum and scutum, coxae and capitulum (ventral aspect),
tarsus 4, spiracle and anal groove. Specimen found with the ? figured above.
(Original, G. H.F.N, del.)

<-\^:^

Fig. 177. I. hexagonus larva : dorsum, venter, tarsus 3 ; hypostome highly magnified.
Drawn from specimen (N. 82) raised in the laboratory, Cambridge, from ? taken in
Northumberland, 1904. (Original, G. H. F. N. del.)

182 Grenus Ixodes

Geographical Distribution.

Owing to the possible confusion, especially of immature stages, with
other closely allied species, it appears inadvisable to cite the hosts and
localities given by most authors. According to Neumann, 1899, p. 131,
this tick is widely distributed in Europe, and occurs on many species
of animals (mammals and birds).

Our collection contains $ s, o s and larvae from the following places
and hosts [observations b}^ other authors are added in square brackets] :

EUROPE : Scotland : Mr W. Evans' collection in Edinburgh con-
tains OS from the polecat, Ross-shire, xii. 1907, which we determined.
[Mr Evans (i. 1907, p. 35) has already reported finding this tick (?) on
this host in January, whilst larvae were found -on Erinaceus ewopaeus,
both in Ross-shire (Neumann det.).] England: (N. 404) $s, os, from
Mustela putorius, Holme, Peterborough, III. 1908 (W. Farren). (N. 60,
446, 599, 918, 1193) from Erinaceus europaeus : $ s, Cambridge, iv. 1897
(N. C. Rothschild); os, Cambridge, vii. 1904 (G. H. F. Nuttall); ?s,
OS, Madingley, near Cambridge, vii. 1908 (W. A. Harding); $,
Grantchester, near Cambridge, viii. 1906 (N. D. F. Pearce); os,
Longnor Hall, Leebotwood, Salop, ix. 1910 (R. F. L. Burton).
(N. 1160) Larvae from young Canis vulpes, Boxworth, Cambs., ill. 1900
(N. C. Rothschild). (N. 262) % , fiom Canis familiaris, Cambridge,
V. 1907 (G. S. Haynes). (N. 399) ?s, os, from Mustela erminea,
Histon, Cambs., v. 1908 (G. H. F. Nuttall). (N. 1067) ?, from Canis
vulpes, Boxworth, Hants., i. 1898 (N. C. Rothschild). (N. 600) from
Mustela putorius, Boxworth, iv. 1900 (N. C. Rothschild). In W. F.
Cooper's coll., $, from ferret, Canterbury, Kent, iv. 1908 (Spanton).
From Mustela erminea in the following places: (N. 611) os and larvae,
Lyndhurst, Hampshire, III. 1901 (G. Tate) ; (N. 603) Wigginton, Herts.,
V. 1903 (Barrett); (N. 607, 1159) os, Tring, Herts., iv. and IX. 1902;
(N. 602, 610), ?s, OS and larvae, Tring, Herts., v. 1902, and v. 1903;
all received from Hon. N. C. Rothschild. (N. 7) ?, gorged, removed
from a tvoman, Caxton, Cambs., VI. 1905 (Dr Giles, presented by
Dr Wherry); (N. 604) Mustela vulgaris, Tring, Herts., III. 1902
(N. C. Rothschild) ; (N. 668) $ s, o s, from the same host and place,
1908 (F. J. Cox); (N. 128) from Shrewsbury, IX. 1906 (R. F. L.
Burton); (N. 1194) ?, os, Claverdon Leys, Warwick, ix. 1910
(E. G. Wheler); (N. 456, 598, 601) ?s, from Lutra vulgaris; no
particulars (N. C. Rothschild and W. A. Harding). Wales : speci-
mens presented by the Hon. N. C. Rothschild from Mustela puto7'ius,

/. hexagonus var. cookei 183

Aberystwyth: (N. 746) larvae, viii. 1900; (N. 606) ?s, os, larvae,
XI. 1902; (N. G09) os, xii. 1902. Ireland: (N. 612) ?s, os, from
Mustela e^-niinea, Inismore, iii. 1902 (N. C. Rothschild). Belgium :
(N. 597) from Erinaceus europaeiis (E. A. Boulenger). Germany :
(N. 1168) $s, OS, from ferrets, Kittendorf, Mecklenburg-Schwerin,
VI. 1910 (G. V. Oertzen's gamekeeper) ; (N. 608) from Erinaceus
europaeiis, Dernburg, ix. 1898 (N. C. Rothschild); (N. 1186b) $,
from dof/, Godesberg, vii. 1910 (Baron v. Rigal's gamekeeper). France :
We have examined 2 ^/'s and a ^ kindly sent to us by Prof. Neumann ;
the specimens were collected by Simon, in the Foret de la Londe,
Seine Inferieure. We possess a ^Z* of this lot. [Italy : Canestrini, 1890,
p. 500, states that it occurs by preference on dogs ; he has examined
$s from dog, fox and hedgehog in Trentino, Veneto, Liguria and
Toscana. Canestrini, p. 502, records " creniilatus " from Borgotaro and
Trentino, occurring on Meles taxus.]

AFRICA: In W. F. Cooper's coll., ?, from wild boar, Algeria, 1908
(S. Williamson).

AMERICA : (N. 284) $ , from Sciurus sp., La Honda, California, ii.
1895 (ex V. L. Kellogg coll.). [Banks, 1908, p. 31, records this species
( ? s) from sheep, Kansas (Neumann det.) ; from rabbit, Baltimore, Md.
(Neumann det., ex Hassall coll.). He does not refer to his earlier
statement (1904, p. 331) that it occurs in California (Santa Clara
County, Palo Alto and Mt Shasta).] We have 2 $s from S. America
found on Myopotamus coypii (ex Rothschild coll., no particulars).

Ixodes hexagonus var. cookei (Packard, 1869).

Syn. : Ixodes cookei Packard, 1869 a, \i. 67.

Ixodes hexagomis var. longispinosus Neumann, 1901, p. 283.

Ixodes cookei Packard, in Banks, 1908, pp. 28, 29, PI. IX, Figs. 1-5. (Larva,

dorsum; palp of o (surely wrong!); $ capitulum, tarsus 1.) Banks, 1908,

p. 53, moreover, includes the following in his synonymy of /. cookei.
Ixodes cruciarius Fitch, 1871, p. 366. Banks, 1904, gives a poor figure (No. 81)

of cajjitulum, scutum and spiracle.
Ixodes hexagomis in Salmon and Stiles, 1902, p. 467 (not /. hexagonus Leach,

fide Banks).

Female : Coxa I provided with a long spine, which partially overlaps
coxa II. The short spines of coxae II-IV a little shorter than in the
type.

Evidently the N. American form of /. hexagonus.

184 Grenus Ixodes

Neumann examined 43 ^/'s and 127 $s from various mammals in
diflferent parts of N. America, and could find no other constant characters
distinguishing them from the type. Banks considers that the porose
areas of the % are shorter and less triangular than in the type, but in
the specimen we have seen this is not noticeable.

Neumann (1899, p. 130) re-described /. hexagonus, giving it a long
spine on coxa I. Later (1901, p. 283), he divides hexagonus into three
varieties : the type, longispinosus and inchoatus. Banks (1908, p. 32),
points out that inchoahis had already been named canisuga by Johnston
in 1849. We agree with this opinion, but consider the form sufficiently
different from hexagonus to deserve specific rank. There remains the
question of hexagonus var. longispinosus. Banks (1908, p. 29) has
compared Packard's type of /. cookei with Neumann's types of I.
hexagonus var. longispinosus, and finds them identical. Dr L. 0. Howard
has very kindly sent us two alleged $ s of /. cookei from the collection
of Marx, identified by Banks. Unfortunately, the two specimens are
quite clearly of different species, but one of them is no doubt the form
intended. It agrees very closely with the typical European hexagonus,
except for the stronger spine of coxa I, and is certainly no more than
a variety of hexagonus. Banks is, we believe, correct in recognizing its
identity with Neumann's var. longispinosus, but as Packard's /. cookei
has priority, it becomes /. hexagonus var. cookei.

We therefore have :

Ixodes hexagonus Leach.

„ „ var. cookei Packard (Syn. var. longispinosus Neumann).

„ canisuga Johnston (Syn. hexagonus var. inchoatus Nn.).

N.B. We are convinced that the /. hexagonus in many collections
of ticks will prove, on further examination, to be /. canisuga.

Neumann, 1901, p. 283, states that "var. longispinosus" was estab-
lished by him on the basis of 42 (/s and 113 $ s, found on Lutra, Mustela
vison, sheep, Texsis; Spermophilus; Felis domestica,'M.a,me; /oa;, Colorado ;
weasel, porcupine and marmot (Smithsonian Institution and Bur. Animal
Industry, Washington, D.C.).

Banks, 1908, p. 31, states that specimens from the pocket gopher,
Iowa, and Spermophilus, District of Columbia (Hassall coll.), labelled
/. hexagonus by Neumann, are referable to cookei.

We have a $ in our collection (N. 717) from Iowa, U.S.A.
(ex Marx coll., presented by the U.S. Department of Agriculture), and
have determined a $ from a dog, Calabogie, Ontario, for Dr C. G. Hewitt,
Government Entomologist, Ottawa (R. M. Reid coll., v. 1908).

/. nitens

185

15. IXODES NITENS Neumann, 1904.

Fig. 178 (original).
Lit. : Neumann, 1904, pp. 459, 460 (no figure).

Male : unknown.

Female : Body short oval, narrower in front, broader towards the
posterior third, brownish, L. 2'5 to 3'.5 mm., W. Vb to 1"9 mm. (partly
fed). Scutum oval-lozenge-shaped, with antero-lateral borders almost
straight, slightly emarginate, longer than wide (1 x 0'8 mm.), scapulae
pointed ; glossy ; cervical grooves obsolete, lateral grooves faintly
indicated ; some very sparse and very fine punctations in the anterior
angles. Dorsum bearing a very few short scattered hairs ; marginal
groove shallow. Venter almost smooth ; vulva facing coxae IV ;
genital grooves straight and diverging ; anal grooves rounded in
front of the anus, scarcely diverging. Spiracles small, oval, whitish.
Capitulum narrow, 7.50 /u, long ; base triangular, posterior border
concave, twice as long as broad dorsally ; auricula forming a flattened
retrograde horn pointing outward at 45°, Porose areas oval, separated,
broader than long. Hypostome long, narrow, sharp, with numerous
anterior denticles, then 3 | 3 followed by 2 j 2 rows of teeth. Palps
long, narrow, cultriform, the second article almost double the third.
Legs : coxa I with two spurs, the internal longer, scarcely reaching
to coxa II ; a very short spur externally on coxae II-IV. Tarsi long,
slender, tapering gradually ; pad almost as long as claws.

Fig. 178. I. nitens ? : capitulum (mutilated) and scutum, capitulum in ventral aspect,
part of venter, spiracle, tarsus 4, anal groove. The figures to the left sketched, thoB?
to the right drawn to scale. (Original, from the type, G. H. F. N. del.)

Nymph: Similar to the $, but genital pore and porose areas
absent.

Our description differs somewhat from Neumann's.

This species is based on two $ s and one o found on Mus macleaH at
Christmas Island, Pacific Ocean (N. C. Rothschild coll.). Our figure is
drawn from a type in our possession (N. 360).

N. I. 13

186

Genus Ixodes

16. IXODES BICORNIS Neumann, 1906.

Fig. 179.
Lit. and Icon. : Neumann, 1906, pp. 196, 197, Fig. 1 (reproduced).

Male : unknown.

Female: Body oval, red-maroon, 4x 2*1 mm. (partly fed), broadest
toward the posterior third. Scutum oval, 1'7 x 1*5 mm., dark brown,
glossy, almost smooth, with extremely fine punctations, excepting about
20 large ones along the posterior border: cervical grooves barely visible;
lateral grooves marked by a slightly prominent ridge. Dorsum : very short
hairs near the lateral borders; marginal groove deep, limiting a prominent
fold. Venter bearing long, whitish hairs or else glabrous Vulva between
coxae IV. Anal grooves short, convex, divergent. Spiracles whitish,
circular. Capitulum long (1 mm.); base pentagonal, with sides slightly
diverging in front, cornua slight ; porose areas circular or slightly
elongated transversely, the interval almost equal to their diameter ;
auricula forming a stout spine which is slightly longer than broad.
Hypostome long, narrow, pointed, dentition 4 | 4 followed by 3 | 3. Palps
long (0*85 mm.), narrow, cultriform, article 1 salient ventrally, article 2
one and one-half times as long as 3. Legs long, maroon-brown, coxa I
with two spurs, the internal longer than the article, and partly covering

Fig. 179. I. bicornis ? : part of venter ; hypostome. (Neumann, 1906, Fig. 1.)

/. auritulifs 187

coxa II ; the external spur pointetl, slightly longer than broad, almost
parallel to the internal ; a short spur at the postero-cxternal angle of
the other coxae, much reduced on pair IV. Tarsi long, narrow, tapering
obliquely.

Description based on 3 $ s, from Atoyac, State of Guerrero, Mexico :
2 found on Felis onca L., the other on a child. This species, which is
called " CoDchuda" at Atoyac, is considered to inflict "a bite fatal to
children "(A. Duges coll., now in Neumann coll., Toulouse). /. bicornis
is very closely allied to /. fuscipes Koch (/. spinosus Neumann). We
possess (N. 687) 1 $ from Felis concolor (ex Rothschild collection ;
no further particulars as to origin) which we refer to this species.
Banks (1908, p. 27) may be right in regarding /. bicornis Neumann as
synonymous with /. diversifossus Neumann, 1899. In the absence of
a figure of /. diversifossus, we would point out that the descriptions
differ as follows :

/. diversifossus. I. bicornis.

Scutum: 1'3 x 1 mm. 1*7 x 1*5 mm.

Palps: Article 2 scarcely longer Article 2 one and one-half times as

than 3. long as 3.

Hypostome : 2 | 2 or 3 [ 3 4 | 4, followed by 3 | 3.

17. IXODES AURITULUS Neumann, 1904.

Figs. 180 and 181.

Lit., Icon, and Synon. : Ixodes thoraciais Neumann, 1899 ; not Ixodes

thoracicus C. L. Koch, 1844, as described in Neumann, 1899, p. 149.
Ixodes thoracicus Neumann, 1899, pp. 149-151, Figs. 22, 24 (reproduced).
I.vodes auritulus Neumann, 1904, p. 450.

Male : unknown.

Female : Body short oval, swollen, 7 x 5-5 ram., dark reddish brown,
legs reddish yellow. Scutum oval, with lateral borders almost straight
along their anterior half, 1-3 x 1 mm., glossy brown in front and
on the sides, yellow between the cervical grooves; the latter shallow,
almost attaining the posterior border ; lateral grooves indicated by a but
slightly prominent thickened border ; punctations very numerous, very
fine, equal ; short hairs near the lateral and anterior borders. Dorsum:
grooves normal; some short hairs. Venter: hairs more numerous.

13-2

188

Genus Ixodes

Vulva facing coxae III. Sexual grooves almost straight, divergent.
Anus rather near the posterior border ; anal grooves parallel. Spiracles
small, rounded, whitish, placed in front of half the body-length. Capit-
uluni of medium length (0*7 mm.) ; base broader than long, trapezoid, the
posterior border straight and blackish, the laterals convergent in front ;
porose areas large, circular; well-marked cornua; auriculae stouter,
closer together than cornua. Chelicera (?). Hypostome (Fig. 181)
rounded or truncated in front, of uniform width, bearing 6 | 6, then 5 | 5
rows of pointed teeth, the external rows longest. Palps: article 1 large,
forming a horn-like protrusion directed forward, concave externally
for the insertion of article 2 ; article 2 longer than 3, from which it
is not clearly separated. Legs medium ; coxae brownish, glossy,
bearing hairs at their posterior border; coxa I small, bifid, the outer
spur strong ; a short spur on the postero-external angle of coxae II-IV.
A short spur at the posterior distal border of the trochanters. Tarsi
long, narrow ; tarsus 1 scarcely shorter than articles 4 and 5 united,
without apparent pseudo-articulation ; tarsus 4 slightly shorter and
thicker ; all tarsi taper gradually distally ; pad | the length of claws.

Fig. 180. I. auritulus ? : capitulum and scutum ; V ventral and D dorsal aspects.

(Neumann, 1899, Fig. 22.)
Fig. 181. I. auritulus ? : hypostome, x 65. (Neumann, 1899, Fig. 23.)

/. hrunneiis 189

Description based on 4 $s collected at Punta- Arenas, Straits of
Magellan, S. America, by Lebrun, from an undetermined bird (Paris
Museum).

In the British Museum there are 2 $ s, from Trupialis militaris, San
Sebastian Settlement, Tierra del Fuego (Crawshay coll., October, 1906).
The better specimen is mutilated, articles 2-4 of the palps being absent.
The sides of the scutum are more angular than in Neumann's figui-e of
the type, and correspond more to our figure of /. brunneus (Fig. 182).

18. IXODES BRUNNEUS Koch, 1844.

Figs. 182, 183.

Lit. etc. : Lvodes brunneus C. L. Kcxjh, 1844, p. 232 ; 1847, p. 99, PL XX, Fig. 74

( $ from N. America).
Ixodes avisugus Berlese, 1890, fasc. LV, No. 5. Latin description. Figures of

dorsum, venter, capitulum and scutum.
Ixodes frontalis (Panzer), in Neumann, 1899, p. 133, Fig. 11 (reproduced).
Ixodes avisugus Berlese, in Canestrini, 1890, p. 505.
not Ixodes bifiircatus Neumann, 1899, p. 122, as stated by Neumann, 1901, p. 347,

in relation to a 9 specimen he sul)sequeutly named Dermacentor bifurcatus

(Neumann, 1904, p. 453).
Ixodes brunneus Koch, in Neumann, 1904, p. 454, who gives description, sent

to him by Dahl, of the type (1 ? ).
Ixodes kelloggi Nuttall and Warburton, 1907, p. 396, Figs. 6-8 (reproduced).
Ixodes brunneus Koch, in Banks, 1908, p. 26, PI. Ill, Fig. 9 ( $ capitulum and

scutum ; figures inaccurate).
'i Acarus frontalis Panzer, 1795, fasc. 59, Fig. 23.
1 1xodes pari Leach, 1815, p. 399, found on Parus major.
1 1xodes pallipes Koch, 1835-1844, Heft 39, Fig. 10, found on Sitta europaea

(Larva).
1 Ixodes sturni Pagenstecher, 1861, p. 40 (Larva).

Note : The foregoing synonymy is doubtful with regard to the names
preceded by (?), but in these cases the ticks were found on birds, and
may well have been brunneus. In the absence of any accurate figures, it
is easy to understand the confusion which has arisen regarding this .species.
The type, a single ? (Berlin Museum), has been mounted in balsam.
Professor W. Donitz, who has kindly compared the structures in the
type with specimens we sent him, agrees with us as to the identity of
kelloggi with brunneus. Banks (1908, p. 26), after seeing our figures
of kelloggi, expressed a similar opinion. We agree with Neumann in
degrading avisugus to a synonym ; our specimens agree with Neumann's
specimens of " I. frontalis (Panzer) " Neumann, 1899.

190 Genus Ixodes

The following description is the one which we gave of kelloggi, with
a few slight additions :

Male : unknown.

Female (fully fed) : Z. 7 to 8, W. 5-2 mm. Body brown, with
numerous white hairs. Scutum (Fig. 182) very long, 1'4 x 1*1 mm.,
somewhat diamond-shaped, with a few conspicuous white hairs ; scapular
angles sharp, no lateral grooves ; cervical grooves well marked but not
deep, slightly converging at first, then diverging to reach the margin
nearly half-way along its postero-lateral border ; several large shallow
punctations, in part confluent, from some of which hairs have fallen.
Venter : with numerous white hairs ; spiracles slightly oval, brown,
macula central ; vulva between coxa III ; anal grooves parallel or
slightly divergent, rather short, terminating far from the posterior
border. Capituliim (Figs. 182, 183) of medium length (08 ram.) ; base
triangular, cornua slight ; porose areas large, transverse, of indefinite
form; palps rather long and narrow, article 2 the longest; blunt auriculae;
ventral base broad and rectangular posteriorly ; hypostome pointed, 3 | 3,
followed by 2 I 2 ^ Legs (Fig. 182 B) medium, coxa I with two spurs wide
apart, coxae II-IV with short outer spurs and inner edge trenchant ;
tarsi tapering (as in 7. pilosus ^ , Fig. 217).

Nymph : resembles the % (Neumann).

Larva : Scutum as broad as long (Neumann).

The species occurs on birds in Europe, Africa and North America.
The type in the Berlin Museum (1 gorged $) was found on Fringilla
alhicollis Gmelin, from North America. The larva described by Koch
(1844) as /. pallipes was found on Sitta coesia by Panzer (1795), and
Leach (181.5) found the $ on Parus major. Berlese (1890) states that
he frequently encountered this species on Passerines, fixed to the head
near to the auditory orifices, and in the country about Florence and
Venice on Anthus arhoreus, A. pratensis, Erythacus rubicula, Meinda
nigra. Canestrini (1890) possessed two specimens found on Praticola
ruhetra and Emberiza schoeniclus.

The description oifrontalishy Neumann (1899) is based on specimens
collected in Corsica on Turdus (E. Simon coll.); 3 ?s from Crex pra-
tensis from the Lower Loire (Trouessart coll.), from Saxicola rid)icola

1 In Neumann's figure (Fig. 183 of frontalis, here reproduced) the dentition ranges
from 4 I 4 to 3 1 3, 2 I 2. The shape of the hypostome is different from ours, possibly owing
to pressure of the coverglass on the specimen. His Fig. C (basis capituli and coxae I) is
obviously drawn from a balsam mount, and consequently somewhat misleading, as such
preparations generally are.

/. hrimneiis

191

(Paris Mus.), Turdus aonalaschkae Pallasi, from Baltimore (Bureau of
Auimal Industry) ; 1 o from a goose in Paris (Trouessart coll.) ; 1 larva
from Tardus merula from Saintes,

Fig. 182. I. hrunneus ? : ^, capitulum and scutum, x 20 ; D, coxae I-IV ; C, capitulum
in ventral aspect, x 70. (Nuttall and Warburton, 1908, Figs. 6-8 of I. kelloggi.
G. H. F. N. and E. W. del.)

Fig. 183. I. hrunneus ?: A, digit, x 220 ; B, hypostome, x80; C, ventral aspect of
basis capituli and coxae I, x 40. (Neumann, 1899, Fig. 11, of /. frontalis (Panzer).
See footnote to p. 190. )

192 Greniis Ixodes

We possess, or have examined, females from France : (N. 523) from
Passer montanus, St Ge'nis cle Malgoire (Gard), XI. 1908 (A. Hugues
coll.; presented by Hod. N. C.Rothschild); England: (N. 224) from owl,
Cambridge, 1904 (G. S. Graham-Smith) ; California : (N. 279^ from
Cyancitta stelleri frontalis, xii. 1895 (R. C. Snodgrass) ; (N. 280') from
thrush, Palo Alto, xi. 1895 ; (N. 282) from Piranga ludoviciana Sunol,
V. 1896 ; (N. 283) from Habia melanocephala Sunol, v. 1896 ; from " a
bird\" and from Carpidacus sp., Palo Alto, V. 1891 (Snodgrass coll., now
in Leland Stanford University coll.). We are indebted to Professor
V. L. Kellogg for the specimens from California.

A specimen ( $ ) has recently reached us from Africa, having been
found on a dove (Hajilopelia johnstoni Shelly) at Mlange Mountain,
Nyasaland (5500 ft.), v. 1910, by S. A. Neave (No. 150, Ent.
Committee ; determined by us).

Banks (1908) records 2 $ s from a tufted tit, Raleigh, North Carolina
(Brimley coll.) ; 1 $ from a hermit thrush, Baltimore, Maryland (Hassall
coll.) " this being the specimen named by Neumann Ixodes frontalis " ;
1 $ from a " chipping bird" probably at Amherst, Mass.

Note: Neumann (1899) describes a form which he regards as a
variety, as follows, the same being a young $ specimen found at Toulouse
on Ramphocelus coccineus, from Santa Fe de Bogota (Lacomme coll.) :
Body 5x3 mm. (capitulum included), reddish brown. Scutum oval,
longer (1*6 mm.) than broad, darker reddish brown, glossy, with sides
rounded, posterior angle broad; cervical grooves broad, shallow, only
attaining one-half the length ; lateral grooves indicated by their external
prominence ; slightly curved, attaining the limits of the posterior angle.
Punctations large, very numerous, only absent in front between the
cervical grooves. Surfaces finely shagreened, the dorsal glabrous, the
ventral beariug a few white hairs. Spiracles whitish, in front of half
the body-length. Capitulum 1 mm. long. (Digit 145 /u. long.) The rest
as in the type.

19. IXODES CAVIPALPUS Nutt. and Warb. 1907.

Figs. 184-180.
Lit. and Icon. : Nuttall and Warburton, 1907, p. 396, Figs. 1-5 (reproduced).

1 Described by us as /. kelloggi.

/. cavipalpKs

193

Male (Figs. 184, 185) : L. 19, If. 1-2 mm. Body long-oval (Fig. 184).
Scutum accurately long-oval, broadest in the middle, glossy, hairless ;
marginal fold very narrow in front and gradually broadening posteriorly ;
no lateral grooves; cervical grooves' and punctations very faint. Venter
brown with long hairs ; se.xual orifice between coxae III ; pregenital
shield longer than broad, narrow in front ; anal grooves slightly
divergent ; anal shields slightly broader in front ; spiracles oval, white.
Capitulum (Fig. 184 J., B) short (0'5 mm.), like that of /. ricinus in
general appearance ; basis capituli with median posterior point ventrally ;
hypostome with six well-marked transverse ridges of teeth giving the
effect of dentition 3 | 3 ; large basal teeth. (Digit ; see Fig. 185 B.)
Legs moderately strong ; coxa I with a slight spur, coxae II-IV
unarmed, but for a very slight tubercle at the external angle of coxa
IV ; tarsi tapering gradually.

_ A B

Fig. 184. I. cavipalpus <? : dorsum ; A and B : capitulum in dorsal and ventral aspects.
(Nuttall and Warburton, 1908, Figs. 1 and 2.)

A B

Fig. 185. I. cavipalpus: {A) i and ? spiracles, respectively, measuring 370 yu and 270 /x/.
orientated so that the tick's capitulum was to the left and the dorsum above ; the ?
spiracle shows a slight angularity postero-dorsally. {!>) i and ? digits, respectively,
138 /x and 140 /xZ., viewed from the dorsal surface. (Nuttall and Warburton, 1908,
Figs. 4 and 5.)

' Accentuated in Fig. 184.

194

Genus Ixodes

Female (Figs. 185, 186), more or less fed : L. 2-9 to 5, W. 1-8 to
3'3 mm. Body brown, with a few scattered hairs. Scutum {I. 1'3 to 1"4,
w. 1 to 1*2 mm.) dark brown, long oval with slightly sinuous posterior
border ; cervical grooves shallow, almost reaching the margin ; no lateral
grooves ; numerous fine shallow punctations. Vente?- brown, with
scattered hairs, long at the sides ; vulva between coxae IV ; anal grooves
nearly reaching the posterior border and slightly divergent ; spiracles
round, whitish. Capitulum (Fig, 186) fairly long (09 mm.), base tri-
angular, with piriform porose areas far apart ; palps with article 2
concave externally (whence " cavipalpus ") owing to lateral prominence
proximally ; hypostome long and narrow with a crown of small teeth
followed by dentition 2 j 2. (Digit; see Fig. 185 B.) Legs as in ^f, but
coxae I-II close together, III distant from II, and IV more distant from
III ; coxa I pointed, II-IV with straight posterior borders.

Fig. 186. /. cavipalpus ? : capitulum in (A) dorsal and (B) ventral aspects. (Nuttall
and Warburton, 1908, Fig. 3.)

Description based on (N, 245) 2 ^/'s and 7 $s from a baboon (Cyno-
cephalus habuin), Kansanshi, N.W. Rhodesia, 1. 1907 (Dr A. Yale Massey) ;
(N. 353) 1 (/" from a native haby, Benguella Hinterland, Angola, xii.
1907 (Dr F. Creighton Wellman) ; and (N. 394) ^b, % s from the last-
named source, 1908.

This species strongly resembles /. pilosus (p. 221), but may be
readily distinguished from it by the anal grooves and the absence of
a lateral groove on the $ scutum.

/. cuujiistns

195

20. IXODES ANGUSTUS Neumann, 1899.

Figs. 187-190 (original).

Lit. etc. : Ixodes angustus Neumann, 1899, p. 136 (no figure).

not Ixodes angustus Neumann, 1901, p. 284, Fig. 4 (digit of 9 ). In this pai)er
Neumann re-described what he took to be angustus from 2 $ s found on
Didelphys azarai, Buenos Ayres (C. Berg coll.). Banks (1908, v. infra) has
pointed out tliat this second description relates to an entirely different form
which Neumann (1910, p. 30) has since recognized as /. loricatus. In con-
sequence of this mistake Neumann (1910) suppresses the species angustzis,
but this does not appear to us to be permissible, for the type specimen first
described (1899) still exists, and is a perfectly distinct species.

Ixodes angustus Neumann, in Banks, 1904, p. 331 ; 1908, p. 29, PI. IV, Fig. 2
( $ capitulum, scutum, coxae and spiracle ; sketchy, but not very inaccurate).

Male^ (Fig. 187) : oval, rather narrower in front, broadest in the
middle. Scutum (1"4 x '9 mm.) convex, marginal fold moderate;
numerous fine, shallow punctations and short white hairs; a rather
well-defined pseudo-scutum ; cervical grooves shallow, divergent ; lateral
grooves indicated by slight marginal ridges on the pseudo-scutum.
Venter : genital orifice opposite the second intercoxal space ; pregenital
plate of irregular form, short; median plate sub-trapezoid, longer
than broad, narrower in front ; anal plate with sides very slightly
diverging ; adanal plates rhomboidal. Spiracle sub-circular, its ventral
borde'- somewhat pointed. Capitulum: base trapezoid, broader in

Fig. 187. I. angustus s : dorsum, venter, spiracle and tarsus 4. From specimen
(N. 1065) found on squirrel, Mt Lehman, British Columbia, S. Hadwen coll.
(Original, G. H.F.N, del.)

^ Banks, 1908, p. 30, appears to have seen the male ; his description covers five lines
and is quite inadequate. The o and larva have not been hitherto figured or described.

196

Genus Ixodes

front, punctate ; palps short, with very convex inner contour and
slightly convex outer contour ; article 2 broader anteriorly than long,
article 3 about equal in length to article 2, article 1 somewhat sharply
prominent ventrally ; hypostome small, armed only on its anterior third
with 3 I 3 rather obscure sub-equal teeth, about 4 per file. Legs : coxae
all bidentate, the inner spur of coxa I fairly long and strong; small
internal and external spurs on coxae II-IV, internal spur on IV almost
obsolete. Tarsi moderate, tai-sus 4 tapering obliquely ; pad long.

Female (Fig. 188) : Body elongate, with numerous fine punctations
and short white hairs. Scutum (15 x I'l mm. Nn.) much longer than
broad, broadest at the anterior third, where there are distinct lateral
angles ; cervical grooves shallow, long, diverging gradually ; lateral
grooves indicated by straight ridges reaching the border near the
posterior end of the scutum ; numerous small punctations, most evident
posteriorly ; scapular angles very sharp and prominent ; a few scattered
hairs. Vente7': vulva between coxae III ; anal groove rather ogival in
front, with sides nearly parallel ; spiracle transversely oval. Capitulum :
base triangular dorsally, slightly longer than broad, the posterior border
straight, cornua absent. Porose areas large, of rather indefinite contour,
the interval less than their length. Palps long, article 2 half as long
again as article 3, article 3 rather pointed in front. Hypostome long,
sharply pointed, 2 | 2 then 3 | 3 sharp teeth. Legs as in the </", except
that the 'internal spine on coxa I is rather shorter and coxae III and IV
are without the internal spurs.

Fig. 188. I. a7igustus ? : capitulum and scutum, capitulum in ventral aspect, coxae,
anal groove, spiracle and tarsus 4. Drawn from specimens N. 506 and 991.
(Original, F. M. H. del.)

/. angiistus

107

Nymph' (Fig. 189): differs remarkably from the % in several
respects. Scutum as broad or broader than long, with lateral angles, and
with lateral grooves not attaining the posterior border. GcipituluTn :
base with pointed cornua, and large, laterally projecting auriculae ;
palpal article 1 projecting forward and inward in a long pointed process.
Legs : coxae resembling those of adults ; tarsi short, slightly humped
distally, where they taper abruptly.

Larva' (Fig. 190): resembles the o to some extent. Scutum broad,
without lateral grooves, the cervical grooves attaining the posterior
border; emargination very slight. Capitulum: base devoid of cornua
and auriculae ; palpal article 1 projecting forward as in the o, but also
outward and backward ventrally.

Fig. 189. Fig. 190.

Fig. 189. /. angxistuH o : capitulum and scutum ; capitulum in ventral aspect with

coxae, tarsus 4, anal groove and spiracle. Drawn from specimens (N. 1134) found on

squirrels, Mt Lehman, British Columbia, Canada, 1910 ; S. Hadwen coll. (Original,

G. H. F. N. del.)
Fig. 190. 1. angustus larva : dorsum and venter, tarsus 3. Same origin as the nymph

in Fig. 189. (N. 1134. Original, G. H. F. N. del.)

Described from 1 ^ (N. 106.5) from Sciurus hudsonvus douglasi,
Mt. Lehman, British Columbia, Canada (S. Hadwen coll., iii. 1910), and
several $ s found on the same host and in the same locality (T. Bowhill,
XII. 1908, and S. Hadwen, xii. 1909 and ix. 1910, N. 506, 991, etc.).
Mr S. Hadwen has sent us (N. 119.5) a cT and % which copulated in
confinement, the j/* having been taken on Lepus dalli and the $ on the

1 Were it not for the kindness of S. Hadwen, D.V.Sci., who has raised this tick
through the various stages and sent us the specimens he has raised, we should not have
been in a position to associate the nymphal and larval form with the adults. It is true
that Dr Hadwen had found the immature forms on the same hosts with the adults, but
they differ so much from the latter that we could not refer them to angustus ou the
strength of their structure.

198 Genus Ixodes

squirrel aforementioned, viii. 1910. We have also received a $
(N. 678) from Tamias toivnsendi, Chilliwack, British Columbia
(ill. 1900, N. C. Rothschild coll.), a $ (N. 284) from Sciurus sp.. La Honda,
California (ii. 1895, V. L. Kellogg coll.), a ? (N. 715) from ? host, Glacier
Bay, Alaska, iv. 1909 (gift of the U.S. Department of Agriculture), and
finally a ? from "a mouse, Jeffrey, N.B., C. H. McNutt, x. 1908" (sent to
us by C. G. Hewitt, Ottawa, Canada, for identification). Neumann (1899)
first described the species from 1 $ (mutilated, without complete
capitulum and legs) from Neotoma occidentalis, Shoshone Falls, Idaho
(coll. Bureau of Animal Industry, Washington, D.C.). Banks (1904)
records 2 $ s " from Siskiyou County, California, also received from
Massett, British Columbia."

See further under Notes on Biology (p. 315).

21. IXODES CALEDONICUS Nuttall, 1910.

Figs. 191-193.
Lit. and Icon. : Nuttall, xii. 1910, pp. 408-411, Figs. 1-3 (reproduced).

Male : unknown.

Female (gorged): Body 7'8 x 4 mm., oblong, with sides almost
parallel, posterior border broadly rounded, covered with a few short
white hairs. Scutum (1'3 x 1 mm.): glossy, sub-oval, longer than broad,
emargination slight, cervical grooves beginning as deep pits (giving the
effect of sharp scapulae), then deep and distinct for about two-thirds the
length, but slightly divergent and fading away toward the postero-
lateral borders ; no lateral grooves ; very fine, uniform punctations,
except for a few larger ones along the anterior and antero-lateral
borders ; a few very small, short hairs. Capitulum : base sub-rectangular,
broader than long, with slightly concave raised and trenchant dorsal
ridge continuous with the slight (trenchant) cornua; porose areas not
depressed, large, ovoid, almost confluent. Palps short, far apart basally,
converging and rounded distally, with thick internal border and broadly
rounded tips, articulations between articles 2 and 3 obsolete; thumb-like
in profile ; ventral surface of basis capituli flattened, pentagonal, blunt
auriculae protruding laterally; palpal article 1 with ventral angle ; hypo-
stome inclined ventrally, rounded in front, dentition 2|2, with 8-9 blunt
teeth per file, and a narrow unarmed median ridge. Vente?^: vulva slightly
posterior to a line connecting the posterior borders of coxae II ; genital
grooves rounded in front, diverging slightly, then sub-parallel, finally
diverging slightly to the posterior border ; anal grooves rounding the

/. caledonkas

199

anus anteriorly, then parallel, and slightly divergent behind. Spiracle
circular, macula median. Legs: coxa I visible dorsally, bidentate ;
with stout external spur and short internal spur ; a stout external spur
on coxae II and III, smaller on coxa IV ; trochanters 1-3 with short
spur at postero-external angles; tarsus 4 long, tapering obliquely at
the distal third of its terminal portion ; claws much longer than the
pad.

Fig. 191. I. calcdonicus ? : caiDitulnm aud scutum, capitulum in ventral aspect with
coxae, anal groove, tarsus 4 and sijiracle. (Nuttall, 1910, Fig. 1, F. M. H. del.)

Nymph (Fig. 192) : differs but slightly from the ? : a few short
(caducent) hairs on the scutum, longer hairs, in moderate numbers, on
the body. Scutum as broad as long (0'7 x 07 mm.) with lateral angles
rounded, with antero- and postero-lateral borders sub-rectilinear; cervical
grooves attaining the posterior border. Capitulum with marked,
trenchant cornua protruding outward and continuous with the turned
upward dorsal ridge. Hypostome 2 | 2, with eight pointed teeth in the
external files. Venter : spiracle bluntly oval. Otherwise resembling
the ?.

200

Genus Ixodes

-D

Fig. 192. I. caledonicus o : capitulnm and scutum with dorsal aspect of first leg-pair,
ventral aspect of capitulum with coxae, tarsus 4, anal groove and spiracle. (Nuttall,
1910, Fig. 2, F. M. H. del.)

Larva (unfed, Fig. 193) : Body 0'8 mm. long, resembles the $
and o in its chief characters (coxae, trochanters, tarsi). Scutum more
rounded (0*4 x 0'4 mm.) than in the o, with deep cervical grooves.
Capitulum : hypostome 2 j 2, with six teeth in the external files.

¥5.

Fig. 193. I. caledonicus larva : capitulum and scutum, ventral aspect of capitulum with
coxae, tarsus 3, anal gi-oove. (Nuttall, 1910, Fig. 3, F. M. H. del.)

Described from 1 ^ (N. 961) found on rocks below a dove's nest,
Fastcastle, Scotland, 6. ix. 1909, by Messrs J. F. Cormack and
J. Waterston and communicated by Dr J. H. Ashworth (Edinburgh) ;
1 o and 3 larvae (N. 1142) found on young domestic pigeon, from a
dovecot at Dunipace, Stirlingshire, Scotland, 18. IV. 1910, and 1 $
and 4 OS (N. 1200) from the same source, 9. viii. 1910, communicated
by Mr William Evans (Edinburgh).

/. fossnlatus 201

22. IXODES FOSSULATUS Neumann, 1899.

Fig". 19-i (original).
Lit. ; Neumann, 1899, p. 120 (no figure).

Male: unknown.

Female : Body oval, narrowed in front, broadly rounded behind,
5x3 mm. Scutum oval, 1"3 x 1 mm., with rounded borders, glossy;
scapulae pointed, cervical grooves attaining the posterior border ; lateral
grooves but slightly curved, joining the border at about the posterior
third of the length ; margin raised ; many fine equal punctations, coarser
posteriorly in the median field. Dorsum shagreened by numerous
punctations ; glabrous, without marginal groove ; three grooves in the
posterior half. Venter shagreened like dorsum ; vulva narrow, facing
coxae IV ; anal grooves curved, not reaching the posterior border.
Spiracles distant from coxae IV, placed in front of the middle of the
body-length, oval, with short axis directed forward, macula slightly
eccentric. Capitulum with dorsal base twice as long as broad ; with
slight cornua ; porose areas large, ovoid, with long axis oblique, with an
interval equal to their width. Auricula forming a spine directed
backward and outward. Chelicera and hypostome ? Palps long, narrow.
Legs normal. Coxae small, not extending beyond the anterior quarter
of the body-length ; coxa I with two spurs, the internal longer; coxa II
conic.il internally ; a short spur at the postero-external angle of coxae
II-III, almost obsolete on coxa IV. Tarsi long, tapering abruptly near
their tips ; pads large, almost as long as the claws ; the latter short and
thick. Colour : body and legs reddish brown.

Fig. 194. I. fossulatus ? : Ccapitulum (mutilated) and scutum, ventral aspect of capitulum,
tarsus 4, coxae, spiracle and anal groove. Sketch from type specimen from Ecuador.
(Berlin Mus. Original, G. H. F. N. del.)

Our description of the type, a single dry specimen, from Ecuador,
S. America (Berlin Alus.), differs from that of Neumann in several
respects.

N. I. 14

202

Genus Ixodes

23. IXODES ACUTITARSUS (Karsch, 1880).

Fig. 195.

Lit., Synon. and Icon. : Haenudastor acutitarsus Karsch, 1880, p. 142 (Latin

description in five lines ; quite insufficient).
Ixodes laevis Neumann, 1899, p. 148, Fig. 21 (i-eproduced) ; recognized as

/. acutitarsus by Neumann (1901, pp. 285, 290) after examining Karsch's

type.
Ixodes acutitarsus Karsch, in Neumann, 1899, p. 180; 1901, p. 286.
Eschatocephalus acutitarsus Karsch, in Neumann, 1901, p. 290.

The following description is based partly on Neumann's /. laevis (Paris
Mus.), which we have examined. We see no reason why the species should have
been regarded as an Eschatocephalus, since the legs are not of inordinate length.

Male : unknown.

Female (unfed) : dark brown, very large, 7".5 x 3"8 mm. (Karsch's
type measured 8 x 3"3 mm.) ; body oval, dorsum convex, broadest behind
coxae IV, Scutum cordiform, as broad as long (2'5 mm., Nn.), the
antero-lateral borders slightly convex, one-half the length of the
postero-lateral borders which are straight or but slightly concave ;
rounded posteriorly ; cervical grooves parallel, then divergent, shallow,
attaining the postero-lateral borders ; traces of lateral grooves ; surface
glossy, a few fine punctations situated toward the anterior angles.
Dorsum convex, smooth ; marginal groove deep and broad, limiting a
broad marginal fold which is slightly incised by the dorsal extension of
the ventral grooves ; a few very short hairs. Venter : vulva facing
coxae IV, sexual grooves subrectilinear, divergent ; anal grooves rounded

Fig. 195. Z. acutitarsus ? : (1) dorsum ; (2) digit, 265 fi L, and (3) coxae. (Neumann,
1899, Fig. 21 of I. laevis Nn.)

/. ijigas 203

in front, widely divergent behind ; short hairs, mostly behind the coxae.
Spiracles large, subcircular, with macula eccentric (anterior). Gapitidum
long (2 mm., Nn.), with base slightly broader than long; dorsal ridge
subrectilinear (no cornua) and lateral borders parallel; porose areas
oval, with long axis oblique, the interval nearly equal to their width.
(Digit ; see Fig. 195 (2).) Hypostome narrow, lanceolate, as lung or
slightly longer than the palps; dentition 4 | 4, then 212 rows of 10-12
teeth, which are widely separated by a median ridge. Palps long ;
external border concave posteriorly, article 1 projecting outward like a
knot, article 2 twice as long as 3. Legs large, strong (pair 4 extend
one-half their length beyond the posterior border of the body in unfed % ),
and bearing short hairs. Coxa I bidentate, the internal spur thick and
overlapping coxa II, the external spur short ; a short external spur on
coxae II-IV. Tarsus 4 tapering gradually ; pad about one-half as
long as claws.

Origin: Karsch's type came from Japan (1 $ , D. Hilgendorf coll.),
Neumann's /. laevis came from Sikkim, India (1 $, Harmand coll.,
Paris Mus.). Neumann has since recorded 2 $s from Sikkim (Berlin
Mus.), and we have seen 2 $s from Japan and S. Formosa (Brit. Mus.).

24. IXODES GIGAS Warburton, 1910.

Fig. 196.

Lit. and Icon: "Warburton, 1910, p. 397, Figs. 1 and 2 (reproduced).

Male: L. 5, W. 3 mm. Body oval, broadest near the hind end,
marginal fold uniformly broad. Scutum chestnut coloured, darker on
the scapulae and sides, glossy and smooth except for a few small
punctations on the scapulae and between the cervical grooves ; the
latter parallel at first, then sharply diverging ; no lateral grooves.
Capituhmi moderate ; base rather long, pentagonal, without cornua.
Palps of medium length, article 2 rather prominent laterally and about
twice the length of article 3. Hypostome 2 ] 2, seven or eight teeth
per file. Venter : pregenital plate elongate, rather indented anteriorly ;
median plate much longer than broad, the sides slightly diverging ;
anus rather near the posterior border ; anal plate with sides curved
and slightly diverging ; genital orifice facing the third intercoxal space.
Spiracles rather large, oval or slightly reniform. Legs long ; coxae I

14—2

204

Genus Ixodes

bidentate, like that of a Rhipicephalus or Hyalomma, protruding in
front of the body and visible dorsally ; coxae II-IV with a slight
external spur and a blade-like internal edge; all the coxae close
together and occupying little more than the anterior third of the
body-length ; tarsi long and tapering, only slightly gibbous dorsally.
Legs 4 extend beyond the posterior end of the body by their two distal
articles.

Fig. 196. I. gigas i : dorsum and venter. (C. Warburton, 1910, Figs. 1 and 2.)
Tlie anal grooves are somewhat too divergent in the figure.

Female : unknown.

Described from two specimens in the India Museum, Calcutta
(No. -Y§- and No.?), taken at Punkabani, Darjiling District, E. Himalayas
(no host recorded).

This fine species — the largest male Ixodes known — may very likely
prove to be the cT of Ixodes acutitarsus (Karsch, 1880), but it is unsafe
at present to attribute it to that species. Its coxal armature is unique
in this genus.

25. IXODES RUBICUNDUS Neumann, 1904.

Figs. 197 (Nn.) and 198 (original).

;Lit. and Icon. : Neumauu, 1904, pp. 460-462, Fig. 2 (reproduced).

Howard, C. W., viii. 1908, p. 97, PL IV, Fig. I (a, b), translation from Neumann
and copy of his figure.

Male (Fig. 197) : Body 2"3 mm. long (capitulum included), contour
oval, narrower anteriorly, 1"2 mm. broad toward the posterior third,

/. ruhicundus

205

maroon-brown. Scutum convex, glossy, glabrous, marginal fold broader
posteriorly than on the sides ; cervical grooves narrow, superficial,
divergent, extending to about the middle of the body-length ; numerous
punctations, small, unequal, regularly distributed. Venter: genital
pore broad, between coxae III. Pregenital plate long, triangular;
median plate much longer than broad, with sparse, shallow punctations ;
anal plate oblong, sides parallel, longer than wide, very finely punctate ;
adanal plates longer than broad, sides parallel ; no hairs. Spiracles
circular. Capitulum short (0-5 mm.) ; with trapezoid base, broader
anteriorly and broader than long, no cornua, posterior angles pro-
truding on ventral surface, separated by a similar median protuberance.
Chelicera thick, 135 fx, long (dorsal process with two stout, suc-
cessive teeth, the posterior stouter ; external article with three teeth,
the anterior small, the posterior very stout). Hypostome broad, the
teeth fused on each half in four transverse ridges of three or four
denticles, followed by a row of four or five teeth and on each side by
a very stout tooth. Palps short, articles 2 and 3 of about the same
length. Legs of medium length, brick-red. Coxa I with internal spur
almost obsolete, coxae II-III unarmed but trenchant, coxa IV also
trenchant but with a slight postero-external spur. Tarsi of medium
length, tapering gradually.

Fig. 197. I. ruhicundus s : venter and hypostome. (Neumann, 1904, Fig. 2.)

206 Genus Ixodes

Female (gorged, Fig. 198): Body elongate oval, may attain L. 10 mm.,
W. 6*5 mm. toward the posterior third, brick-red in specimens of small
size, maroon-red in others. Scutum a short oval, scarcely longer than
broad (1"6 x 1-4 mm.), with lateral borders at first slightly convex and
diverging, then rectilinear and converging to the rather narrow posterior
angle ; emargination slight, cervical grooves well marked, reaching
almost to the lateral borders at their re-entering angles ; lateral grooves
well marked up to the lateral border, numerous fine punctations,
slightly unequal, rarer in the lateral fields ; surface glossy, glabrous,
maroon-brown. Dorsum with sparse hairs (caducent) ; punctations very
superficial; three posterior grooves. Venter with similar punctations
and hairs ; vulva between coxae IV^; anus fairly anterior; anal grooves
long, parallel, united in a flat curve in front of the anus. Spiracles
small, circular, whitish. Capitidum 0*85 mm. long; dorsal base sub-
rectangular, much wider than long ; porose areas piriform, oblique,
converging anteriorly, placed near the lateral borders and separated by
a space equal to their width ; auriculae slight and blunt. (Chelicera
with digit 140 yu, long ; dorsal process long, parallel to the shaft ; external
article with 5 teeth growing progressively larger antero-posteriorly.)

Fig. 198. I. ruhicundus ? : capitulum (mutilated) aud scutum, spiracle, tarsus 4, anal
groove, coxae and base of capitulum, ventral aspect. From a gorged ? from Ovis aries,
Neumann coll. 1253. (Original, G. H. F. N. del.)

Hypostome long, narrow, sublanceolate, dentition 3 | 3, the internal file
short, of 6 or 7 teeth, the median almost as long as the external ; the
latter extending from apex to base, with 15-16 teeth. Palps long,
fairly slender, concave externally, second article longer than third.
Legs tapering, brick -red ; coxae I, II and III flat, unarmed, posterior
border trenchant ; coxae IV more swollen, with a small tuberosity on
the external third of the posterior border. Tarsi long, slender, tapering
gradually.

1 In the specimen examined by us the vulva is placed just behind coxae III.

/. simplex 207

Description based on two ^a and seven $s found on slieep in Eastern
Cape Colony (Lounsbury coll.). We have examined specimens kindly
lent us by Prof. Neumann and others (in W. F. Cooper's coll., determined
by us) from goats, Albert District, S. Africa, 1905 (H. E. F.). This
tick is supposed to inoculate an obscurely defined disease characterised
by anaemia.

Ixodes rubicundus var. limbatus Neumann, 1908.

The subspecies, according to Neumann (iv. 1908, p. 22), differs
from the type as follows:

Male : unknown.

Female: Scutum smaller (1"3 x 1'15 mm.) ; reddish brown, lighter
along the lateral borders ; cervical grooves very shallow ; lateral grooves
indicated by a slight external elevation ; punctations finer, subequal.
Capitulum smaller (0'73 mm. /.); hypostome with fewer teeth per
file.

Description based on six $s from Ovis aries and Capra hircus,
Katanga, Congo Free State (coll. of the Special Committee, Katanga).

26. IXODES SIMPLEX Neumann, 1906.

Fig. 199 (original).
Lit. : Neumann, 1906, pp. 197, 198 (no figure).

Male : unknown.

Female : Body oblong, with lateral borders parallel, slightly nar-
rowed on a level with the spiracles, rusty yellow, 4x3 mm. Scutum
longer than broad, 1*2 x 0*9 mm., broader toward the middle, the postero-
lateral borders slightly concave, red-brown or yellowish, glossy, with
widely separated punctations, which are fine and larger in the lateral
fields ; cervical grooves obsolete ; no lateral grooves. Dorsum covered
by many long hairs. Venter : hairs short and fewer, except on the
sides. Vulva between coxae III. Anal grooves relatively short,
divergent, ogival in front. Spiracles brownish, subcircular. Capitulum
short ; base triangular, without cornua ; porose areas elongate trans-
versely, separated by a median depression, interval equal to half their
width; auriculae absent. Hypostome and chelicera? Palps short,
articles 2 and 3 of nearly equal length, about as broad as long ; article 1

208

Genus Ixodes

cylindrical. Legs slender, long. Coxae flat, all unarmed. Tarsi long,
tapering obliquely.

Fig. 199.

I. simplex ? : capitulum and scutum. Sketch from tjjje in British Mus.
(Original, G. H. F. N. del.)

Description based on one gorged $ , habitat unknown ; one $ and
one O from Rhinolophus ferrum-equinum (Schreb.) from Shanghai; one
$ from Vespertilio sp. (? from Gaboon, French Congo : British Museum).

27. IXODES JAPONENSIS Neumann, 1904.

Figs. 200, 201 (original).
Lit. : Neumann, 1904, pp. 458, 459 (no figure).

Male : unknown.

Female : Body oval, narrow in front, broader at the posterior third,
brownish yellow, L. 2, W. 1'5 mm. Scutum subcircular, as broad as long
(I'l mm.); cervical grooves far apart; lateral grooves limited by a very
marked ridge; very fine punctations; some very long hairs. Dorsum
bearing some long scattered bail's ; a distinct marginal groove. Venter
with numerous long hairs; vulva facing the last intercoxal space.
Genital grooves straight, divergent. Anal grooves slightly ogival, with
sides diverging greatly. Spiracles large, sub-circular, whitish. Capit-
ulum O'oo mm. long; base pentagonal, broader anteriorly, posterior
margin straight, with slight cornua, nearly twice as broad as long on
the dorsal surface ; porose areas well defined, oval, longer than broad,
the space between them equal to their breadth ; ventral surface

/. japouensifi and rauUi((/((

209

uniform, slight auriculae. Hypustonic long, niirrovv, 4 , -i near the tip
(corona), then 2 1 2 with a median unarmed ridge. Palps medium,
article 2 scarcely longer than 8. Leys : coxae unarmed ; coxa I with
internal angle sharp, all, especially coxa II, trenchant on their posterior
bordei'. Tarsi long, slender, tarsus 4 tapeiing gradually, the others
somewhat humped; pad almost as long as the claws.

Fig. 200.

Fig. 201.

Fig. 200. I. japonensis ? : capitulum in dorsal and ventral aspects. Drawn from the

unique type in the Paris Mus. (Original, G. H. F. N. del.)
Fig. 201. I. japonensis ? : dorsal aspect of body, venter, spiracle and tarsus 4. The

same specimen as the one to which the preceding figure relates. The scutum is

injured, there being a triradiate crack in the centi'e, indicated in the figure. (Original,

G. H. F. N. del.)

Our description is based on one $ (type) collected in the vicinity of
Tokio, Japan, by J. Harniand, 1901 (Paris Museum).

28. IXODES CANISUGA Johnston, 1849.

Figs. 202-209 (original).

Lit. and Synon. : Ixodes canisuga Johnston, 1849, p. 371 ; Evans, 1907, p. 36 ;
Banks, 1908, p. 32.
Ixodes phimheus, in Wheler, 1899 ; see Wlieler, 1906, p. 413.
Ixodes hexagonus var. inchoatus Neumann, 1901, p. 283; Wheler, 190G, p. 413.

We feel justified in reviving the overlooked species of Johnston,
to which attention was drawn by Mr William Evans (1907) to whom
we are indebted for an abstract of the author's original description.
The latter is not accompanied by a figure and is very brief, but he
distinguishes canisuga from hexagonus (to which it is "nearly allied"),
and describes the scutum of the former as "heart-shaped, with base
forward and widely emarginate, chestnut-brown, punctured, obsoletely

210

Genus Ixodes

foveolate on each side." Johnston's specimens were taken from the
pointer, and were sent to him as "the dog-tick." Neumann (1906), in
correspondence with Mr Evans, doubtfully refers to canisuga as possibly
synonymous with hewagonus var. inchoatus. Judging from the numerous
specimens which have reached us from Scotland, England and Ireland,
canisuga appears to be pre-eminently " the dog-tick " of this country
and we have no doubt, despite his insufficient description, that Johnston
had this species in view. Banks (1908, p. 32) appears to have come to
the same conclusion,

Male (Figs. 202-205): Broad oval. Scutum: 1-9 x ll to 2-3 x
I'o mm., oval, only slightly narrower in front, marginal fold fairly
broad ; very finely punctate, cervical grooves chiefly visible as elongate
divergent depressions, far from the anterior border ; lateral grooves
absent ; two other elongate, parallel depressions on either side of the
middle of the scutum. Emargination moderate ; scapular angles blunt
and rather rugose. Venter : genital orifice facing the second intercoxal
space, pregenital plate ill-defined ; median plate fairly broad, widest
just behind the spiracles; anal plate with sides only slightly divergent;
adanal plates long, with sides neai'ly parallel. Spiracle nearly circular.
Capitulum : very small ; base rectangular, broader than long dorsally,
the posterior border slightly concave ; no cornua ; a slight median ridge
or prominence. Palps very short, convex dorsally, article 2 barely
longer than article 3. Hypostome (Fig. 204) bifid, with about eight

Fig. 202. I. canisuga <? : dorsum and venter, x 20. The slight protuberances at the
postero-lateral angles of coxae II-IV omitted through an oversight. Specimen
collected in Northumberland, 1905, by E. G. Wheler. (N. 1079 ex W. F. Cooper coll.
Original, G. H. F. N. del.)

/. canisnga

211

fairly distinct lateral teeth on either side and corresponding ventral
crenulations (varies somewhat in structure, see Fig. 203). Lerjs : coxae
practically unarmed, coxa I only bluntly prominent at its internal angle,
slight protuberances at the postoro-lateral angles of coxae II-IV.
Tarsus 4 fairly long, prominent dorsally at some distance from its
extremity.

Fig. 203. I. canisuga i : capitulum in dorsal and ventral aspects, x 43. In the ventral
view the capitulum is tilted slightly backward. From the same specimen as Fig. 202.
(Original, G. H. F. N. del.)

Fig. 204. Fig. 205.

Fig. 204. I. canisuga <J : hypostome. Drawn from specimen (N. 1128) from Gowanburn,

N. Tyne, 1901. Presented by Mr E. G. Wheler. (Original, G. H. F. N. del.)
Fig. 205. I. canisuga 3 : legs 1-4 in profile, x 20. (Original, G. H. F. N. del.)

212

Genus Ixodes

Female (Figs. 206, 207) : unfed, 2-1 mm. l, may attain 8x5 mm.
Scutum : 1 X O'D to I'l x 1 mm., cordiform, scarcely longer than broad,
broadest much in front of the middle, postero-lateral borders straight
or somewhat concave; cervical grooves shallow, wavy, reaching the

Fig. 206. Pig. 207.

Fig. 206. I. canisuga ? : capitulum and scutum ; capitulum in ventral aspect. Found

on dog, Tynron, Scotland, by J. McMonnies, vii. 1908. (N. 447. Original,

G. H. F. N. and E. W. del.)
Fig. 207. I. canisuga ? (replete) : venter, spiracle, tarsus 4. Same specimen as in

Fig. 206. (Original, G. H. F. N. del.)

posterior border, lateral grooves short, the ridge external to them
terminating just beyond the lateral angles; punctations large and
confluent in the lateral fields, giving a rugose effect, moderate and more
discrete in the median field ; emargination moderate ; scapular angles
blunt. Dorsum only slightly punctate, with very short, white hairs.
Venter: vulva facing the second intercoxal space; genital grooves nearly
parallel at first, then separating, and terminating parallel to one another.
Anal grooves rounded or slightly ogival in front (often discontinuous),
with parallel sides. Spiracle sub-circular, but with major axis transverse.
Capitulum : base rectangular dorsally, twice as broad as long ; no
cornua; slight auricular ridges ventrally ; porose areas large, deep, oval,
the interval less than their diameter. Palps short, broad, article 2 about
equal in length to article 3 (asymmetry in Fig. 206). Hypostome 2 | 2,

/. eanisuga

213

about 7-8 teeth per tile, with a wide unarmed median area. Legs :
coxae as in the ^ ; tarsi distinctly humped.

Nymph and Larva (Figs. 208, 209): strongly resemble the $,
especially in the shape of the scutum, the legs, and ventral grooves.

Fig. 208. I. caniauga o : capitulum and scutum ; coxae and ventral aspect of capitulum ;
tarsus 4 ; anal groove and spiracle. Specimen from dog, Tynron, Scotland,
J. McMonnies coll., vii. 1908. (N. Ul. Original, G. H. F. N. del.)

Fig. 209. I. eanisuga, larva : dorsum, venter and tarsus .3. Specimen (British) received
from Mr E. G. Wheler. (N. 1044. Original, G. H. F. N. del.)

Our collection contains numerous specimens collected chiefly in this
country: — Scotland: (N. 414, $s), from cZo^, Thornhill, Dumfriesshire,
V. 1908 (J. R. Wallace); (N. 422, 447, ?s, os) from dogs, Auchenbrack,
Tynron, V. and vii. 1908 (J. McMonnies); (N. 402) $s from sheep,
Blackadie, Sanquhar, v. 1908 (R. Bramwell). [P^vans (i. 1907, p. 35)
found them on dogs; especially young collies, mostly attached about
neck, shoulders and sides, and behind forelegs, Pentlands and Moor-
foot districts, Scotland; os and $s (March, April); no ^T found.
(Det. by Neumann)]. England: (N. 1123, (/), Gowanburn, N. Tyne,
V. 1901, and (N. 104(), $) from dog, same locality, v. 1901 (E. G.
Wheler); (N. 1079, cf, %) from dog, Alnwick, Northumberland
E. G. Wheler); (W. F. Cooper's coll. % ), from dog, Stanhope, Durham,

214 Genus Ixodes

IX. 1905 (D. Robertson); (N. 501, %), from liorse, Cambridge,
V. 1908 (G. A. Banham); (N. 1068, (/s, ?s) from nest of GotiJe
riparia, Lyndhurst, Hants, vii. 1900 (N. C. Rothschild); (N. 605)
from Cotile riparia, Warsbrough Bridge, Barnsley, vii. 1906 (A.
Whittaker; ex N. C. Rothschild coll.); (N. 1051, ?), from grass, on
the Downs, Lizard, ix. 1909 (L. Cobbett) ; (N. 19, %\ from dog,
Longner Hall, Shrewsbury, iv. 1906 (R. F. L. Barton.) The last-
named gentleman has, moreover, sent us from the same locality (N. 499,
larva) from mole, XI. 1908; (N. 1137, ?s, o), from ferrets, vi. 1910,
and (N. 1141, 1143, many j/s, $s, os) from nests of three sand-martins
(G. riparia) on the banks of the River Severn, Shrewsbury, vii. 1910.
Ireland: (N. 62, $s) from dog, Sligo, ix. 1905 (M. F. Nuttall).
Germany : (N. 768, $ s), from dog, Kittendorf, Mecklenburg-Schwerin,
viii. 1908 (G. H. F. Nuttall). France: [Neumann, 1901, records (?)
" inchoatus" from the focc (R. Blanchard coll., Paris); from the
dog, Toulouse (Neumann coll.); and badger (Railliet coll., Alfort)].
N. America: [Banks, 1908, p. 32, records the species (2$s) as occur-
ring either on the rerf squirrel ov fox, but does not state the locality].
See further under Notes on Biology (p. 316).

29. IXODES TEXANUS Banks, 1909.

Figs. 210, 211 (original).

Banks, vi. 1909, p. 172, Figs. 16, 17 ( § capitulum, coxa I, tarsus 4
and spiracle. Inaccurate).

Male : unknown.

Female : Scutum as long as broad \ hexagonal, with posterior border
broadly rounded, broadest about the middle. Cervical grooves broad,
shallow, nearly parallel, far apart, reaching the posterior border, the parts
external to them, and especially the scapulae, being rugose, the area
between them pitted with deep punctations, more or less confluent.
No lateral grooves. Scapulae moderately prominent. Dorsum with
very short white hairs ; punctations inconspicuous. Capitulum : base
sub-trapezoid, narrower in front, very rugged, its sculpture rendering
the porose areas ill-defined. Porose areas irregular, separated by about
their breadth, the interval rugged ; no distinct auriculae. Palps rather
short and broad, with straight external and very convex internal contours,
the internal contour very rugose, giving a beaded effect ; article 1

^ The scuta of our three specimens measure respectively l"lxl-2, 1-2 x I'l, and
1x1 mm. Our most replete specimen measures 4-9 x 3-3 mm. Banks gives the body-length
at 6-5 mm., but does not state if this is the maximum length which may be attained in
gorged females.

1. texaniis

215

rather sharply prominent ventrally. Hypostome thick and strong, with
2 I 2 files of strong teeth along its whole length. Venter : vulva between
coxae III ; anal grooves rounded in front and very slightly diverging.
Spiracles small, sub-circular or short oval. Legs : coxae almost unarmed,
except that coxa I is slightly prominent at its internal angle, and there
is a faint indication of an external spur on all the coxae. Their
posterior borders are trenchant. Tarsi rather short and humped,

Fig. 210. I. texanus ? : capitulum and scutum, part of venter, spiracle, anal groove and
tarsus 4. Found on coon, Mt Lehman, British Columbia, i. 1910, by S. Hadwen.
(N. 992. Original, G. H. F. N. del.')

Fig. 211. I. texanus $ : capitulum and scutum, capitulum in ventral aspect, coxae,
anal groove, spiracle and tarsus 4. Drawn from specimen found on grey squirrel.
King's river, California, vrii. 1896. (N. 270. Pieceived from V. L. Kellogg and identified
by N. Banks. Original, F. M. Howlett del.»)

^ The differences shown in these figures are partly due to individual variations in the
specimens and partly to the position (this applies more particularly to the palps) in which
they were drawn.

21(3 Genus Ixodes

descending almost perpendicularly in tarsi 1-3, and with a slight
slope in tarsus 4.

Described from (N. 270) 2 $s taken from a Californian grexj squirrel,
King's river, California, vii. 1896 (V. L. Kellogg's coll.), and (N. 992)
1 % from Procyon lotor, at Mt Lehman, British Columbia, Canada,
I, 1910 (S. Hadwen). The two Californian specimens, which we were
unable to identify, were sent to Banks, who recognized them as a species
he has since described very briefly as /. texanus, from a raccoon, Live
Oak County, Texas (Mitchell and Bishopp).

30. IXODES STILESI Neumann, 1911.

Fig. 212.

Lit., Icon, and Synon. : Ixodes elegans Neumann, 1910, p. 191, Fig. 1 (reproduced).
Ixodes stilesi Neumann, 1911. Species renamed, elegans being preoccupied
(personal communication ; sec List of Condemned Species, p. 284).

Male unknown :

Female (unfed) : Body oval, sides sub-rectilinear, maroon-brown,
5 to 7 X 2 to 3 mm., slightly constricted on a level with the spiracles.
Dorsum bearing short sparse hairs, faint marginal grooves, and three
shallow longitudinal grooves. Replete specimens attain 9x7 mm.
(5 mm. dorso-ventrally),. ovoid or piriform. Scutum elongate, l"5-l"6.o x
1-1'14 mm., scapulae prominent, sides diverging as far as their anterior
fourth, then slightly converging and rounded behind ; dark brown, with
light median field, very glossy ; cervical grooves shallow, barely attaining
the posterior third ; lateral grooves faintly indicated by a straight ridge;
a few small punctations, chiefly in the lateral fields and along cervical
border, bearing minute hairs. Venter: hairs longer and more numerous
than on dorsum ; vulva facing coxae III ; anus distant from the posteiuor
border; anal grooves long, straight, parallel, uniting in a semicircle
anteriorly ; spiracles slightly oval, transverse. Capitulum 1*2 mm. I.,
base, viewed dorsally, pentagonal, lateral borders parallel ; cornua short
and pointed; porose areas oval, longer than broad, the interval barely
equal to their width; auriculae slight. Hypostome long, narrow, 3 | 3,
14-16 stout teeth per file, the inner files little shorter than the external ;
digit 210 fx I. (see Fig. 212). Palps 1 mm. I., narrow, cultriform ;
article 1 with veiitral point and a long basal hair ; article 2 nearly twice
as long as 3. Legs of medium length ; coxa I bifid, the spurs far apart,
somewhat flat, the inner spur slightly longer; a short postero-external
spur on coxae II-IV ; tarsi slightly humped distal ly ; pads large, almost
as lon<x as claws.

/. stilesi

217

Fig. 212. I. stilesi ? : capitulum and scutum, left digit in dorsal aspect, palp in
profile, inner aspect. (Neumann, 1910, Fig. 1.)

Described (condensed after Neumann) from 14 ? s (2 replete), found
on Pudua humilis (Cervidae), Chili (E. Porter coll., Paris Mus.).

31 . IXODES NEUMANNI n.sp. Nuttall and Warburton.

Figs. 218, 214 (original).

Lit. and Synon. Ixodes eudyptidis Maskell, 1885, in Neumann, 1899, pp. 128, 129
(no figure).
not Ixodes signatus Birula, 1895, p. 357.
not Ixodes parvirostris Neumann, 1901, p. 284.
Ixodes praecoxalis'^euvsi?inx\, 1899, p. 121; 1902, pp. 119-121; 1904, p. 451 (no

figures).
Ixodes intermedms Neumann, 1899, p. 1.32; 1904, p. 451 (no figure),
not Ixodes evdyptidis var. signata Neumann, 1904, p. 451.

Note: This species was described by Neumann, 1899, p. 128, as
Ixodes eudyptidis Maskell, without, as it seems to us, sufficient reason,
Maskell's description and figures being so exceedingly poor that it
is impossible to recognize the species with which he was dealing.
Maskell's type has not been available for study, consequently the only
safe procedure seems to be to describe the tick under consideration as a
new species, and we name it after Professor Neumann who utilised it

N. I. 15

218

Genus Ixodes

in his description of /, eudyptidis. Our description differs in some
respects from that of Neumann, who kindly lent us his types for purposes
of study.

Neumann (1904, p. 451) has himself degraded 7. praecoxalis and
I. intermedins to the rank of synonyms of his I. eudyptidis Maskell.
We recognize /. signatus Birula ( = /. parvirostris Neumann, 1901,
and I. eudyptidis var. signata Neumann, 1904) as a distinct species,
after examining the type specimens so named.

Male : unknown.

Female: Body attains 8 x 5"5 mm. Scutum dark brown, much
longer than broad {VI x 1'2 mm.^), hexagonal", with rounded posterior
border, broadest rather in front of its anterior third, slightly corrugated
on its anterior and antero-lateral borders, elsewhere marked with very
small discrete punctations ; cervical grooves shallow, broad, wavy (Fig.
213), the anterior commencement of lateral grooves faintly indicated ;
scapulae rounded and not prominent in front. Short white hairs present
in anterior portion, probably caducent elsewhere. Dorsum with a few

Fig. 213. I. neumanni ? : capitulum and scutum ; ventral aspect of capitulum, ana
groove, coxae, spiracle and tarsus 4. (There should be a slight ventral spur on
trochanters 1, 2 and 3). From New Zealand, ex Trouessart coll., 1894. (Neumann
coll. 710. Original, G. H. F. N. del.)

1 The scutum of I. intermedins Neumann, 1899, p. 132, measures 1-5x1 mm., that of
I. praecoxalis Neumann, 1899, p. 121 ; 1902, p. 119, measures 1-2 x 1 mm.

^ The postero-lateral borders are more rectilinear than is shown in the figure.

/. nemnanni

219

white hairs ; punctations inconspicuous. Venter : very finely striate,
without punctations, but with nunaerous short, white hairs ; vulva
facing the second intercoxal space. Anal grooves semicircular in front,
with sides reaching the posterior border and approaching each other
somewhat midway. Spiracles large, transversely oval. Capitulum:
(0"8 mm. I.) base twice as broad as long, with parallel sides and straight
posterior border ; destitute of cornua ; porose areas pear-shaped, trans-
verse, the broader end internal, interval less than their narrower
diameter, well-marked blunt auriculae, visible dorsally. Palps of
medium length, broad, outer border straight, inner border roughly
parallel with it but with an indentation at the junction of articles 2
and 3. Article 2 longer than article 3. Hypostome long, very slightly
spatulate, with large corona behind which there is a broad unarmed
median area ; dentition 4 1 4 in front to 2 | 2 behind, the outermost
teeth (10 per tile) much the strongest^; this and the next row extend
nearly the whole length of the hypostome ; the two internal rows short
and progressively smaller. Legs medium, coxa I bidentate, the inner
spur broad and curved ; trochanter I without dorsal projection but
with very slight ventral spur; coxae II-IV trenchant and with slight
postero-external spur ; trochanters 2 and 3 slightly spurred ventrally
at their distal extremities. Tarsi fairly long, tapering abruptly ; claws
and pads small.

Nymph : resembles the $ , but for the absence of the vulva and
porose areas, and the smaller spiracle (Fig. 214).

Fig. 214. I. nemnanni o: capitulum and scutum; parts of venter, spiracle and tarsus 4.
Same origin as preceding. (Original, G. H. F. N. del.)
1 Neumann (1902, p. 119) gives the dentition of I. praecoxalis as 4 14, that of I.
intermedins as 2 | 2.

15—2

220

Genus Ixodes

Our description is based on 1 $ and 1 O, part of a lot of 7 ?s and
12 o s, collected in New Zealand by Suter and presented to Neumann by
Trouessart in 1894.

Neumann's specimen of I. intermedius (1 ?) was collected by
Dr Schauinsland from Phalacrocorax sp., and /. praecoxalis (2 $ s, 1 o)
from jtEstrelata cooki in New Zealand.

32. IXODES PERCAVATUS Neumann, 1906.

Figs. 215, 216.
Lit. and Icon. : Neumann, 1906, pp. 200, 201, Figs. 4, 5 (reproduced).

Male : unknown.

Female : Body oblong, lateral borders parallel, reddish brown,
6x3 mm. Scutum oval-lozenge-shaped, broadest slightly in front of
the middle of its length, 1-6 x 1'3 mm. ; cervical grooves very evident,
attaining the posterior border; no lateral grooves; punctations very
fine, numerous only peripherally, where there are some short hairs.
Dorsum with fairly numerous short hairs ; traces of a marginal groove.
Venter- with fewer hairs. Vulva between coxae III. Anal grooves long,
parallel, shortly ogival in front. Spiracles whitish, subcircular, slightly
elongate transversely. Capitulum short (shorter than in the figure);
base trapezoid, broader behind, with slight cornua ; porose areas very
large ^ almost occupying the whole width, tangential in the median line ;

Fig. 215.

Fig. 216.

Figs. 215, 216. I. percavatus ? : capitulum and scutum ; forepart of venter. (Neumann,

1906, Figs. 4 and 5.) 1

1 N.B. Neumann's figures are slightly inaccurate in two respects : the porose areas
occupy a greater proportion of the basis capituli, leaving only a narrow band in front ;
and the spur on trochanter 4 is smaller (not larger) than that on trochanter 3.

/. percavatus var. rothschilcU 221

auricula forming a stout retrograde process. Digit ? Hypostome
narrow, rounded distally, 4 | 4, teeth equal. Palps short, article 2 barely
longer than 8, article 1 triangular ventrally, with an anterior point
internally and a posterior point. Legs of medium length, dark red-
brown. Coxae I-III bear spurs at the postero-internal angle, the spurs
diminishing in size and absent on coxa IV ; coxa I bears an external
spur which is slightly longer than broad, short, pointed, almost trans-
verse ; a shorter spur at the postero-external angle of coxae II-IV.
A 7'etrograde ventral spur on the distal border of trochanters 1-4 (see
Fig. 216). Tarsi of medium length tapering obliquely, those of pair
1 long and slender.

Description based on four dried gorged $s, slightly mutilated, from
Nightingale Island, of the Tristan d'Acunha group (W. of the Cape of
Good Hope ; Brit. Mus.). Species allied to Ixodes neumanni Nuttall
and Warburton.

Ixodes percavatus var. rothschildi n. var. Nuttall
and Warburton.

Male : unknown.

Female : differs from the type in the following respects : Cornua
strong, and recurved dorsally ; porose areas smaller, the interval equal to
half their width ; no retrograde spur on trochanter 4.

Nymph : resembling the $ in all respects, except sexual charac-
teristics.

Described from (N. 634) 2 $s and 3 Os, taken from a puffin (ex
Rothschild coll., no locality recorded).

33. IXODES PILOSUS Koch, 1844.

Figs. 217-221 (original).

Lit. and Icon. : Koch, 1844, p. 233; 1847, p. 105, PI. XXI, Fig. 79 ( ? )
Neumann, 1899, p. 151 ; 1901, p. 288.
Loimsbury, 1900a, p. 47; 1900c, p. 32; 1904, p. 29.
Mally, 1904, p. 7.
Howard, C. W., 1908, p. 94.
Donitz, 1910, p. 435, PI. XVI b. Fig. 6 (? : capitulum, ventral aspect; good).

The " Paralysis Tick " or " Russet Tick " of Cape Colony.

Male (Fig. 217): long-oval, 2*8 x 16 mm. Scutum oval, leaving a
narrow marginal fold ; cervical grooves shallow, parallel at first, then

222

Genus Ixodes

diverging ; no lateral grooves ; numerous unequal punctations, especially
evident in the median field. Emargination moderate, scapular angles
blunt. Venter : genital orifice between coxae III ; pregenital plate
twice as long as broad : median plate long, its postero -lateral borders
very oblique ; anus very near the posterior border ; anal plate almost
pear-shaped, rounded in front, narrowing behind ; adanal plates with
their oblique anterior border much longer than their posterior border.
Spiracles very large, long-oval, Gapituliim : base trapezoid dorsally,
broader anteriorly, finely punctate, produced to a median retrograde
point ventrally ; palps fairly long and broad dorsally, the outer contour
straight, the inner convex ; hypostome strongly armed for half its length
with 4 I 4 teeth, about 5 per file, terminating basally with 4 strong teeth,
2 on either side, the external the stronger. Legs strong, coxa I with
short trenchant internal spur, coxa IV with a slight indication of an
external spur ; coxae II and III without spurs, but with posterior border
very trenchant

Fig. 217. I. pilosus i : dorsum and venter, spiracle and tarsus 4. The <? to the left
(N. 927) is unfed, that to the right (N. 1111) is replete. Specimens from Cape Colony,
received (N. 927) from C. P. Lounsbury, and (N. 1111) from A. H. N. Fillers.
(Original, G. H. F. N. del.)

Female (Figs. 218, 219) : Body red-brown, with numerous rather
long, white hairs. Scutum longer than broad, broadest at its anterior
third, cordiform ; lateral grooves straight, meeting the lateral borders
just behind the rounded lateral angles ; cervical grooves fairly distinct

/. 2^i^osus

223

for about two-thirds of the scutal length, and often perceptible to the
posterior margin, at first converging, then diverging gradually ; emargina-
tion slight, scapular angles short but fairly sharp ; punctations nunnerous,
discrete, fairly deep, of moderate size. Venter : vulva between coxae IV ;
genital grooves nearly straight, slightly diverging ; anus very posterior ;
anal groove short, horseshoe-shaped ; spiracle fairly large, transverse-
oval. Capitulum (Fig. 219): base sub-triangular, much broader than long,
with slightly concave posterior border and with sides parallel for a short
distance ; cornua absent ; porose areas elongate-pear-shaped, the broader
ends internal, the interval fairly broad ; fairly distinct auricular ridges
ventrally. Palps long, the external contour noticeably concave, article 2
half as long again as article 3 which terminates bluntly. Hypostome as
long as the palps, 2 [ 2, but a short additional internal row of teeth (3 per
file) on either side in the anterior portion ; median unarmed area fairly
large, especially posteriorly. Legs fairly strong, coxae rather elongate,
coxa I with very slight internal spur, coxae II-IV without spurs, but
with trenchant posterior borders, especially coxae II and III ; tarsi
tapering gradually, the terminal much longer than the basal segment.

Fig. 218, 1. piloms ? (unfed) : dorsal aspect of body, venter, spiracle, tarsus 4.
(N. 927. Original, G. H. F. N. del.)

224

Genus Ixodes

Fig. 219. I. pilosus ? : capitulum in ventral and dorsal aspects. From the same ? as in
Fig. 218. (Original, G. H. F. N. del.)

Nymph (Fig. 220) : resembles the $ , especially iu the horseshoe -
shaped anal groove and a slight external concavity of the palps. The
scutum is somewhat more angular, and the coxae have a fairly distinct
external spur.

Fig. 220. I. pilosus o (unfed) : dorsum, part of venter, tarsus 4, spiracle. Specimen
from Cape Colony, sent by C. P. Lounsbury, 1906. (N. 921. Original, G. H. F. N. del.)

Larva (Fig. 221): the palps are slightly concave externally. The
anal groove is generally obsolete, but when visible it conforms in shape
to that of the $! and o.

See further under Notes on Biology (p. 316).

/. pilosiis

225

Fig. 221. I. pilosus larva (unfed) : venter, dorsum, tarsus 3, digit. Same source as o in
Fig. 220. (N. 919. Original, G. H. F. N. del.)

Origin: Koch's type, a $, came from S, Africa. Neumann (1899)
described the species from 3 cT and 5 % specimens from Cape Colony
(E. Simon coll.), and subsequently reported (1901), after examining
specimens from C. P. Lounsbury's collection, that pilosus is found
abundantly in Cape Colony on cattle, mules Jwrses and pigs. Lounsbury
(1900 a) observed specimens in copula on cattle, horses, goats and hogs,
and (1900 b), records them as occurring at Stellenbosch ; he successfully
reared large numbers of larvae on goats ; he, moreover, states (1904) that
the "russet tick" is common on dogs at Stellenbosch. Mally (1904)
states they occur on sheep in the E. Province, Cape Colony. Howard
(1908) adds man, the cat, leopard and bush-buck to the list of hosts,
and Donitz (1910) records the species from the civet cat at Umtali.
According to Howard and Lounsbury, it occurs in grass districts both
in Cape Colony and the Orange Free State. Mr W. F. Cooper's
collection contains $ s from the duikerbok, Gonubie Park, S. Africa, xii.
1908.

Our drawings have been made from numerous specimens of all stages
kindly presented by Mr C. P. Lounsbury, in 190G (N. 919-923), and we
have received other specimens from Cape Colony through the courtesy
of Messrs E. G. Wheler (N. 924-927) and A. H. Noel Pillers (N. 1111).

Neumann has described a variety of 1. pilosus under the name of
/. pilosus howardi, which appears to us doubtfully good for the following
reasons :

We possess numerous specimens received from Lounsbury (in whose
collection Neumann, 1901, p. 288, found Koch's I. pilosus in abundance),
and these were sent to us as co-types of /. pilosus. On measuring
them it was found that they agreed better with Neumann's description
of the var. Jioiuardi.

Neumann, however (in a letter dated July 27th, 1910), admits that

226 Genus Ixodes

he was mistaken in the length of the </ I. pilosus, which averages
2*8 mm, (and not 3"15 mm, as given in his description), so that the
varietal differences almost disappear. The only specimen he was able
to send us of /, pilosus var, hoiuardi was a microscopic preparation of
the %, which appears to us identical with the specimens of I. pilosus
from Louusbury's collection. His description of the variety is as follows :

Ixodes pilosus var. howardi Nn. 1908.

Lit. : Neumann, 17. vi. 1908, p. 125.

Howard, C. W., 17, vi. 1908, p. 126 ; also viii. 1908, p. 95, PI. IV, Figs, a-k ;
describes $ , $ , larvae, eggs ; figures ^ and ? and details thereof somewhat
sketchily.

Male: resembles the type, but is smaller, 2-3 x 1-2 mm. (type measures
3*15x1 '6 mm.; average length 2'8 mm., see top of this page).

Female : differs from the type as follows : Scutum longer than broad
(1*27 X 1*1 mm.), oval; cervical and lateral grooves less marked; punctations
finer, less deep, more numerous ; hairs less numerous on dorsum and venter.
Anal grooves with branches closer together and forming an open ellipse posteriorly.
Capitulum smaller, 0*7 mm. (instead of 0'85 in type ; porose areas smallei*, shallower,
wider apart. Legs much feebler, especially the coxae. General colour, unfed, orange-
red.

Description based on 2 (J sand 12 $s, found on dogs at Leydsdorp, Transvaal, by
Dr Copland and at Durban, Natal, by Dr Theiler, and sent to Neumann by C. W.
Howard (Pretoria). The latter subsequently took specimens in various parts of the
Transvaal, i.e. in the Marico, Rustenburg, Pretoria, Barberton and Zoutspansberg
Districts, and at Piet Retief and Durban in more or less dry or very arid places.
Howard states that "most of the localities are in the low bush veldt, but those
places on the high veldt (about 4000 feet) where the tick is common, are those
parts which are covered with bush, and the tick is apparently lacking in the high
grass."

The principal host is the dog, but Howard has found specimens on a cat,
Erinaceus frontalis and Rhinolophus sp.

34. IXODES LUNATUS Neumann, 1907.

Figs. 222, 223.
Lit. and Icon. : Neumann, v, 1907, pp, 88-90, Figs. 1 and 2 (here reproduced).

Male : unknown.

Female : L. 2-7, W. 1"8 mm., body oval, at least twice as broad
behind as in front, brownish, broadest towards the posterior third.
Scutum 1'57 X 1-39 mm., narrow, elliptical, maroon-brown, glossy,

/. lunatus

227

glabrous ; cervical grooves faintly marked, extending two-thirds the
length ; lateral grooves indicated by a scarcely visible linear ridge ;
punctations very fine, numerous, widely separated. Dorsum covered
by fairly numerous short hairs ; no trace of a marginal groove.
Venter with similar hairs, vulva between coxae IV ; anal groove horse-
shoe-shaped ; spiracles, sub-circular, brownish, nearer to the vulva than
to the anus. Capitulum long (mutilated in the type, but about 0*94
mm. long); basis capituli triangular, with slight cornua ; porose areas
medium, oval, longer than broad, the interval equal to their length, a
slightly salient ridge running externally to them ; auricula forming a
pointed retrograde process ; digit ? hypostome ? Palps 0"84< mm. long,
narrow, cultriform, article 2 slightly longer than 3. Legs slender, long
(pair 4 about 2*5 mm. long) ; coxa I bears two sharp spurs, divergent,
separated, the inner longer ; a shorter spur at the postero-external angle
of coxae II-III, very short on coxa IV ; tarsi long, tapering obliquely,
pair 1 long and slender ; pad almost as long as the claws.

Fig. 222. Fig. 223.

Figs. 222, 22.3. I. hutcttus ? : dorsum and venter. (Neumanu, 1907, Figs. 1 and 2.)

Description based on 1 ^ from Hallomys audeberti Jent. (Muridae),
Madagascar (Leyden Natural Hist. JVIuseum).

228

Genus Ixodes

35. IXODES RASUS Neumann, 1899.

Figs. 224-226 (Fig. 225 original).
Lit. and Icon. : Neumann, 1899, pp. 137-139, Figs. 12-14 (reproduced); 1901, p. 285.

Male (Fig. 224): Body oval (2-7 x 1-6 mm., Nn.), broader behind,
with posterior border rounded. Scutum convex, reddish brown, marginal
fold narrow (01 mm.), uniform. Cervical grooves faint, at first parallel,
then divergent. Very numerous fine, equal punctations, distributed
over the whole surface. Venter : sexual orifice broad, opposite the second
intercoxal space; pregenital plate quadrangular and finely punctate;
pre-anal plate very large, broad posteriorly, punctate ; anal plate circular,
the anus eccentric, nearer the anterior rim of the circle. Spiracles large
and long-oval, almost twice as long as broad, slightly tapering behind.
Capitulum short (0'55 ram.), slightly longer than broad, broadest just
behind the palps. Hypostome (Fig. 224 C) short and broad, sometimes
slightly indented, dentition 4 | 4 to 3 | 3, the teeth increasing in size
posteriorly, the rows are irregular, and the internal teeth are flat and
blunt, while the external teeth are pointed, the last externals very large.
(Digit; see Fig. 224 B.) Palps short, broadest at junction of articles 2
and 3, which are almost equal in length. Legs long, the last two articles
of pair 4 extending beyond the abdomen. Coxae large, the first three
noticeably glossy, the last punctate. A short, blunt spur at the postero-
internal angle of coxa I and at the postero-external angle of coxa IV.
Coxae II and III unarmed.

Fig. 224. I. rasits <? : A venter ; B digit, x 210 ; C hypostome, x 135.
(Neumann, 1899, Fig. 12.)

/. 7'asus

229

Female (Figs. 225-226) : Butly rather sh(jrt-uval, broadest at the
posterior third. Scutum (I'd x 1*5 mm., Nn.), reddish brown, oval,
glossy, dark brown, with very numerous fine, equal punctations ; cervical
grooves superficial but nearly attaining the posterior border; lateral
grooves barely indicated by a slightly prominent margin. Dorsum with
three posterior longitudinal grooves, and with numerous punctations
studded with short hairs. Venter with similar punctations and hairs.
Vulva between coxae IV ; anal groove circular, with anus eccentric, as
in </. Capitulum long (1'3 mm., Nn.), base an equilateral triangle, with
slight cornua. Porose areas oval, size variable, converging in front,
the interval about equal to their breadth ; auricula forming a stout
retrograde process (variable in size, see Fig. 22.5). Hypostome lanceolate ;
teeth 4 1 4, then 3 | 3, finally 2 | 2, the externals stout and pointed,
the internals blunt ; the inner files far apart. Palps flat, cultriform,
the outer border nearly straight, the inner convex ; article 2 twice
as long as 3. (Chelicera, see Fig. 226 A.) Legs of medium length ;
coxae I and II near together, the coxae somewhat separated. The coxae
resemble those of ^, but the spurs tend to become obsolete in large $s.

Fig. 225. I. rasus ? unfed : dorsum, part of venter, tarsus 4, spiracle. Details (to
right) showing variation (p.a.) in size of porose areas ; a in size and form of auricula;
in coxa I (note spur) and vulval flap or apron, as observed in another specimen out
of the same lot. Collected from mayi and dog, Kumasi, Ashanti, by W. Graham,
X. 1907. (N. 928. Original, G. H.F.N, del.)

230

Genus Ixodes

Fig. 226. I. rasns ? : A digit, x 210 ; B hypostome, x 70 ; C, D tarsi 1 and 4.
(Neumann, 1899, Figs. 13 and 14.)

Tarsi long, tapering fairly gradually ; tarsus 1 nearly as long as articles
4 and 5 together ; pad almost as long as the claws.

Neumann (1899, p. 139, and 1901, p. 285) records specimens from the
Congo (from Hyrax sp., cT ? in copula, Mocquerys coll., now in Neumann
coll.) ; from the Cameroons (Zenker and Conradt coll.) ; from German
East Africa (Kummer and Conradt coll.), one specimen having been
taken from an Ichneumon ; and from Togo (Berlin Mus.).

Neumann, 1907 a, p. 18, reports 1 %, from Kilimandjaro, Kibonoto
(1300-1900 metres), 17. x. 1905; 1 %, from Kilimandjaro, 8. x. 1905;
1 %, from Meru (3000-3500 metres), Regenwald, collected by the
Sjostedt Zoological Expedition, German East Africa.

We have received examples (N. 877 d) from Uganda, taken from
cattle, which came from Bukedi to Mpumu ix. 1909 (Sir D. Bruce) ;
from (N. 503) Obuasi, S. Ashanti, from leopard, and (N. 928) from
Kumasi, C. Ashanti, from man and dog (Dr W. M. Graham, 1907-8).

Our description differs from that of Neumann.

36. IXODES UGANDANUS Neumann, 1906.

Figs. 227, 228.

Lit. and Icon. : Neumann, 1906, pp. 198-200, Figs. 2, 3 (reproduced).

Male (Fig. 227) : Body 2*2 x 1*2 mm., contour oval, narrow in front,
broadest toward the posterior third, maroon-brown. Scutum convex,
glossy, glabrous, marginal fold narrow, about as broad on the sides as

/. uganrlanffs

231

behind ; cervical grooves narrow, superficial ; punctations many, distant,
very fine (iDconspicuous). Venter : genital orifice broad, between
coxae III. Pregenital plate rectangular, longer than broad, emarginate
in front ; median plate pentagonal, large, very broad behind, very finely
punctate ; anal plate almost circular, forming a very short point behind,
coarsely punctate ; adanal plates somewhat longer than broad, with
many punctations. Spiracles large, oval, much longer than broad.
Capitulum short (0"5 mm.), with pentagonal base, broader than long, the
sides diverging in front, no cornua, the postero-ventral border un-
dulating; no auriculae. Hypostome (Fig. 227) short, broad, 2 stout
marginal teeth, the posterior stouter; the other teeth forming saw-like
ridges, as figured. Palps short, articles 2 and 3 of about equal length.
Legs of medium length, dark. Coxae broad, contiguous; a short, blunt
spur at the postero-internal angle of coxa I and at the postero-external
angle of coxa IV. Tarsi of medium length, tapering gradually.

/=^^^

Fig. 227. I. ugandanus i : venter and hypostome. (Neumann, 1906, Fig. 2.)

232 Genus Ixodes

Female (Fig. 228) : Body short oval, 6x4 mm., dark brown.
Scutum oval-lozenge-shaped, longer than broad (1'3 x I'l mm.), broadest
slightly behind the anterior third, dark brown, glossy, slightly emar-
ginate^ in front, very fine punctations, slight lateral grooves^ cervical
grooves very superficial and extending beyond half the scutal length.
Dorsum with many clearly visible punctations, few scattered hairs,
tegumentary folds very marked and undulating ; traces of three posterior
grooves. Venter bears similar punctations, hairs and folds ; vulva small,
between coxae IV ; anal groove encircling the anus, converging behind
and uniting in an elongated point. Spiracles larger than coxa IV,
slightly oval, with long axis transverse to the body axis. Capitulum
narrow, 092 mm. long. Dorsal base rectangular, much broader than
long; very slight cornua; porose areas oval (but variable), oblique,
slightly converging in front, almost tangential to the posterior border,
the interval equal to their width ; no auriculae. Hypostome long,
narrow, lanceolate, with 2 (? 3) marginal rows of teeth on each side,
leaving the middle portion free. Palps long, slender, article 2 at least
twice as long as 3. Legs slender, relatively short, maroon-browD.
Coxa I prolonged by an internal spur, which is fine, about three times
as long as broad at the base ; coxae II-III unarmed ; coxae IV with a
very small external tuberosity. Tarsus 4 tapering obliquely, with a
slight swelling at its base.

Fig. 228. I. ugandanus ? : capitulum and scutum. (Neumann, 1906, Fig. 3.)

1 Neumann states that the ? scutum is not emarginate and does not possess lateral
grooves ; that the spiracle is small, and the porose areas oval.

/. cor (lifer 233

Neumann's original description was based on I ^ and 1 % in coitus,
found on Aulacodus sp., from Ugan(1a, collected by E. Degan (British
Mus.). Neumann, 1907 a, p. 18, has since reported 1 cT, 1 $, from
Kilimandjaro, Kibonoto, Regenwald (2000-3000 metres), collected by
the Sjostedt Zoological Expedition, German East Africa. We have
received specimens of this tick (N. 1215, $s), taken from a large rodent,
Oshogbo, Eastern Province of S. Nigeria, W. Africa, 28. ii. 1910
(J. J. Simpson coll., for Entomol. Res. Committee); (N. 1216,? o),
taken from sheep, Ibesha, W. Africa, 7. II. 1910 (J. J. Simpson).

We have modified Neumann's description of the $, in the light of
the numerous specimens we have received.

Ixodes ugandanus subsp. djaronensis Neumann, 1907.

Lit. : Neumann, 1907 a, p. 18 (no figure).

Male : Body 1*7 mm. I. Capitulum 0"3 mm. I., auriculae almost
obsolete ; digit 90 /x I., with external article strongly bidentate and
having two minute distal teeth.

Female: Body attaining 4*6 mm, (capitulum included) x 3'3 mm.
Colour varying from yellowish white to maroon-brown. Scutum
0"95 X 0*68 mm. Capitulum 0*6 mm. I., auriculae slight.

The foregoing is abstracted from the author's description. The sub-
species is stated to differ from the type in being smaller, and having
slight auriculae.

Described from 1 (/* and 10 $s, found on Genetta suadelica Mtsch.,
23. VII. 1905, Kilimandjaro, Kibonoto, and 1 $,from Meru, Regenwald
(3500 metres), collected by the Sjostedt Zoological Expedition, German
East Africa. We possess (N, 1222) 1 $, found on Procavia brucei,
Harar, Abyssinia, 1901, collected by Dr H. Brumpt, which we refer to
this subspecies.

37. IXODES CORDIFER Neumann, 1908.

Fig. 229.

Lit. and Icon. : Neumann, vii. 1908, p. 73, Fig. 1 (reproduced).

Male : Body oval, twice as broad behind as in front ; 35x2 mm.
(width at spiracles). Scutum brownish yellow, glossy, slightly convex,
covering the whole dorsum ; cervical grooves very short and very

N. I. 16

234

Genus Ixodes

shallow ; punctations fine, numerous, finer and less numerous longi-
tudinally over the whole median third, the punctations absent in two
symmetrical, longitudinal and sinuous depressions occupying the anterior
two-thirds, where they are limited outwardly by a linear prominence
indicating a lateral groove. Venter concave ; hairs few and very short ;
genital orifice broad, between coxae II ; pregenital plate short, rather
broader than long ; median plate quadrangular, narrow in front, much
longer (1-7 mm.) than broad (1*1 mm. posteriorly) ; anal plate cordi-
form, as broad as long, with anterior border straight, sides convex
and forming a point behind, with fine punctations ; adanal plates twice
as broad at the posterior border as in front \ finely punctate and fused
behind the point of the anal plate ; spiracles large, oval, much longer
than broad. Gapitulum yellowish, short (0*6 mm.), with pentagonal
dorsal base, as broad as long ; lateral borders slightly diverging in front,
ending at the insertion of the palps ; posterior borders straight, without
salient angles; no auriculae; hypostome short, broad, 2| 2, with rounded
teeth. Palps short, broad ; article 1 transverse ; articles 2 and 3
apparently fused, constricted at the base and very broad dorsally along
most of the length. Legs dirty yellow, long, stout; coxae broad.

Fig. 229. /. cordifer s : venter and tarsus 4. (Neumann, 1908, Fig. 1.)
1 Neumann's figure, reproduced here, appears to be inaccurate in this respect.

/. holocf/dns

235

contiguous, with four long hairs at the posterior border ; a spur at the
postero-external angle, increasing in size from I-IV, where it is about
thrice as long as broad ; a short flat spur at the postero-internal angle
on coxae I-III, decreasing in size backward. Tarsi long, tapering
abruptly (step-wise) ; claws long, pad almost as long.

Female : unknown.

From description by Neumann, based on 1 cT from Sekroe, New
Guinea, collected by S. Scliadler (Leyden Mus. Nat. Hist.). In general
form this species approaches /. holocyclus (which is Australian) rather
than /. rasus (which is African).

38. IXODES HOLOCYCLUS Neumann, 1899.

Figs. 230-232.

Lit. and Icon. : Neumann, 1899, pp. 151-155, Figs. 24-26 (reproduced).

Male (Fig. 230): Body oval, broader behind, posterior border
rounded, 3x2 mm., reddish yellow all over. Scutum convex, marginal
fold narrow ; cervical grooves short, continuous in front with the lateral
grooves which attain the posterior border. No punctations nor hairs.
Venter : sexual orifice broad, facing the second intercoxal space. Pre-
genital plate hexagonal, one-third broader than long. Anal plate longer
than broad ; anal grooves at first straight and parallel, then curved and
converging backward without uniting at the posterior border. Median
plate very long, very broad behind, covered with short hairs. Adanal
plates in the form of oblique parallelograms. Spiracles almost twice

Fig. 230. I. holocyclus c? : 1, ventral aspect ; 2, hypostome, x 75.
(Neumann, 1899, Fig. 24.)

16-2

236 Grenus Ixodes

as long as broad, tapering much behind, whitish. Capitulum short
(0*6 mm.), its base quadrangular, scarcely broader than long. (Digit, see
footnoted) Hypostome (Fig. 230) short and broad, with some blunt
denticles in front, then 2 | 2 files of 5-6 blunt teeth. Palps as in $ but
shorter, scarcely twice as long as broad. Legs with coxae contiguous,
with spur slightly stouter than in $ ; legs thicker, with shorter articles,
more enlarged distally.

Female (Fig. 231, unfed): Body flat, oval, yellowish, 3*2 x 1-7
mm., a complete marginal groove limiting a marginal fold of equal width
throughout. When fully gorged, the body is oval, broader behind, con-
tracted on a line with the spiracles, the largest specimen 15x11 mm.,
and 9 mm. thick. Integument finely striated, with short hairs, few and
scattered. Scutum reddish yellow, almost oval, slightly longer (2"8 mm.)
than broad (2 mm.), flat in the median field, the sides convex ; cervical
grooves convergent, short and superficial; lateral grooves clearly marked,
attaining the posterior border, accentuated by an external ridge ;
punctations very numerous, subequal, confluent in places. Dorsum :
grooves normal. Venter : vulva behind the level of coxae III (variable).
Sexual grooves very divergent, then curving toward each other at
the level of the anus, ending on the posterior border. Anus rather
posterior, in an elongate depression ; anal grooves parallel, then
convergent behind, where they unite in a point on the median line.
Spiracles very small, circular, reddish, placed toward the middle of the
length, distant from coxae IV. Capitulum long (1*3 mm.), reddish yellow ;
base short, pentagonal, broader than long, posterior border fairly straight,
lateral borders sub-parallel, then convergent in a cone anteriorly ; porose
areas deep, oval, elongate in the direction of the axis ; auricula forming
a retrograde horn. (Digit, see footnote 2.) Hypostome (Fig. 231) long,
narrow, lanceolate, numerous small sharp teeth, followed by 3 | 3 files of
12-13 stout teeth, increasing^ in size from within outward in each trans-
verse series, the median files far apart. Palps much longer than hypo-
stome ; article 1 thick, salient outward near the base ; article 2 at least
twice as long as 3, the whole palp cultriform, broadened dorsally, concave

1 Digit 160 fi long; dorsal process (?) ; external article 4-toothed, the two anterior teeth
very small, the others strong.

2 Digit 210 fi long; dorsal process a longitudinal crescent, with retrograde teeth,
prolonged down to the lower third of the digit by its shaft of insertion ; external article
elongate, with five teeth of increasing size.

^ Neumann states they diminish outwardly, but this does not agree with the figures
nor with our specimens.

/. holocj/clus

237

outside : article 4 very small, subterminal. Legs reddish yellow, long.
Coxae large, trapezoid, separated by intervals almost equal to their
width, a postero-external spur decreasing in size from coxae I to IV;
long white hairs on the posterior coxal border. Similar hairs on the
ventral border of the other articles. Tarsus 1 as in /. ornithorhynclii (see
Fig. 237) ; tarsus 4 almost four times as long as broad, abruptly tapering,
the pseudo-articulation nearer the proximal extremity of the article ;
claws short, the pad almost reaching their tips.

Fig. 231. I. holocycbts ? (partially gorged) : 1, capitulum and scutum in dorsal aspect ;
2, venter ; 3, bypostome, x 75. (Neumann, 1899, Fig. 25.)

Nymph (Fig. 232) : differs from $ as follows : Body oval, 2-6 x
] "3 mm. (broadest on a line with the spiracles). Scutum 1 x 0'7 mm.
Capitulum 0'88 mm. long (dorsally) ; base more conical. (Digit, see
footnoted) Hypostome narrower, sharper, with 2 [ 2 files of 13-14 teeth,
the external stout, pointed, the internal blunt Palps without apparent
articulation separating articles 2 and 3. Legs 4 extending beyond
the posterior border of the body to at least the length of the tarsi ;
coxal spurs feebler.

Description based on 2 $ s and 1 (/ collected from Australian sheep ;
1 % from Phascogale penicillata (Marsupial Tree-shrew) at Port
Stephens, Australia ; 1 O from Sciurus variabilis in India (British
Museum) ; 1 gorged $ from a dog in Queensland (Bur. Animal
Industry, Wash,); 2 gorged ^s of unknown origin (Hamburg Museum);
1 unfed ? from N. S. Wales, and 1 o from E. India (?) (Berlin
Museum) ; 1 microscopic specimen of a O from a red squirrel from

1 Digit 115 fx long ; dorsal process a subterminal book at tbe end of a narrow sbaft.

238

Genus Ixodes

Fig. 232. I. holocyclus o : dorsal aspect of capitulum, x 88 /^ Z.
(Neumann, 1899, Fig. 26.)

India (R. Blauchard coll.). Neumann (1901, p. 288) states it has
been found on calves in N. S. Wales (Bur. of Agr., N.S.W.). We have
received (N. 644) 2 $s from W. Australia, vii. 1900, from Macropus
sp., also (N. 643) 2 os from W. Australia which we attribute to this
species (ex Rothschild coll.). W. F. Cooper's collection contains
specimens ( $ ) from Lismore, Tweedline, Queensland, x.1907 (S. R. Scott)
and (t/", $ ) received from the Dept. of Agriculture, Brisbane, Queensland,

39. IXODES SCHILLINGSI Neumann, 1901.

Figs. 233 (Neumann) and 234, 235 (original).

Lit. and Icon. : Neumann, 1901, pp. 288, 289 (? , no figure).
Neumann, 1910, pp. 170-172, Fig. 10 (cJ, reproduced).

Male (Fig. 233) : Body oval, 2-3 x 1-38 mm., broadest in front of the
spiracles about midway along body-length. Scutum maroon-brown,
slightly glossy, convex, glabrous, marginal fold bearing very short hairs ;
cervical grooves doubtful ; very many large, subequal punctations,
absent only on three small areas situate on the posterior third ; two
symmetrical punctate depressions midway along the length ; deeply
emarginate ; scapulae prominent. Venter flat, very short equidistant
hairs ; sexual orifice facing the second intercoxal space ; median plate
pentagonal, narrowing in front, longer than broad (0'89 x 7'2 mm.);
anal plate small, racket-shaped, with quadrangular prolongation behind

/. scliillingsi

239

adanal plates with straight anterior border, the convex external and
concave internal borders uniting in a point behind ; all the plates bear
shallow medium punctations. Spiracles large, oval, orientated lengthways
to body. Capitulum dark, short (0"5 mm. I.) ; dorsal base pentagonal,
as broad as long, lateral borders slightly diverging in front, posterior
border straight, no cornua ; auriculae pointed, with median point arising
from ventral ridge (tricuspid appearance) ; digit 135 //, I. (see Fig. 233).
Hypostome short, broad, with six transverse rows of 6-8 rounded teeth
and a large basal tootii on each side equal in length to the others
collectively. Palps short, broad dorsally ; article 2 broader than long,
barely as long as 3. Legs maroon-brown, pair 4 very long, the last two
articles extending beyond the posterior border of the body. Coxae with
trenchant posterior border; coxa I triangular, with very small spur
postero-internally ; 11 and III subquadrangular, likewise with small spur
as on I ; IV longer than broad, with small postero-external spur. Tarsi
long, tapering abruptly (see Fig. 233) ; pad almost as long as claws.

Fig. 233. I. scliillingsi 3 : venter, left digit in ventral aspect, tarsus 4.
(Neumann, 1910, Fig. 10.)

Female (Fig. 234) : Body relatively short, narrowed in front, much
dilated toward the middle of its length in front of the spiracles, rounded
posteriorly ; L. 4'3 mm., W. 3"5 mm., yellowish white. Scutum reddish
brown, much longer than broad (1'5 x 11 mm,), broadest toward the

240

Genus Ixodes

anterior third ; emargination slight ; cervical grooves superficial at first,
then fairly deep and diverging slightly to the middle of the postero-
lateral border ; no lateral grooves, punctations very numerous, deep,
equal. Dorsum bearing a few scattered hairs, finely striated. Venter
with hairs more numerous, similar striations. Vulva facing coxae IV ;
sexual grooves nearly straight, slightly divergent, then slightly conver-
gent posterior to the anus. Anus situated far posteriorly ; anal plate
forming an ellipse, disappearing posteriorly in a short prolongation.
Spiracles subcircular, whitish, very posterior, far from coxae IV.
Capitulum 09 mm. long; base an equilateral triangle; porose areas
subtriangular, broadened transversely, large, shallow, the interval wide ;
auricula forming a sharp retrograde hook. Hypostome long, narrow,
lanceolate, pointed anteriorly, with pointed teeth, numerous and small
in front, then 3 j 3, followed by 2 | 2 files of 12-13 teeth extending almost
to the base, the internal files much separated, especially posteriorly.
Palps long, narrow, widest in the middle of article 2, scarcely extending
beyond the hypostome, much separated at their base, then curving
inward ; article 3 elongate, nearly as long as article 2. Legs of medium
length. Coxa I with a very sharp, narrow, internal spur and short
strong external spur ; II and III trenchant on their posterior border ;
the trenchant edge ending abruptly externally; coxa IV with a slight

Fig. 234. I. schilUngsi ? : capituham and scutum, capitulum in ventral aspect with
coxae, tarsus 4, spiracle, slightly magnified view of venter, s indicating position of
spiracle. Drawn from (N. 1171) co-type ; presented by the Berlin Museum.
(Original, G. H. F. N. del.)

/. schillmf/si

241

indication of a spur at the postero-extenial angle. A very slight tooth,
ventrally, at the distal extremity of the trochanters of each leg. Tarsi
long, especially those of pairs 1 and 4, tapering fairly abruptly near to
their extremities ; pad almost as long as the claws.

Nymph (Fig. 235) : resembles the $ but the scutum is broader ;
the punctations on the scutum, though deep, are not so numerous,
and there is a slight external spur on all the coxae.

Fig. 235. I. scMllingsi o: capitulum and scutum, ventral aspect of (mutilated) capitulum
with coxae, anal groove, spiracle and tarsus 4. Specimen from the same source as
preceding. (Original, G. H. F. N. del.)

Our description of the % and o is based on (N. 1171) 1 % and 2 os
presented by the Berlin Museum from among Neumann's types (12 $s,
etc.) from Colobus caudatus, a monkey, from German East Africa
(Schillings coll.). According to Schillings, as stated by Neumann, this
tick fixes itself exclusively on the eyes, especially in young monkeys,
one eye being often lost. Neumann, 1907 a, p. 18, reports 2 $s and
1 o, found on Colobus caudatus, Kilimandjaro, Kibonoto, 3. vii. 1905,
and 1 $ found in the same locality on grass, 12. vii. 1905, collected
by the Sjostedt Zoological Expedition, German East Africa. Neumann
(1910, p. 171) has recently described the ^f from a specimen found in
copula, collected by G. Vasse in 1907, in Mozambique, Province of
Gorongoza (Paris Mus.) ; the description we give of the cT is condensed
from Neumann, who states that the 2 $s collected with the ^ were
somewhat larger than the types (7 ram. I., including capitulum).

242 Genus Ixodes

40. IXODES ORNITHORHYNCHI Lucas, 1845.

Figs. 236-238 (Fig. 236 original).

Lit, and Icon. : Lucas, 1845, p. 58, PI. I, Figs. 3, 3 a, 3 b, $ dorsum and venter.
Pagenstecher, 1861, p. 40, PL II, Fig. 25, hypostome and digit (nymphs).
Neumann, 1899, pp. 142-144, Figs. 18, 19 (reproduced).

Male : unknown.

Female (Fig. 236, gorged) : Body long and narrow, with lateral
borders almost straight and parallel, attaining 7x4 mm., body, capitu-
lum and legs earthy yellow. Scutum considerably shorter than broad
(1"2 X 19 mm.), only slightly emarginate ; antero- and postero-lateral
borders very divergent, slightly convex, posterior angle very broad and
rounded ; cervical grooves shallow, divergent, broad, disappearing an-
teriorly ; very many very fine equal punctations ; no lateral grooves.
Integument very finely wrinkled, covered by fine short hairs. Dorsum :
a complete marginal groove only visible in the unfed tick close to the
border (Fig. 236). Venter : vulva narrow, facing the second intercoxal
space ; sexual grooves very long, sinuous, only slightly divergent,
attaining the posterior border; anus situated far behind, anal grooves
slightly or not divergent. Spiracles large, almost circular, milky white.
Capitulu7n short, with pentagonal or subtriangular base at least twice
as broad as long; porose areas very large, rounded. (Digit, see Fig. 238 C)
Hypostome (Fig, 238 H) short, bearing 3 | 3 files of 7-8 teeth. Palps
thick, with base distant from the hypostome ; article 2 about twice as
long as 3. Legs long, grouped in the anterior third of the body in
gorged specimens. Coxae elongate, unarmed, punctate, flat, trenchant
at the posterior border ; legs long, relatively narrow, haired. Tarsus
1 (Fig. 237) thrice as long as broad, cylindrical, slightly humped near
the tip, with obsolete pseudo-articulation ; tarsus 4 four times as long
as broad ; the basal pseudo-article broader than the distal, which tapers
dorsally. Claws short, pad two-thirds the length of the claws.

Nym.'ph (Ixodes ot'nithorhynchi Fa.gensteQ,her) : resembles $. Body
relatively broader (3 x 1'5 mm.), oval, the colour varying from brownish
yellow to earthy yellow. Gapitulum with base relatively shorter and
broader, without porose areas. Digit 85 yu. long, as in $ . Hypostome
with 2 I 2 files of 5 teeth.

Neumann was the first to describe the species adequately from
(a) six ^s, from the Paris Museum, of which five were from the
Orriithorhynchus (Lucsis coll.), and one from the Marianne Islands, collected

/. ornithorhyncM

243

Fig. 236. I. ornithorhynchi ? : to left, an unfed ¥ , dorsum, venter and spiracle. Type
in Paris Mus. To right, a replete ? , dorsum and coxae (the latter not to scale),
from specimen in Megnin coll. ; body 5-5 mm. I. (Coll. Lab. de Parasitol., Paris.
Original, G. H. F. N. del.)

Fig. 238.

Fig. 237. /. ornithorhynchi ? : tarsi 1 and 4, x 45. (Neumann, 1899, Fig. 19.)
Fig. 238. I. ornithorhynchi ? : C, left digit viewed dorsally to one side, 150 fj. I. ; H, bypo-
stome, x75. (Neumann, 1899, Fig. 18.)

by A. Marche. (Lucas states that this species attaches itself especially
to the sides of the belly near the anal and genital regions, and also on
the back); (b) 30 os, collected by Jules Verreaux in Tasmania in 1843
(Paris Museum). (Pagenstecher records several from Ornithorhynclms ;
he took them for females) ; (c) three $ s, from the Oniithorhynchiis
(Megnin coll.). Neumann (1901, p. 285) states that this tick has also
been found on the legs of Ornithorhynchus (coll. Bureau of Agriculture,
N. S. Wales). Our figures are drawn from Lucas' types and specimens
in the Megnin collection (now in the Lab. de Parasitol., Paris).

244 Genus Ixodes

41. IXODES TASMANI Neumann, 1899.

Figs. 239 (Neumann) and 240 (original).
Lit. and Icon. : Neumann, 1899, pp. 144, 145, Fig. 20 (reproduced).

Male : unknown.

Female (unfed): Body flat, oval, reddish yellow, the scutum, capi-
tulum and legs being darker; 3-2 x 1-6 mm. (greatest breadth on a line
with coxae III). Dorsum with complete marginal groove, limiting a
marginal fold of uniform width. When gorged, the body is ovoid,
swollen dorsally, almost as thick as broad, broader behind, 10 x 7 mm.,
yellowish brown, glabrous, or bearing short hairs. Integument trans-
versely striated, covered by small, regularly disposed varrucosities.
Scutum^ slightly broader (1'5 mm.) than long, the antero-lateral borders
divergent, subrectilinear, with rounded lateral angles, and short, wavy
postero-lateral borders soon disappearing in the broadly rounded posterior
border ; cervical grooves shallow, divergent ; many fine and equal
punctations ; no lateral grooves. The dorsum shows traces of the usual
grooves. Venter: vulva narrow, facing the antero-internal angle of
coxae III ; sexual grooves divergent, very long, extending slightly
over on to the dorsum. Anus toward the posterior third ; anal grooves
parallel, then approaching each other slightly behind, where they also
extend over on to the dorsum. Spiracles whitish, transversely oval.
Capitulum short (750 fi), base twice as broad as long, hexagonal (due,
apparently, to the fusion of the enlarged basal article of the palp with
the basis capituli) ; porose areas large, oval, close together. Digit ?
Hypostome (Fig. 239 B) markedly spatulate, with broad corona, bearing
3 1 3 files of 9-10 teeth, of about equal size in each transverse row,
decreasing in size from in front backward, and continued by 4-5 rows
of blunt and salient teeth. Palps flattened, much separated and narrow
at their base, very broad at their distal half; the articulations between
articles 2-3 obsolete, article 1 forming a curious structure surrounding
the hypostome. Legs of medium length, grouped, in gorged specimens,
on the anterior quarter of the body, the tips of the fourth pair scarcely
extend beyond the spiracles, the movable articles being cylindrical.
Coxae trenchant ; tarsi almost identical with those of /. ornithorkynchi
(Fig. 237). Claws short ; pad almost reaching their free extremities.

^ There is a noticeable variation in the shape of the scutum in the types. Those
collected by Verreaux in 1843 (Paris Mus.) have a scutum very much as figured by
Neumann (see Fig. 239^).

1. tasmanl

245

Neumann's description was based on one unfed and two gorged $s,
collected by Verreaux, the ornithologist (1847), in Tasmania; one unfed
and one gorged $ from the Island of St Pierre (Ponafidin ?) (Paris
Mus.), and one gorged $, of unknown origin (Berlin Mus.). Hosts
probably birds.

We have somewhat altered the description given by Neumann,
after having examined the types, one of which we figure.

tf"-S'''v^Sr^^^\'

Fig. 239. /. tasmani ?: /J, capitulum and scutum ; B, hypoatome, x 105 :
C, tarsus 1, x 70. (Neumann, 1899, Fig. 20.)

Fig. 240. I. tasmani ? (type, unfed) : dorsum and venter, spiracle. Verreaux coll., 1847;
a pinned specimen. (Paris Mus. Original, G. H. F. N. del.)

246 Genus Ixodes

42. IXODES TENUIROSTRIS Neumann, 1901.

PI. V, Figs. 1, 2, original. Figs. 241-244 (242-244 from
Neumann, 241 original).

Lit., etc. : Ixodes tenuirostris Neumann, 1901, p. 286, Figs. 5, 6 ( $ , reproduced).
Ixodes tenuirostris Neumann, 1902, p. 119, Fig. b {^ , reproduced).
Ixodes tenuirostris in Wheler, 1890, p. 40; 1906, p. 414, Figs. 23, 24 ($ capituluin,

(^ dorsum, photographed).
? Ixodes trianguliceps Birula, 1895. See Notes on Doubtful Species (p. 293).

Male (PI. V, Fig. 1 ; Figs. 241, 242) : Body regularly oval, small,
reddish yellow all over. Scutum : 1*5 x 0*8 mm., glossy, convex, mar-
ginal fold narrow ; very slightly emarginate ; cervical grooves very
faint, shallow, divergent ; no lateral grooves ; some very short sparse
hairs on the posterior border; punctations extremely minute and
inconspicuous. Venter: sexual orifice large, facing the posterior borders
of coxae II ; median plate scarcely longer than broad ; anal plate very
narrow anteriorly, very broad posteriorly, almost triangular, with sides
very diveigent and subrectilinear ; adanal plates trapezoid, considerably
broader in front than behind ; a few inconspicuous short hairs. Spiracle
rather large, subcircular, with its posterior border rather straight.
Capitulum short (0-33 mm. Nn.), with base slightly broader in front
than behind, shagreened, with lateral and posterior margins ridge-like.
Hypostome short and broad, some denticles distally, followed by 2 | 2
files of 3 to 4 short stout teeth succeeded by squamiform teeth. Palps
short, thick, convex dorsally, much separated at the base, with long
axes converging in front. Legs : coxae rather short and broad, without
spurs, but with posterior borders somewhat trenchant; tarsus 4 rather
long, tapering gradually.

Fig. 241. Fig. 242.

Fig. 241. I. tenuirostris s : capitulum in dorsal aspect. From the type at the British

Museum. (Original, sketch by G. H. F. N.)
Fig. 242. I. tenuirostris <? : posterior part of venter. (Neumann, 1902, Fig. 5.)

TICKS

PLATE V

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Figs. 1, 2. Ixodes tenuirostris.

Fig. 1. (f venter. (Origiual, E. G. Wheler, phot.)

Fig. 2. ? venter, mounted specimen, xll. (Original, E. G. Wheler, phot.)

Figs. 3, 4. Ixodes putus.
Fig. 3. s dorsum, x 7. (Wheler, 1906, Fig. 26.)
Fig. 4. ? dorsum, x 7. (Wheler, 1900, Fig. 25.)

(All specinieus of British origin.)

1

/. tenuirostf'is

247

Female (PI, V, Fig. 2; Figs. 243, 244): Body lead-coloured,
elongate-oval, may attain 8x5 mm. in fully gorged specimens. Scutum:
0'8 X 07 mm., broadest near the posterior end ; antero-lateral borders
subrectilinear, divergent, united by a broadly rounded posterior border ;
emargination slight ; surfiice uniform, glossy, without cervical grooves,
punctations inconspicuous ; lateral grooves straight, parallel to the
borders, very fine. Dorsum covered by fine hairs, short, scattered ; in
the posterior third a horseshoe-shaped groove open posteriorly, and in
the space which it circumscribes a median groove extending more
posteriorly. Venter with hairs somewhat more abundant. Vulva large,
facing coxae III. Sexual and anal grooves very divergent, the latter
ogival in front of the anus; spiracles whitish, rather obliquely oval,
narrowed antero-dorsally. Capitulum small (0'5 mm. Nn.); base (+ article
1 of palps which are fused with it) subtriangular dorsally, twice as broad
as long, the sides rounded, the posterior border somewhat concave, porose
areas round, covering the greater part of the dorsal surface, the interval
equal to about half their diameter. Hypostome narrow, with 2 | 2 files
of 7-8 blunt teeth. Palps narrow, the first article in the form of a stout
spur, directed transversely outward and fused with the base, article 2
very narrow at its base and widening gradually, twice as long as 3.
Legs weak, short. Coxae as in the cT ; tarsi slender, tapering' gradually ;
pad almost as long as claws.

Fig. 243.

Fig. 244.

Fig. 243. I. tenuirostris ? : capitulum and scutum (too long). (Neumann, 1901, Fig. 5.)
Fig. 244. I. tenuirostris ? : capitulum and coxae I. (Neumann, 1901, Fig. 6.)

248 Genus Ixodes

Nymph : resembles the $ in its essential characters, except that
the spur on article 1 of the palp is more salient ventrally than laterally.

Larva : resembles the o, but the scutum is comparatively shorter
and broader.

This species appears particularly to infest small Mammalia, Neumann
first recorded the tick (three $s, two os) from Evotomys glareolus,
Island of Riigen (Lemm coll., Berlin Mus.) and (one $ ) from " Arvicola
pratends" Painswick, Gloucestershire, England (E. G. Wheler's coll.,
1893). The cT, first described by Neumann, was found on Evotomys
glareolus, Swansea, Wales (type, C. Oldham coll., British Mus.). Evans
(1907, p. 36) found the o on Arvicola amphibius var. ater in Kincardine-
shire, Scotland (October). We have received specimens from the
following hosts and localities :

Great Britain : (N, 623, $ ) from Microtus agrestris (field vole),
Tring, Herts.^ (N. 1165, $ s), from field vole, Longner Hall, Shrewsbury,
VIII. 1910 (R. F. L. Burton). (N. 622, $), from Microtus amphihius
(water vole), England, iv. 1892^. (N. 620, $) from Mus minutus
(harvest mouse); no particulars^ (N. 356, larvae), from Sorex minutus
(pigmy shrew), Grippetts, Gloucestershire, i, 1908 (E. A. Wilson).
(N. 621, os), from Sorex vulgaris (common shrew), Bevendean, Sussex,

VI. 1899\

Switzerland : (N. 624, 625, 1069, a ^ captured with mouthparts
buried iu $ 's body close to anus, $s and o s) from Microtus arvalis,
Campfer, St Moritz, Engadine, vii. 1904 (K. Jordan)^. (N. 619, ^ and
$), from Evotomys glareolus (bank vole), Campfer, St Moritz, Engadine,

VII. 1904 (K. Jordan) 1. (N. 1161, $ ), from Tarasp, summer of 1901 1.

43. IXODES FECIALIS Warburton and NuttaU, 1909.

Fig. 245.

Lit. and Icon. : Warburton and Nuttall, 1909, pp. 58, 59, Figs. 1 and 2

(reproduced).

Male : unknown.

Female (gorged) : Body long, cylindrical, 8x4 mm., dark brown, with
yellow scutum, capitulum and legs. Scutum- rather longer than broad
(1*5 X 1*3 mm.), broadest near the posterior end ; uniformly punctate
except along the cervical grooves, which are otherwise obsolete ; the

1 Ex N. C. Eothschild coll.

2 Recalling a conventional form of heraldic shield, hence /eciaZ/s.

/. fecialis

249

sides straight and diverging posteriorly ; lateral grooves indicated by
a prominent ridge of darker colour, sparsely punctate and parallel with
the lateral border; scapulae blunt; emargination moderate. Dorsum
minutely punctate, and thickly studded, especially in the posterior
portion, with very short white hairs. Gapitulum : base, a triangle,
longer than broad, with wing-like lateral projections hollowed dorso-
laterally to receive the palps ; porose areas long-oval, rather near
together, very close to the posterior border, which is somewhat sinuous ;
palps medium, flat, the second article twice as long as the third ; hypo-
stome truncate, with parallel sides, 2|2 equidistant files of stout teeth, 9
per file. Venter : vulva between coxae IV ; spiracle rather small, trans-
verse oval, somewhat pointed dorsally, far from coxae IV ; anal groove
long, ill-defined in front but apparently somewhat ogival ; the sides
nearly parallel. Ze^s yellow, slender. Coxae unarmed but with posterior
border somewhat blade-like ; coxa I with strong anterior projection em-
bracing the basis capituli collar-wise, its posterior edge straight ; coxae
II-IV with posterior edge increasingly curved, coxa IV being sub-
circular. Tarsi tapering gradually.

Described from (N. 650) one gorged ? from Dasyurus geoffroyi,
Cranbrook, W. Australia, ill. 1900, presented by the Hon. N. C.
Rothschild.

Fig. 245 A.

Fig. 245 B.

Fig. 246.

Fig. 245. I. fecialis 9 ; A, capitulum and scutum; B, ventral aspect of capitulum with
coxae I. (Warburton and Nuttall, 1909, Figs. 1, 2. Sketch by C. W.)

Fig. 246. /. fecialis, var. aegrifossus ? : capitulum and scutum. (Warburton and
Nuttall, 1909, Fig. 3. Sketch by C. W.)
N. I. 17

250 Genus Ixodes

Ixodes fecialis var. aegrifossus Warburton and
Nuttall, 1909.

Fig. 246.

Lit. and Icon. : Warburton and Nuttall, 1909, pp. 59, 60, Fig. 3 (reproduced).

Male : unknown.

Female (Fig. 246) : differs from the type in the following respects :

Scutum narrower (1"4 x 1 mm.), more pointed posteriorly, with punc-
tations almost obsolete. Gapitulum : base with the median triangular
region distinctly marked off by ridges from the lateral regions ; porose
areas hardly visible {aegrifossus) ; hypostome ?

Described from (N. 339) three mutilated $s taken from an opossum,
Tamberine Mountain, Logan river, S.E. Queensland, iv. 1907, and
labelled " Scrub-ticks," sent by Professor R. T. Hewlett (London), who
received them from the Bacteriological Institute, Brisbane. We also
possess (N. 1213) a ? found on Perameles obesula, at Bannister,
W. Australia, 21. viii. 1900 (ex N. C. Rothschild coll.).

N.B. The porose areas were at first believed to be absent, the median
area of the basis capituli appearing to be uniformly finely granular in
texture. They were distinctly though faintly apparent, however, in a
microscopic preparation.

These ticks possess a type of scutum, broad posteriorly, which
appears to be somewhat characteristic of Australian forms. It is
observable in 7. australiensis, I. ornithorhynchi and /. tasmani, and the
scutum of the nymph of /. vestitus is of this shape, though, in the adult
female, it tends to become more circular. I. tenuirosb^is alone of extra-
Australian species conforms to this type. The species here described
most clearly resembles /. vestitus in general characteristics.

44. IXODES AUSTRALIENSIS Neumann, 1904.

Figs. 247-249 (Figs. 247, 248, original).
Lit. and Icon. : Neumann, 1904, pp. 456, 457, Fig. 1 (reproduced).

Male : unknown.

Female : Body a short oval, usually broadest on a level with
coxae IV, more or less dark brown; L. 2*5 to 325 mm., W. 19 to
2*7 mm. Scutum hexagonal, with rounded angles, shorter than broad

/. atistraliensis

251

(1 X 1-35 mm.), glossy; cervical grooves superficial, reaching the posterior
border; no lateral grooves; punctations fine, numerous; longitudinal
parallel striae mixed with the punctations, well marked especially in
the lateral fields. Dorsum uniform, very finely punctate, glabrous, with
a complete marginal groove. Venter smooth ; vulva broad, between
coxae III. Aiial grooves closed posteriorly; oval, tangential to the

Fig. 247. I. australiensjs ? : eapituluni and scutum, capitulum in ventral aspect.
Specimen found on Bettongia lesueuri, Kogonup, W. Australia, 1900. (N. 646.
Original, G. H. F. N. del.)

Fig. 248.

Fig. 249.

Fig. 248. I. australiensis ? : part of venter, tarsus 4, spiracle. (The same specimen as

in Fig. 247. Scale = 1 mm. Original, G. H. F. N. del.)
Fig. 249. /. anstraliensis ?: capitulum in ventral aspect, coxae I. Obviously drawn

from a mounted specimen. (Neumann, 1904, Fig. 1.)

17—2

252 Genus Ixodes

anal ring anteriorly, forming a short point at their union posteriorly.
Spiracles small, circular, whitish, placed somewhat posteriorly to the
middle of the length. Capitulum very short (540 /x), with very short
sub-rectangular base, at least twice as broad as long on the dorsal
surface, as long as wide on the ventral surface ; auricula a small
tuberosity; excavate on the anterior border; porose areas medium,
somewhat triangular, separated by an interval equal to their breadth ;
in front of them a triangular median pit. Hypostome spatulate,
dentition 3 | 3 or 4 | 4. Palps much separated at the base, claviform ;
article 1 large, hollowed out on the inner side to surround the chelicerae
and hypostome like a cuff; the second article convex externally at the
base, its articulation with the third but slightly visible. Legs of
medium length. Coxae flattened, not winged, increasing in size from
I to IV, all bearing a sharp spur at the postero-external angle, internal
angle unarmed. Coxa I visible dorsally. Tarsi long, tapering gradually;
pads half as long as claws.

The species was described by Neumann from five $s, from Canis sp.,
West Australia (N. C. Rothschild coll.). The specimens we possess are
doubtless the types, for they were derived from the Hon. N. C.
Rothschild's collection. Our specimens, two $ s (N. 645, 646), were
found on Canis familiaris, at Cranbrook, IV. 1900 (J. T. Young), and
on Bettongia lesueuri (marsupial rat-kangaroo), at Kogonup, XI. 1900.
Our figures are drawn from the latter.

45. IXODES VESTITUS Neumann, 1908.

Figs. 250-253 (Figs. 251-253 original).

Lit. and Icon. : Neumann, iv. 1908, p. 7, Fig. 2 (reproduced), Fig. 3 (part
of venter ; omitted).

Male : unknown.

Female (Figs. 250, 251) : Body oblong, with lateral borders parallel,
reddish yellow, L. 37 mm. (capitulum included), W. 22 mm. Scutum
cordiform^ with rounded sides, broadest (r35 mm.) toward the middle,
r25 mm. long; cervical grooves absent, replaced by two elongated
depressions (Fig. 250) separated from the anterior border by a distance

1 The antero-lateral borders of the scutum in the type are very ill-defined. They
appeared to us almost rectlliuear (as in the accompanying figures of the nymph and
larva) but Neumann's figure, here reproduced, coincides with the appearance of a specimen
in our possession.

1. vestitus

253

equal to their length, and midway between the centre and the lateral
border ; lateral grooves very fine, without external raised area ; puncta-
tions fine, unequal, fewer in the median field, abundant external to the
lateral grooves ; surface glabrous, glossy. Dorsum covered by stiff
whitish hairs, which are absent just behind the scutum and in three
longitudinal grooves; of these the laterals extend along the whole body-
length ; the median groove is shorter. Venter covered by similar hairs
posterior to the vulva and coxae ; no hairs in the genital grooves and
over the greater part of the space bounded by the anal grooves; vulva
facing the second intercoxal space ; anal grooves parallel, wide apart,
relatively short, curving in front of the anus; spiracles whitish,
subcircular. Gapitulum short (O'T mm.); base (+ article 1 of palps
which are fused with it) twice as broad as long, without cornua ;
porose areas deep, oval, considerably broader than long, almost con-
tiguous; no distinct auriculae. Hypostome narrow, with parallel
borders, 2 | 2, 7-8 stout teeth per file, the median files much separated ;
chelicerae (?) ; palps relatively short ayd broad, article 1 very broad,

Fig. 250. /. vestihis ?: capitulum and scutum. (Neumann, 1908, Fig. 2.)

254

Genus Ixodes

almost discoidal, salient outwardly, fused with the base ; article 2
constricted basally, then broadening abruptly, almost twice as long as
article 3 ; article 4 ver}^ small, terminal. Legs'^ very short and thick,
coxae unarmed, flat, transversely elongated, with posterior border
trenchant ; coxa I prolonged forward, the anterior portion slightly
visible dorsally ; tarsi short, humped near the tips, which taper some-
what abruptly.

Fig. 251. I. vestitus ? : capitulum iu ventral aspect, palp (lateral view), coxae, anal
grooves, spiracle. Leg 4 (less magnified). Found on Diemenia superciliosa (brown
snake), Herdman's Lake, near Perth, W. Australia. B. H. Woodward coll. (N. 642.
Original, G.H. F. N. del.)

Nymph (Fig. 252) : resembles the $ except in the shape of the
scutum, which is of the I.fecialis type, being broadest near its posterior
extremity. The dorsum is thickly clothed with strong white hairs.

1 Neumann notes that leg 3, and especially coxa III, on the right side are atrophied in
the type ; the coxa is only half as large as the one of the opposite side.

/. vestitus

255

Fig. 252. I. vestitus o : dorsum (the capitulum being slightly foreshortened) ; palp (side
view), part of venter, anal grooves, capitulum (dorsal aspect), spiracle and leg 4 : same
origin as the ? in Fig. 251. (N. 642. Original, G. H. F. N. del.)

Fig. 253. I. vestitus larva : dorsum, venter, leg 3. Same origin as the ? in Fig. 251.
(N. 642. Original, G. H. F. N. del.)

Larva (Fig. 253) : resembles the o. Four longitudinal rows of
white hairs on the dorsum.

Description based on one $ from Myr"mecohiusfasciatusWa.terhouae
(Dasyuridae), West Australia (Neumann's type, British Museum). Our
collection contains (N. 64-2) $ , O and larvae from Diemenia superciliosa
(a snake), Herdman's Lake, near Perth, W. Australia (B. H. Woodward,
ex N. C. Rothschild coll.).

256 Genus Ixodes

46. IXODES PUTUS (Pickard-Cambridge, 1878).

Plate V, Figs. 3 and 4 ; Text-figs. 254-260 (Figs. 254,
255, 260, original).

Lit. etc. : Hyalomma puta Pickard-Cambridge, 1878, p. 222, PI. XIII, Fig. 4 (o).
We are indebted to the author for sending us one of his types to examine
in VII. 1910. It proved to be an unfed o, bleached pale yellow by long
immersion in alcohol. His figures, which are very sketchy, represent the
dorsum, venter, and dorsal aspect of the o's capitulum. Without having
seen the type it would have been impossible to refer it to the species
described as puhis by Neumann.

Ixodes borealis Kramer and C. J. Neuman, 1883, p. 518, PI.

Ixodes fimhriatus Kramer' and C. J. Neuman, 1883, p. 518. Cited by L. G.
Neumann, 1899, p. 127, as a good species. We are, however, convinced,
from the authors' description, that Jimbriatus=putus (J. (It measured
4 X 3"5 mm.)

Ixodes putus (Cambridge) in Neumann, 1899, pp. 125-127, Figs. 7-9 (reproduced),
description of ?; 1901, p. 283.

Ceratixodes put\(s (Cambridge) in Neumann, 1902, p. 117, Fig. 4 (capitulum of ^ ,
dorsal and ventral aspects, palp in profile, somewhat inaccurate, consequently
not reproduced. Lahille, 1905, p. 26, reprints descrijjtion of (J, $ and
o given by Neumann ; on pp. 138-148, more detailed description (original),
gives many measurements, etc., PI. XIII, coloured Figs. 1-13 of § and o
with details.

Ceratixodes putus Cambridge, in Wheler, 1906, pp. 415, 416, Figs. 25 ( $ ) and
26 ((?) photographs (reproduced).

Ixodes borealis Kramer and Neuman, in Evans, 1906, pp. 85, 86, Fig. of 5
(reproduced).

1 Ixodes uriae White, 1852, p. ccx, found on U^-ia troile, Baffin's Bay (P. C.
Sutherland).

1 1xodes eudyptidis Maskell, 1885 (see Notes on Doubtful Species, p. 291).

Male (PI. V, Fig. 8 ; Text-fig. 254) : very large (3-7 mm. L,
including capitulum), larger than the unfed %, narrow in front, the
lateral borders diverging for about one-third of the length, then parallel,
the posterior border only slightly convex. Scutum (33 x 2'1 mm.),
broadly oval, very convex, with postero-lateral borders slightly concave,
marginal fold moderate ; cervical grooves fairly long, shallow, divergent,
terminating in a large shallow depression on either side, behind which
two smaller circular pits are often visible ; no lateral grooves ; very
numerous unequal punctations, and very short white hairs, generally
caducent, except on the posterior portion of the scutum. Emargination
very deep ; scapular angles very prominent, but blunt. Dorsum :

1. putus

257

beyond the marginal fold are seen dorsal ly five distinct posterior ridges
(in reality borne by the ventral plates) bearing tufts of strong, equal,
white hairs or bristles, the external ridges being twice the length of the
three median. Venter : Sexual orifice facing the first intercoxal space ;
pregenital plate semilunar, median plate elongate, rather narrow, anal
plate rounded in front, its sides sub-parallel, delimiting the median
tuft of hairs; adanal plates with very convex external border; the anal
and adanal plates deeply punctate. Spiracle very large, very near
coxa IV, subcircular. Capitidum very small ; base twice as broad as
long, broader in front, the posterior border straigiit, the anterior border
excavate ; palps horn-like and curved upwards, tapering to their
extremity, article 1 conspicuous dorsally, article 2 beset with stiff
bristles, longer than article 3, the articulation between articles 2 and 3
very obscure ; article 4 projects ventrally from the base of article 3.
Hypostome very rudimentary, short, bifid, with faint indications of six
teeth, the anterior more external. Legs : coxae contiguous, unarmed,
very convex ventrally; all the articles except the tarsi of legs 1, 2
and 3 extremely stout and strong ; leg 4 conspicuously thinner and
somewhat longer. All the tarsi weak and tapering gradually ; tarsi 2-4
with small terminal spur ; pad one-third as long as claws.

Fig. 254. I. putus i : dorsum, venter and side view ; H, hypostome, spiracle. The apron
or flap over the sexual orifice is indicated by the letter F in the middle figure. From
specimen (N. 968) found on cliffs at Rhosilly, Gower, Glamorganshire, 1909, by
Dr J. W. W. Stephens. (Original, G. H. F. N. del.)

258

Genus Ixodes

Female (PL V, Fig. 4 ; Text-figs. 255-259) : L. 3-3 mm., including
capitulum ; when gorged may attain 11x7 mm. Body light grey when
alive, thickly clothed with white hairs. Scutum : (1"6 x 1*1 mm.) dull
yellow-brown, a few short hairs or hairless, much longer than broad,
broadest near its anterior end ; cervical grooves well marked, long,
reaching the posterior border ; no lateral grooves ; numerous moderate,
deep punctations ; emargination almost absent ; scapulae often some-
what rugose. Venter: very hairy in young specimens: the hairs are
absent in the region corresponding to the anal plate of the male, and
are caducent in the anterior portion in older specimens. Vulva facing
the second intercoxal space ; genital grooves sub-parallel for half their
length, then slightly divergent ; anal grooves slightly divergent; spiracle
(Fig. 259) nearly circular. Capitulum (0"3 mm. I.), base much broader
than long, slightly broader in front ; porose areas oval or semilunar, the
interval very narrow ; palps far apart at their base, article 1 conspicuous
dorsally, article 3 broader than article 2, giving the palps a clavate
appearance (articulations 2, 3, indistinct). Digit (see Fig. 258).
Hypostome strong, well covered with 2 | 2 strong equal teeth, about
8 per file. Legs : long and more slender than in the (^, leg 4 not
conspicuously differing from the others; coxae unarmed, coxae I, II
and III rather elongate and triangular, coxa IV rounded ; tarsi
distinctly humped ; pad short.

Fig. 255

Fig. 256.

Fig. 255. I. putus ¥ : dorsum, venter, right palp in profile, spiracle and tarsus 4. Same
origin as the d in Fig. 254. (N. 968. Original, G. H. F. N. del.)

Fig. 256. I. putus ¥ : dorsum, x 12. (From Evans, 1906, p. 85; described as I. borealis
Kramer and Neuman, 1883.)

/. pnUis

259

Fig. 257. I. putus ? : capitulum in dorsal and ventral aspects. (Neumann, 1899,
Fig. 8, G. Marx del.)

Fig. 258.

Fig. 259.

Fig. 258. /. ■putus 9 : 1, scutum; 2, left digit; 3, hypostome; 4, tarsus 4. (Neumann,

1899, Fig. 9.)
Fig. 259. /. putus ? : spiracle, G. Marx del. (Neumann, 1899, Fig. 7.)

Nymph (Fig. 260): bears an unmistakable resemblaoce to the $.
L. r25 mm. unfed (type), may attain 3'5 mm. when replete.

Fig. 260. I. putus o: capitulum and scutum, capitulum in ventral aspect, coxae,
spiracle and tarsus 4. Same origin as the <? and ? above figured. (N. 9G8.
Original, G. H. F. N. del.)

260 Genus Ixodes

Our description of /. putus is based upon numerous specimens
(t/*, $ , o) found in Great Britain, one $ from Bering Island and one of
the type (o) from Kerguelen Island.

The species has a remarkably wide geographical distribution. As
yet it has only been found parasitic on marine birds or infesting their
breeding places, where it occasionally attacks man.

England : It has been recorded from the following localities by
E. G. Wheler (1906): os on guillemots; $s on puffins, Morthoe,
N. Devon (Pocock) ; ^/s, $ s, on cliffs frequented during the nesting
season by guilleinots and other sea-birds, at Bempton and Buckton on
the Yorkshire coast (Mr Howett, of York, occasionally found sexes in
copula beneath stones) ; on Pinnacle Rocks, Fame Islands (found by
Mr Wheler, who has kindly presented us with some of these specimens :
N. 935, 936, 937). We have adults (N. 454, c/*, $) found on ledges
frequented by guillemots, Bempton Cliffs, Yorkshire, vi. 1908, by
Mr O. Grabham, who states that the jackdaws feed on the ticks with
avidity. Wales: (N. 968, ^^ % o) found on clitfs, Rhosilly, Gower,
Glamorganshire, ^'guillemots, razorbills and puffins breeding there,"
VI. 1909 (sent us by Dr J. W. W. Stephens). Scotland: (N. 459, $)
from seagull, Handa Island, Sutherland, viii. 1908 (presented by
Dr W. L. H. Duckworth) ; Mr W. Evans, of Edinburgh, has kindly sent
us specimens he collected: (o) from a gannet, Bass, vi. 1910, and {%)
from a puffin. Firth of Forth, vii. 1910. He has himself (1906, p. 85)
recorded the occurrence of the tick ( $ , o) from St Kilda, on young
puffins and fulmars, attached to " the bare skin at the angles of the
mouth below the bill, and in the corners of the eyes"; also on stones
and loose turf frequented by the birds. The bites of these ticks are
feared by the natives. Wheler possessed specimens from the Hebrides.
America: Neumann (1899, p. 127) described specimens ($,o) from
the Islands of St Pierre and Miquelon, Newfoundland coast (Baron
coll.); St Paul's Island, Alaska (G. Marx coll., Smithsonian Institution,
Washington, D.C) ; Cape Horn (E. Simon's coll., now Neumann's).
Neumann (1901, p. 283) records o and larvae found on cormorants and
penguins, in Tierra del Fuego and the Antarctic (Belgian Antarctic
Expedition). Lahille (1905, pp. 26, 138-148) recorded specimens ($)
from Phalacrocorax verrucosus (Cab.) Scl. and Salv,, Navarin Island,
Beagle Channel, Tierra del Fuego, ii. 1900, and ($, o) from penguins
(Spheniscus magellanicus [Forst.] Steph.), Port William, Tussac Island
(L. Valette coll.). The ticks were found attached chiefly to the birds'
heads and necks. Lahille was struck by the absence of (/"s. Asia

/. signafus 261

and Australasia : The specimens ((^ and $ ) described respectively as
finihriatus and horealis by Kramer and Neuman were found on Bering
Island, Bering Sea, by tlie Vega Expedition ; we are indebted to the
U.S. Department of Agriculture for a typical specimen (N. 714, $)of
putus from Bering Island. Neumann (1899, p. 127) records ( $ , o) the
tick from Campbell Island, S. of New Zealand (Filhol coll., Paris
Mus.) and from KiNG Island (Tasmania ?). The types (o s) described
by Pickard-Cambridge (1878, p. 222) were found on Pygosceles taeniatus
and in rock crevices, Kerguelen Island, Indian Ocean, by A. E. Eaton
(Transit of Venus Expedition). We have received a $ taken from
a wild duck (N. 617 ; no particulars, ex N. C. Rothschild coll.).
See further under Notes on Biology (p. 317).

47. IXODES SIGNATUS Birula, 1895.

Figs. 261-264 (original).

Lit., Synon. and Icon. : Ixodes signatios Birula, 1895, pp. 357, 358, PI. I, Figs. 10-13
( 9 capitulum in dorsal aspect, tarsus 1, scutum, coxa I, drawn from a
mounted specimen and consequently difficult to recognize from the figures —
that of the capitulum being especially misleading) : recognizable from
tarsus 1, coxa I and scutum.
Ixodes parvirostris Neumann, 1901, p. 284.
Ixodes eudi/ptidis var. signata Neumann, 1904, p. 451.
Ceratixodes signatus (Birula) Banks, 1908, p. 21, PI. II, Figs. 1-3; $ scutum and

coxae, tarsi 1 and 4, capitulum (diagrammatic only). (Why Ceratixodes 1)
Note : As will be seen by reference to /. neumanni N. and W. (p. 217), there has
been considerable confusion about this species. We have examined the type of Ixodes
parvirostris (Nn. coll.) and of /. eudyptidis var. signata (Hamburg Mus.), and find
them to be identical with a mutilated $ specimen (identified by Banks as signatus)
from Pacific Grove, California (U.S. Dei)t. of Agricultiu'e), and with specimens of our
own (N. 290) from the same locality as the last. Our figures are drawn from the
specimens ( $ , o) in the Hamburg Museum and (L) from California.

Male: unknown.

Female (Figs. 261, 262) : Body (6 x 3"5 mm. when replete), elongate,
yellow (in young specimens). Scutum: 1"7 x r2 nmi., yellow-brown
or maroon, subject to considerable variation both of shape and of
texture, normally long-oval, with rounded sides and somewhat pointed
posterior angle, but sometimes considerably narrower and somewhat
angular laterally ; anterior border ill-defined, scapular angles prominent
and often slightly divergent, so that the scutum is somewhat narrower
immediately behind them ; some large confluent punctations or rugosities

262

Genus Ixodes

in the posterior portion of the lateral fields, the rest of the scutum finely
punctate and sometimes rather rugose ; cervical grooves very distinct
for half the scutal length ; no lateral grooves. Venter : vulva just
behind coxae II ; genital grooves sub-parallel ; anal grooves rounded
in front of the anus, the sides somewhat converging to the posterior
border. Spiracle large, nearly circular. Capitulum: base very short
and broad, its posterior border straight with lateral prominences which
hardly amount to cornua, salient laterally in its ventral aspect ; porose
areas elongate piriform, the broader ends internal ; interval very small ;
palps short; far apart at their base, converging anteriorly, excavate
dorsally (especially article 3), article 2 barely longer than article 3 ;
hypostome narrow, lanceolate, 3 | 3 small teeth, then 2 j 2, about 6 sub-
equal, blunt teeth per file, a large unarmed median area. Legs :
yellowish brown, rather long ; coxae with trenchant posterior borders
and with a short, strong, blunt external spur, decreasing in strength
from coxa I to coxa IV ; a very small ventral spur at the distal end
of trochanters 1 and 2. Tarsi long ; tarsus 1 rather humped, tarsus
4 long, especially its distal article, which tapers obliquely far from its
termination ; pads small.

^K:^:^

Fig. 261.

Fig. 262.

Figs. 261, 262. I. signatus ? : capitulum and scutum, left palp in profile, capitulum in
ventral aspect (mutilated), coxae, spiracle, tarsi 1 and 4. The hypostome (highly
magnified) and venter of a second specimen are shown to the right. (From specimens
in the Hamburg Museum. Original, G. H. F. N. del.)

/. signatus

2(53

Nymph (Fig. 263) : very elongate, closely resembling the $ . The
scutum is comparatively somewhat shorter, the basis capituli has ventral
auricular ridges, and the spiracle is transverse-oval.

Fig. 263. I. signatus o : capitulum and scutum, veutral aspect of capitulum (mutilated),
coxae, spiracle, anal grooves and tarsus 4. Same origin as the ? s in Figs. 261, 262.
(Original, G. H. F. N. del.)

Larva (Fig. 264) : the scutum is angular laterally, with postero-
lateral borders concave. The main characteristics of the % are
recognizable in the shape of the capitulum, the armature of the legs,
and the anal groove.

Fig. 264. I. signatus larva (replete) : capitulum and scutum, capitulum in ventral
aspect (foreshortened to show hypostome), parts of venter, tarsus 3. From Brandt's
cormorant. Pacific Grove, California, 1894, ex V. L. Kellogg collection. (N. 290.
Original, G. H.F.N, del.)

264 Genus Ixodes

Described from numerous specimens (N. 290, $ s, o s, larvae) from
Phalacrocoi'acc penicillatus (Brandt's cormorant), Pacific Grove, California,
1894 (V. L. Kellogg coll.) ; a specimen (N. 1227, $ ) from Unalaska,
Aleutian Islands, kindly presented by Professor A. Birula, of St Peters-
burg ; the specimens in the Hamburg Museum and others from the
Neumann collection were kindly sent to us for study.

The types of Birula ($, o) came from the Island of Unalaska
(Wosnessenski coll., 1847). Neumann's description of /. parvirostris
was based on specimens (eight $s) found on Phalacrocorax pelagicus,
Yezo, Japan (Hamburg Mus.).

48. IXODES UNICAVATUS Neumann, 1908.

Figs. 265-267 (Figs. 266, 267, original).
Lit, and Icon. : Neumann, iv. 1908, p. 1, Fig. I ( ? , reproduced).

Male : unknown.

Female (Fig. 265): Body oblong, blood-red, L. 6-5 mm. (capitulum
included), broadest (3*2 mm.) in front of the middle of the length.
Sciitum oval lozenge-shaped, 15 mm. /., broadest (0"8 mm.) in front
of the middle ; cervical grooves shallow, attaining the sides posterior
to the lateral angles ; no lateral grooves ; many sub-equal and fine
punctations ; surface glabrous, glossy, reddish brown. Dorsum finely
striate, giving it a satiny appearance ; a hemispherical protrusion
opposite the scapular angles of the scutum posterior to coxa I ; very
short, fine and sparse hairs. Venter : integument and hairs as on
dorsum; vulva between the posterior borders of coxae II; genital
grooves commencing behind the vulva, diverging at their posterior
third ; anal grooves long, parallel, very slightly approaching each other
behind, united in a semicircle in front of the anus. Spiracles glossy,
smooth, dark, subcircular, very little broader transversely. Capitulum
short (0"65 mm.); base short, almost thrice as broad as long; a small
dorsal point protruding laterally on each side ; without cornua ; porose
areas fused in a single transverse depression, which covers almost
the whole dorsal surface of the base ; ventral surface with a slight
transverse linear protrusion posterior to each palp ; hypostome slightly
lanceolate, 3 j 3 then 2 | 2, the two median anteriorly placed rows being
composed of small teeth, the outer pairs of rows being composed of
8-10 stout teeth; chelicera (?); palps widely separated basally,

/. unicavatns

265

relatively short and broad, uniformly broad along their length ; article 2
half the length of 3 ; article 4 terminal, clearly visible. Le[/s relatively
long and slender, the intermediate articles whitish distally ; coxa I
without internal spur, with an external spur longer than broad and
an anterior spiniform prolongation ; coxae II-IV with external spur
and anterior prolongation as in coxa I, the anterior spine growing
progressively smaller on coxae II-IV ; tarsi long, tapering obliquely
distally.

Fig. 265. Fig. '-^66.

Fig. 265. I. unicavatus ? : capitulum and scutum. (Neumann, 1908, Fig. 1.)
Fig. 266. I. unicavatus o : capitulum and scutum, capitulum in ventral aspect, coxae,
spiracle, anal grooves and tarsus 4. Drawn from specimen found on gannet, Bass,
Scotland, vi. 1910. (W. Evans coll. Original, G. H. F. N. del.)

^

Fig. 267. /. unicavatus larva : capitulum and scutum, capitulum in ventral aspect,
coxae, anal grooves and tarsus 3. Drawn from specimen found on cormorant, Fidra
Point, Firth of Forth, Scotland, xi. 1906. W. Evans coll. Found with ? and o.
(N. 1132. Original, G. H. F. N. del.)

., , 18

266 Genus Ixodes

Nymph (Fig. 266): similar to $, except for the scutum, which is
more rounded.

Larva (Fig. 267): resembling both the $ and o, but the scutum,
whilst comparatively broad, approximates more closely to that of the % .

Neumann described the species from a $ and a O found on
cormorant (Phalacrocorax carbo L.) from the mouth of the Forth,
Scotland, collected by W. Evans, of Edinburgh, Although he mentions
that the lot included a larva, he does not describe it.

Our collection contains specimens from Great Britain: (N. 652) o,
from shag, Plymouth, IX. 1897, and (N. 667), $ s, o s, from Phalacrocorax
graculus, Cromarty, ill. 1898 (W. Hoy); both lots presented by
Hon. N. C. Rothschild. The collection of Mr W. Evans, of Edinburgh,
contains the type specimens ( $ , o and larvae, examined by us) from
the cormorant, Fidra Point, Firth of Forth, XI. 1906.

Note: In a tube labelled " /. eiidyptidis No, 13" at the British
Museum we discovered three $ specimens agreeing very closely with
Neumann's description and figure of /. unicavatus. The specimens
were determined as /. eudyptidis by Neumann in 1905; only one of
the specimens possesses a hypostome. The ticks measure 8x5 mm.,
and were collected from a cormorant by J. E. Harting (locality un-
recorded). The specimens differ from the type as follows : cervical
grooves well-marked ; the hemispherical protrusion opposite the scapular
angles is not visible in all the specimens ; the porose areas show a
slight indication of division in one or two specimens; the palpal
articles 2 and 3 are apparently of equal length, but the separation is
indistinct. In all other respects the specimens agree with Neumann's
description and figure, so that we refer them to /. unicavatus.

49. IXODES LORICATUS Neumann, 1899.

Figs. 268, 269.

Lit. and Icon, : Neumann, 1899, pp. 139-142, Figs. 15-17 (reproduced).
Neumann, 1901, p. 285.

Lahille, 1905, p. 29 (reprints description by Neumann, 1899).
Ixodes angustus Neumann, 1901, p. 284 (see I. angusttis Neumann, 1899, on
p. 195 of this work) ; Neumann, 1910, p. 30.

Male (Fig. 268) : Body oval, narrow, broader and rounded behind,
46 X 23 mm. or 3-7 x 1-8 mm. of the same colour as the unfed %.

/. loricatus

267

Scutum narrow, barely covering more than half the width of the dorsum,
bearing on the pseudoscutum more clearly marked cervical grooves,
but similar lateral grooves and similar punctations to those of $ ; over
the rest of the dorsum the punctations are coarser, distant. A deep
marginal groove ; marginal fold glabrous. Venter : plates highly
chitinised, well defined, maroon ; sexual orifice facing the anterior border
of coxae III ; pregenital plate rectangular, twice as long as broad, with
rounded angles ; median plate elongate, relatively narrow ; anal and
adanal plates elongate, not attaining the posterior border. Spiracles
large, oval, maroon. Scattered hairs over the whole surface, shorter on
the plates. Capitulum 0*7 mm. long; dorsal base slightly broader than
long, similar to that of $ (Fig. 269), the porose areas replaced by a not
clearly defined roughened surface, the lateral points but slightly marked.
Chelicera ? Hypostome and palps similar to those of $ . Legs similar
to those of $ , the coxal spurs slightly weaker.

Female (unfed): Body oblong, sides parallel, 3'8 x 19 mm,, rounded
behind, yellowish white, scutum, capitulum and legs brown. Scutum
oval, extending beyond the middle of the back, 1"8 x 1'2 mm. (emargina-
tion almost absent), glossy, convex, maroon, glabrous ; cervical gi'ooves
slightly marked ; lateral grooves distinct, straight ; punctations many,
equal, very fine. Dorsum with a deep marginal gi'oove, which limits a
thick prominent fold ; numerous ruddy hairs. Venter : vulva facing
coxae III ; sexual grooves broad, straight, divergent ; anal ring maroon,
anus posterior ; anal grooves slightly convergent behind ; spiracles
large, slightly elongate in the ventro-dorsal direction, whitish peri-

Fig. 268.

Fig. 269.

Fig. 268.
Fig. 269.

I. loricatus <? : dorsal and ventral aspects. (Neumann, 1899, Figs. 1.5, 16.)
I. loricatus ? : capitulum, dorsal aspect. (Neumann, 1899, Fig. 17.)

18—2

268 Genus Ixodes

pherally, maroon centrally, placed in front of the posterior third of the
body-length ; hairs longer. Gapitulum (Fig. 269) 1 mm. long, dorsal base
broader than long, divided into three zones by lateral ridges which con-
verge in front; a median trapezoid zone almost completely occupied by the
contiguous porose areas ; two lateral triangular zones with salient angle.
(Digit 125 fi long, resembling that of 7. ricinus.) Hypostome lanceolate,
bearing 2 | 2 files of 10-12 teeth, the marginal stout, pointed ; the internal
weak, blunt and separated from the median line, especially behind.
Palps of medium length, dilated at their internal border, especially
at the second article, which is longer than the third. Legs long, stout,
the two last articles of the fourth pair extending beyond the abdomen.
Coxae broad, brownish, hairy ; coxa I bears a flat spur at each of the
posterior angles ; a similar spur at the postero-external angles of the
others. Tarsi long, tapering somewhat abruptly distally ; pad about as
long as the claws.

A gorged female measures 5*5 x 3"2 mm. ; general colour maroon-
brown ; no marginal groove ; hairs whitish ; anus more anterior and
anal grooves longer than double the interval which separates them.
Four other $s measured 10 x 7 mm.; the greatest width at the posterior
third, the anterior part being narrow ; colour dark brown ; dorsum
glabrous, finely punctate, without characteristic depression.

Nymph: Body oblong, narrowed in front, rounded behind, slightly
constricted on a level with the spiracles, may attain 4'5 x 2 mm., yellow
or reddish yellow all over. Scutum narrow, elongate, with borders but
slightly convex, posterior angle slightly ogival ; 0*67 x 0'46 mm., without
punctations ; cervical grooves long, lateral grooves straight, parallel,
attaining the posterior border. Dorsum excavated by five longitudinal
grooves ; two anterior short, three posterior longer, and a median groove
prolonged further backward ; short, scattered hairs. Ventej' but slightly
concave ; sexual grooves widely separated in front, divergent backward ;
anus distant from the posterior border; anal grooves long, slightly
convergent behind ; hairs as on the dorsum. Spiracles small, rounded,
placed about midway along the body-length. Capitulum short (0"3 mm,),
conforms dorsally with that of $ . Legs similar to those of % ; coxae
distant, with smaller spurs.

Larva (hexapod) : similar to o, 1*9 x 0*8 mm.

Neumann's description is based on two % s and one </ from Didelphys
quica from Brazil, and two os from S. Brazil (Goeldi coll., now in
Neumann coll., Toulouse) ; two os and one larva found on Microdidelphys
sorex (?) from Rio Grande do Sul, Brazil (Brit. Mus.) ; five small <^ s,
and four gorged $ s from Buenos Ayres, Argentine (C. Berg coll.).

/. loricattts var. spinosiis

269

Neumann (1901, p. 283) examined a gorged $ from Tierra del
Fuego (0. Berg coll., now in Neumann's coll., Toulouse). Our collection
contains specimens presented by Hon. N. C. Rothschild, as follows :
(N. 640) from Ateles melanochoerus, Tabasco de la Frontera, Mexico ;
(N. 641) from Didelphys aurita, Brazil, xi. 1901 ; (N. 638) </•, $, from
Didelphys sp., Sapucay ; W. F. Cooper's collection contains a ^T from
Estaiicia Cooper, Paraguay, VI. 190G (Foster).

Ixodes loricatus var. spinosus Nuttall, 1910.

Fig. 270.
Lit. and Icon.: Xuttall, xii. 1910, p. 411, Fig. 5 (reproduced).

Male : unknown.

Female : differs from the type as follows :

It is larger and less hairy generally. The scutum (1*6 x 1"25 mm.)
is broader and less punctate, there being about 20 rather coarse
punctations situated chiefly in the posterior half of the median field.
Capitulum broader (07 mm.) and relatively shorter, more massive, the
ventral surface of the base showing a broad U-shaped depression
extending across the base posteriorly, the arms of the U reaching forward
toward the base of the palps. Venter : spiracles larger (0'55 mm. I.)

Fig. 270. I. loricatus var. spi)wsits ? : capitulum and scutum, ventral aspect of
capitulum with coxae, vulva with commencing genital grooves ; hypostome (highly
magnified) ; spiracle, tarsus 4 and anal grooves. (Nuttall, 1910, Fig. 5. G.H. F. N. del.)

270

Genus Ixodes

than coxa IV, and much nearer to the coxa. Legs more massive, coxa
I with larger postero-external spur prolonged in a point overlapping
coxa II, with sharp internal angle ; coxa II angular internally (round in
the type). Tarsi tapering less gradually.

Described from (N. 647) 3 % s, taken from a large opossum, Tabasco
de la Frontera, Mexico, iu the month of May (ex Hon. N. C. Rothschild's
collection).

50. IXODES COXAEFURCATUS Neumann, 189«.

Fig. 271 (original).

Lit. : Neumann, 1899, p. 127 (no figure).

Male : Body elongate, narrow, sides parallel, 3*7 x 1-7 mm. Scutum
glabrous, marginal fold narrow at the sides and broad behind ; puncta-
tions large, distant ; cervical grooves short, superficial ; faint traces of
lateral grooves; scapulae rounded. Venter: plates glabrous, with
punctations as on scutum ; sexual orifice facing the second intercoxal
space; pregenital plate triangular, longer than broad, with summit
rounded; anal plate almost twice as long as broad, with sides arcuate;

Fig. 271. I. coxaefurcatus i -. dorsum, parts of venter, spiracle and tarsus 4. Sketched
from the type in the Berlin Museum. (Original, G. H. F. N. del.)

7. coxaefurcatns 271

adanal plates with their sides carved arid parallel. Spiracles large,
oval, with long axis directed forward and macula eccentric. Gapituliim
with dorsal base broader than long, posterior border convex ; a median
broad, depressed, roughly circular rugose area. Hypostome with sharp
point, dentition 2 | 2, with 7 teeth per file. Palps short and broad.
Legs of medium length ; coxae bearing long hairs ; coxa I with a
short, stout spur at both posterior angles ; coxae II-IV with a similar
flattened spur at the postero-external angle. Tarsi tapering gradually.
Colour : scutum, shields and legs dark maroon-brown, the venter and
the marginal fold light maroon.

Female : unknown.

Our description is based on an examination of the type, a dried,
pinned, and somewhat mutilated and shrunken specimen from a virgin
forest, Sierra Geral, Brazil^ (Hensel coll., Berlin Mus.).

51. IXODES VESPERTILIONIS Koch, 1844.

Figs. 272-282 (Figs. 272, 273, 277, 280, 281 original). See also
PL VI, Figs. 1 and 2 ($s).

Synonymy : Ixodes vespertilionis Koch, 1844, H. 37, Fig. 9 ( $ ) ; 1844, p. 232, listed ;

1847, p. 21.
Ixodes flavipes Koch, 1844, H. 39, Fig. 2 (/ $); 1844, p. 232, listed; 1847, p. 21.
Ixodes troglodytes Schmidt, 1853 {vide Frauenfeld, 1853, p. 57) ( ^ ).
Eschatocephalus gracilipes Frauenfeld, 1853, p. 57, PI. ((J : dorsum, venter, capit-

ulum ; 5 : dorsum and capituhim. Sketchy, but resembhng /. vespertilionis);

1857, PI. II ( ? ).
Haemalastor gracilipes Frauenfeld, 1854, p. 23, PI. Figs. 4, 5 ( $ ).
Sarconissus flavipes Koleuati, 1856, p. 21, 1857, p. 21, PI. I, " ? ," immature

stages. See Neumann, viii. 1910, p. 191.
Sarconissus hispididus Kolenati, 1856, p. 21 ( 9 )•
Sarconissus hrevipes Kolenati, 1856, p. 21 ( ? ).

Sarconissus kochi Kolenati, 1856, p. 21 (?) ; 1860, p. 573, Ph II, Fig. 4 (c^).
Sarconissus flavidus Kolenati, 1856, p. 21(9).
Sarconissus exaratiis Kolenati, 1856, p. 22; 1860, p. 757, PI. I, Fig. 2 (Neumann

det. VIII. 1910, p. 191).
Haemalastor gracilipes (Frauenfeld) Kolenati, 1860, p. 573. Neumann, det. viii.

1910, p. 191.
Eschatocephalus frauenfeldi L. Koch, 1872, p. 26, PL II, Figs. 29-32 ((^); 1877,

p. 150.

1 Neumann erroneously states tliat the specimen came from Siwa, Egypt.

272 Genus Ixodes

Eschatocephahis seidlitzi L. Koch, 1872, p. 26 (cj); 1877, p. 151.

Ixodes longipes Lucas, 1872, p. Ixxiv. (^J).

Ixodes siculifer Megnin, 1880, p. 132 (j^).

Haemcdastor vespertilionis (C. L. Koch) Neumann, 1899, p. 169; see further
under Literature.

Eschatocephalus vespertilionis (Koch) Neumann, 1901, p. 290; 1902, p. 116.

ISarconissus nodulipes Kolenati, 1860, p. 576, PI. I, Fig. 3.

'\ Eschatocephalus flavi pes (Koch) in Bonnet, 1908, p. 325 (probable).
Lit. and Icon. : Besides the foregoing, see :

1854. Frauenfeld, p. 29 ((?).— 1858. Heller, PI. II, Figs. 7-9; PI. Ill,
Fig. 21.— 1877. Murray, p. 195 {Sarconyssus).—\SdQ. Canestrini, p. 504,
PI. XLIII, Fig. 5 ( ? : dorsum, poor).— 1892. Berlese, fasc. LXI, No. 9 ( ? ,
gorged : dorsum, capitulum and scutum, fairly accurate) ; Marx, p. 235 {Sar-
conyssus). — 1897. Rollinat and Trouessart, p. 136. — 1899. Neumann, p. 169,
Figs. 27-34 ( $ : Fig. 27, venter ; Fig. 28, ventral aspect of capitulum (these two
figures not reproduced), hypostome and digit, leg 1, foot ; $ '■ capitulum in
dorsal and ventral aspects, hypostome and digit ; larval digit (all of these repro-
duced)).—1901. Neumann, p. 290.-1902. Neumann, p. 116.-1906. Wheler,
PI. VIII, Fig. 19 « ( 9 : photograph of dorsum of unmounted specimen, repro-
duced).—1909. Blanchard, p. 96, Figs. 107-110 (after Neumann).— 1910.
Donitz, p. 400 {Eschatocephalus discussed).

See further under Eschatocephahis and Haemalastor^ pp. 134, 281.

Male (Figs. 272-276) : Body long-oval, narrower in front, yellow.
Scutum (3*5 X 2*1 mm.), very long and narrow, indistinctly defined, not
covering the whole dorsum (which is concave in the specimen described),
convex along the median line; emargination slight; faint, rapidly
diverging cervical grooves commencing at some distance behind the
anterior border; no lateral grooves in the usual situation. Coarse
punctations forming a single longitudinal median row and two lateral
rows; numerous very fine punctations over the rest of the scutum. Venter
(Fig. 272): genital orifice facing the second intercoxal space; median
plate pentagonal, fairly broad ; anal plate long-oval ; adanal plates very
long ; all the plates with numerous punctations. Spiracles very large,
subcircular. Capitulum (Fig. 273 ; 0'65 mm. I. dorsally) comparatively
small, base about as broad as long, dorsal ridge convex, the dark raised
ridge extending for a short distance along the lateral borders. Palps
clavate, convex dorsally, with numerous long hairs, article 2 about equal
to article 3. Hypostome (Fig. 274 A) ill-defined, practically smooth, but
with indications of a few small scale-like teeth distally. Digit (see Fig.
274). Legs (Figs. 275, 276) of inordinate length, every article, except
the coxa, being remarkably elongate. Coxae unarmed.

/. vespertilionis

273

Fig. 272.

Fig. 27.3.

Fig. 272. I. vespertilionis i : venter. Specimen from Grotto of Alsasua, E. Simon

coll., 1896. (Neumann coll. 760. Original, G. H. F. N. del.)
Fig. 273. I. vespertilionis <? : capitulum. Same specimen as Fig. 272. (Original,

G. H.F.N, del.)

Fig. 274.

Fig. 276.

Fig. 274. I. vespertilionis <? : A, hypostome, x 84 ; B, B', right digit in dorsal and

ventral aspects, x 238. (Neumann, 1899, Fig. 29.)
Fig. 276. I. vespertilionis <? : ambulacrum, x 110. (Neumann, 1899, Fig. 31.)

Fig. 275. I. vespertilionis i : leg 1, x 18. (Neumann, 1899, Fig. 30.)

274

Genus Ixodes

Female (PL VI, Figs. 1 and 2 ; Figs. 277-279 ; gorged) : Scutum
much longer than broad (1-9 x 1-3 mm.), broadest in the middle; no
lateral grooves, cervical grooves only visible in their hinder portion,
where they diverge and meet the postero-lateral borders of the scutum
at its posterior third. Numerous small shallow punctations. Venter :
vulva between coxae III ; anal groove elliptical in front, with parallel

Fig. 277. I. vespertilionis ? : capitulum (mutilated) and scutum, base of capitulum
(ventral), anal grooves, coxae, spiracle, tarsus 4. Found on bat, Banyuls, France,
by Trouessart, 1896. (Neumann coll. 766. Original, G. H.F.N, del.)

Fig. 278. Fig. 279.

Fig. 278. I. vespertilionis ? : A, hypostome, x 85 ; B, left digit in dorsal aspect, x 225.

(Neumann, 1899, Fig. 33.)
Fig. 279. I. vespertilionis ? : capitulum in dorsal and ventral aspects, x 40. Drawn from

mounted specimen. (Neumann, 1899, Fig. 32.)

/. vespertilioiiis

275

sides ; spiracle as in (/, Gapitidum : base shoiter and biDader than in
the (/*, with salient lateral points ; porose areas large and near together ;
palps normal ; hypostome tapering to a point, 4 1 4, merging to 3 [ 3
posteriorly, the teeth are very pointed and the hypostome posterior to
the basal teeth is constricted (see Figs. 278 A and B (digit), 279).
Legs (PL V, Figs. 1 and 2) long and thin, with the same characteristics
as in (/•, but comparatively smaller.

Nymph (Fig. 280) : resembling the % in all essential parts, the
capitulum with blunt auricular protrusions and no latero-dorsal ridge.

Fig. 280. I. vespertilionis o : capitulum and scutum ; ventral aspect of capitulum, with
coxae, spiracle, anal grooves and tarsus 4. (Same origin as ? in Fig. 277. Original,
G. H.F.N, del.)

Larva (Figs. 281, 282) : resembling the o, but with broad scutum ;
lateral angles and capitulum more elongate and possessing no auriculae.

Fig. 281. Fig. 282.

Fig. 281. I. vespertilionis larva : capitulum and scutum ; ventral aspect of capitulum,

with coxae, tarsus 3. Specimen found on Rhinolophus sp. by Trouessart, 189G.

(Neumann coll. 768. Original, G. H. F. N. del.)
Fig. 282. 1. vespertilionis larva : left digit in dorsal aspect, x 590. (Neumann, 1899,

Fig. 34.)

276 Genus Ixodes

Described from 1 c/*, 1 $, 1 o and 1 larva kindly lent by Professor
Neumann.

This species appears to be widely distributed geographically. It has
hitherto only been found on Cheiroptei^a and in their haunts (in caves,
grottoes and the like).

EUROPE: N. Wales: 1 % from a bat, Cefn, 1896. (Sent to
Mr E. G. Wheler for determination by Professor Newstead, of Liverpool,
being the first specimen, apparently, recorded in this country. Com-
municated in a letter from E. G. W. to G. H. F. N., i. 1906.) Ireland:
1 % from Eden Vale Cave, Co. Clare, vii. 1902 (Dublin Nat. Hist. Mus.,
specimen determined by us.) France : One of us (G. H. F. N.) examined
a number of mounted specimens in the Megnin collection through the
kindness of Professor R. Blanchard (Paris). They had mostly been sent
to Megnin by H. Gadeau de Kerville, of Rouen, during ii-v. 1883 ;
$ s and O from Plecotus auritus, grotte de la Briqueterie, 10. ill. ;
OS and larvae from Rhin. ferrum equinum, grotte Cuvette, 10. iii. ;
larvae from Rhin. hipposideros, grotte de Dieppedalle, Seine inferieure,
10. III. ; larvae, no particulars, 25. ii. ; larvae from Rhin. hippo-
sideros, Carrian de la Londe, 14. v. ; all of these were labelled
I. longipes by Megnin, 1 % , labelled " I. acideifer Megn.," from Rhin.
ferrum equinum, Vincennes, 15 x. 1878 (Gilnicki) ; a larva, from
Vesperugo pipistrellus, Rouen, 20. i. 1884, H. Gadeau de Kerville.
Rollinat and Trouessart, 1897, p. 136, record this tick from Rhin. ferrum
equinum (it was attached to the lip), captured in a cave near Banyuls,
by Dr H. Brumpt (det. by Neumann); Neumann, 1899, p. 176, reports
the tick ( $ , O and larva) on bats from the same place (Trouessart coll.)
and o and larva on bats from Bitche. Neumann, 1899, p. 175, records
9 j/'s from caves and grottoes in the following places : Abeille, Ariege ;
Aspradels and Espezel, Pyrenees ; les Baux, near Aries ; Penne, Tarne ;
Serres, Hautes Alpes ; 3 $ s respectively from Bize and Minerve,
Herault, and from Puivert, Aude ; all of these specimens then in the
Simon collection. Ch. Janet found 2 (/"s in a subterranean chalk-pit,
near Beauvais. Spain: 2 ^/'s from Alsasua, Guipuzcoa and Orduiia,
Biscay, are recorded by Neumann, 1899, p. 175 (Simon coll.). Italy:
% found on Rhinolophus ferrum equinum by Canestrini (1890, p. 504 ;
it measured 8 x 5*4 mm., length of fourth leg= 5*5 mm.). Berlese, 1892,
records the tick from the same host. Germany : C. L. Koch's types of
vespertilionis and flavipes were found on Rhin. ferrum equinum.
L. Koch's 2 (^s were found respectively in the Rosenmliller cave,
near Muggendorf, ix. 1869, and in caves near Almas, Frankischer

TICKS

PLATE VI

Fig. 1.

Fig. 2.

Figs. 1, 2. Ixodes vespertilionis.

Fig. 1. Unfed ? , dorsum, x 7 ; with piece of bat's skin still attached to the mouthparts.

(British specimen. Wheler, 1906, Fig. 19 a.)
Fig. 2. Replete ? , dorsum. Specimen from Co. Clare, Ireland, vii. 1902. (Dublin Mus.,

Original, L. E. Eobinson, phot.)

I

/. vespertilioiiis 277

Jura. Austro- Hungary : o s and larvae have been found on Rhin.
ferrum equiniivi and Rhin. euryale, grotto of Karstes, near Trieste, by
A. Poppe (determined by Neumann, 1899, p. 176). Frauenfeld (1853,
p. 57) found specimens in the Adelsberg cave, Carniola, as Schmidt had
done before him. He states (1854, p. 28) that E. gracilipes iiad been
found in various caves in Carniola and at Adelek in Hungary ; he
mentions the cave of Goba dol, and some were collected at the caves of
Skednenza, Mokrizberg. Joseph found specimens in almost all the
habitats of bats in caves in Carniola, clinging to stalactites, but not
attached to the bats {R. ferrum equinum). Finally, Kolenati's numerous
"species" were either found in the habitats of bats or attached to the
fleshy parts of the flanks of R. feri'um equinum, R. hippocrepis, R. euryale,
R. blasius and R. clivosus, in Moravia.

AFRICA : 1 (/ is recorded from Ain-Fezza, near Tlemcen, Algeria,
having been collected by Simon (Neumann, 1899, p. 175). Whilst
visiting Algiers in February, 1910, one of us (G. H. F. N.) saw a living
$ which had been taken from a bat in Dr Sergent's laboratory at the
lustitut Pasteur.

AUSTRALIA: 1 o in our collection (N. 649), from Vesperugo
tricolor, Kingwilliamstown, S. Australia (Hon. N. C. Rothschild coll.).

278 Genus Ixodes

GEOGRAPHICAL DISTRIBUTION OF THE GENUS.

Some species in the appended list appear to possess special hosts,
judging from their having been found thereon several times. Such
hosts are given in the column to the right :

EUROPE.

ricinus : Distribution general.

Special Hosts.

hexagonus : ,, ,,

canisvga: ,, ,,
brunneus: ,, ,,

Birds.

vespertilionis : „ ,,
tenuirostris : Great Britain, Switzerland,

Island of Riigen.

Bats.

Small mammals

acuminatus : Italy.

putus : Great Britain.

Marine birds.

unicavatvs: Great Britain.

Marine birds.

caledonicus : Scotland.

Birds.

AFRICA.

North Africa :

ricinus : Algeria, Tunisia.

hexagonus : Algeria.

vespertilionis: Algeria. Bats.

nigricans: Madeira (perhaps = ?-?cnnts, see text).
Central Africa:

rasus: E. and W.

ugandanus : E. and W.

ugandanus var. djaronensis : Abyssinia.

schillingsi : E.

rubicundus var. limbatus .- Congo Free State.

cavipalpus : Angola, N.W. Rhodesia.
South Africa :

rubicundus : Cape Colony.

pilosus : Cape Colony.

percavatus : Tristan d'Acunha Islands (W. of Cape Colony).
Madagascar :

iunatus.

ASIA.

ricinus : Transcaucasia, Arabia, Japan.

ricinus var. ovatus : Japan.

acutitarsus : India, S. Formosa, Japan.

signatus: Japan, Unalaska (Aleutian Islands). Marine birds.

japonensis : Japan.

simplex : Shanghai (China).

spinicoxalis : Sumatra.

nitens : Christmas Island (Indian Ocean).

putus: Kerguelen's Land (Indian Ocean), Bering Island

(Bering Sea). Marine birds.

Geographical Distribution 279

AUSTRALASIA.

Special Hosts.
cordifcr: New Guinea.

australiensu : Australia.

iH'stitus : ,,

holocijclus : ,,

fecialis : ,,

fecialis var. a('(jrifossus : Australia.

vespertilionis : Australia. Bats.

tasmani : Tasmania, Island of St Pierre.

ornithorhynchi : Tasmania and Marianne Islands. Ornithorhynchus.

putus : Campbell and King's Islands, Marine birds.

neumanni : New Zealand. Birds.

AMERICA.

North America :

putus : Alaska, Newfoundland. Marine birds.

angttstus : Alaska, Canada, United States.

ricintis: Canada, United States.

richms var. ^ccqmlaris : Canada, United States.

marxi : Canada and United States.

tcxanus : Canada and United States.

hexagonus var. cookei : Canada and United States.

hexagonus : United States.

canisuga : ,,

brunneus : „

signatns: ,, Marine birds.

pratti : ,,

sciilptus : ,,

diversifossiis : ,,

dentatus ; ,,

ricinus var. californicus : United States.

ruhidus : Mexico.

bicornis : ,,

loricatus var. spinosus : Mexico.

minor : Guatemala.

fiiscipes : Panama.
South America :

fossulatus : Ecuador.

elegans : Chili.

boliviensis : Bolivia.

fiiscipes: Brazil.

hexagonus : Locality not stated.

coxaefurcatus : Brazil.

loricatus: Brazil, Paraguay, Argentina, Tierra del Fuego.

auritultts : Straits of Magellan.

putus : Tierra del Fuego, Cape Horn and thereabouts,

Antarctic. Marine birds.

280 Genus Ixodes

LIST OF

CONDEMNED AND DOUBTFUL SPECIES
OF IXODES,

INCLUDING THEIR SYNONYMY AND LITERATURE.

The species are ordered according to the genera to which they have at various
times been referred. The list of good species will be found on p. 346. See also
further under the synonymy of Ixodes, p. 133.

Acarus Linnaeus, 1746.

americanus Linn., 1758, p. 615 ; Fabricius, 1794, p. 428 (given as synonym of
Ixodes americanus by Fabricius, 1805, p. 355, and many subsequent
vf v'liQVs) - Arnblyomma aiiuricanum (Linn.) Koch, 1844.
frontalis Panzer, 1795, H. 59, n. 23 = ? /. brunneus Koch (undeterminable).
hirudo Fabricius, 1795, p. 485 (undeterminable).

holsatus „ 1794, p. 428 = 7. holsatus (Fabr.), condemned, see p. 285.
lipsiensis ,, 1794, p. 427 = 7. Hpsiensis (Fabr.) condemned, see p. 286.
pallipes „ 1794, p. 426 = ? 7. brunneus Koch, 1844.
phimbeus Panzer, 1795, H. 90, n. 22 (undeterminable).
reduvius Linnaeus, 1746, p. 479, see p. 143 of this work.
ricinoides de Geer, 1778, p. 98, PI. V, 16 = 7. ricinus (Linn.) $.
ricinus Linnaeus, 1746, p. 480 = 7. ricinus (Linn). $.
tristriatus Panzer, 1795, H. 59, n. 24 = 7. tristriatus (Panzer), condemned,

see p. 290.
tmdaius Fabricius, 1794, p. 427 (undeterminable).
Ceratixodes Neumann, 1902.

putus (Cambridge), 1878, in Neumann, 1902, p. 117 = 7. putus (Pickard-

Cambridge).
putus Cambridge, in Banks, 1908, p. i\ = Ixodes putus (Pickard-Cambridge).
signatus Birula, in Banks, 1908, p. 21 = /. signatus Birula, 1895.
Crotonus Dumeril, 1822.

ricinus Dumeril, 1822, p. 56 = 7. ricinus (Linn.).
Cynorhaestes Hei-mann, 1804.

reduvius Hermann, 1804, p. 65, PI. VII, Fig. 3, as figured by Hermann, shows
festoons, consequently it cannot be an Ixodes. Genus undeterminable.
(Given as synonym of 7. i-icinus by Neumann.)
ricinus Hermann, 1804, p. 65, possibly =7. ricinus (Linn.).
hermanni Risso, 1826, p. 183 = /. ricinus (Linn.).
Dermanissus Kolenati, 1856.

ruMginosus Kolenati, 1857, p. 20, PI. 1 = 7. rubiginosus (Kolenati), in Neumann,
1899. Condemned, see p. 288.

Condemned Species, etc. 281

Eschatocephalus Fmuenfeld, 1853.

a cut i tarsus (Karsch), in Neumann, 1901=/. ucutitarsus (Karsch).

brei'ipes Neumann, 1899, p. 179 {crassipes Joseph, 1882, renamed) = /. brevipes

(Neumann). See Notes on Doubtful Species, p. 291.
crassipes Joseph, 1882, p. 16. See brevipes above.

Jlavipes (Koch) in Bonnet, x. 1908, p. 325 = probably /. vespertilionis Koch.
frauoifeldi L. Koch, 1872, p. 2G, PI. II, Figs. 29-32 {$); 1877, p. 150=/,

vespertilionis Koch, 1844.
gracilipes Frauenfeld, 1853, p. 55. Pl. = /. vespertilionis Koch, 1844.
seidlitzi L. Koch, 1872, p. 26 ((?); 1877, p. 151=/. vespertilionis Koch, 1844.
vespertilionis (Koch) Neumann = /. vespertilionis Koch, 1844.
(See Sarconysstis.)
Haemalastor Koch, 1844.

acutitarsus Karsch, 1880, p. 142 = /. acutitarsus (Karsch), from Japan.

brevipes Neumann, 1899, p. 179 = /. bi-evipes (Neumann), see p. 291, Notes on

Doubtful Species.
crassipes Kolenati, 1857, p. 437 ; 1860, p. 578, PI. II, 5 ; Neumann, 1899, p. 178,

and 1901, p. 290. Found on Rhinolopkus clivosns, in Egypt. Description

insufficient.
crassitarsus Karsch, 1880, p. \^\ = Amblyomma longirostre (Koch), Neumann,

1905, p. 231, not Amblyomma crassitarsus (Karsch) Neumann, 1901, p. 315;

not Hyaloinma crassitarsus (Karsch), as stated in Neumann, 1899, p. 293.

Original description insufficient ; from Caracas.
longirostris Koch, 1844, p. 233; 1847, p. 4Q = Amblyomma longirostre (Koch)

Neumann, 1905, p. 231, not Hyalomma longirostre (Koch) as stated in

Neumann, 1901, jj. 315.
Hyalomma

p%ita Pickard-Cambridge, 1878, p. 222 — I. putus (Pickard-Cambridge).
Ixodes Latreille, 1796.

aca7itkoglossi Lucas, 1878, p. xxxv ; description insufficient. Specimens from

New Guinea, parasitic on Acanthoglossus bruijnii.
aegyptius (Linn.) Fabricius, 1805, p. 353 ; also Gervais, 1844, p. 244 (who gives

Ac. aegyptius Linnaeus and Cyn. aegyptius Hermann as synonyms) =

Hyalomma aegyptium (Linn.).
aegyptius Audouin, 1827, p. 428, wrongly renamed /. savignyi by Gervais,

1844, p. 245. See savignyi= Hyalomma aegyptium (Linn.).
aequalis Banks, 1909. See Notes on Doubtful Species.
afinis Neumann, 1899, p. 120 = Ixodes ricinus var. scapular is (Say) Nuttall and

Warburton, 1910.
africanus Megnin, 1876, p. 831 = Hyalomma aegyptium (Linn.), jlde Neumann,

1899, p. 286.
albipictxhs Packard, 1867, p. Qb = Dermacentor variabilis (Say), 1821, p. 77;

Banks, 1907, fide Stiles, 1910, p. 31. Packard's description and figures

are very poor. Tick found on moose. /. albipictus also referred to

as attacking man by Holland, 1898, p. 96.
algeriensis Megnin, 1880, p. 121, PI. II. Undeterminable : see Neumann, 1901,

p. 314.

N. I. 19

282 Grenus Ixodes

ameivae Pagenstecher, 1861, p. 41, PI. II, Figs. 23, 24, hypostome and part of
basis capituli with mandibles; probablj' an Amhli/omma or Aponomma,
from Mexico, as rightly suggested by Neumann, 1899, p. 159.
'■'■ americatia'''' (Linn.) Koch, of Murray, 1877, p. 194= amencanits, see below.
americamis (Linn.) Fabricius, 1805, p. Zbb = Ambli/omma americanum (Linn.)

fxie Salmon and Stiles, 1901, p. 475.
americanus Gervais, 1844, p. 2il = Amhlyomma americanum (Linn.) fide

Neumann, 1897, p. 366.
angustus Neumann 1901, p. 2^^^ = Ixodes loricatus Neumann 1899.
annulatus Say, 1821, p. 75 ; 1859, p. \Q = Boophilus anmdatus (Say) Curtice,

1891. Original description useless ; found on Cervus virginianus,

Florida.
aptericola Maskell, 1897, p. 292, PI. XVII, Figs. 7, 8 ; included in Neumann,

1899, p. 163. We find the description and figures so poor as to render even

the determination of the genus impossible. $ tick, found on Apteryx

aiostralis, Dusky Sound, New Zealand.
apteridis Maskell, 1897, p. 291. The figures represent a capitulum of the /.

putus type. The ticks (?(?,$) were found on Apteryx mantelli., inland from

Mt Egmont, New Zealand, and may be the same as the preceding.
'■'■ aqidlae^' (in Hamburg '^\.\\iiQ\vai) = Amhlyomma decoratum Koch, 1844, p. 230;

1847, p. 94, PI. XVIII, Fig. 67. See Neumann, 1899, p. 245.
arcticus Osborn, 1899, p. 553, in Banks, 1908, pp. 23, 53, who states that

the type is lost, and the species only recorded (No. 3500) in the type

catalogue of the Division of Insects, U. S. National Museum. He quotes

Osborn's description as indicating " that the species is a good one." We

consider the description too vague to be of any use, and unhesitatingly

condemn the species. Hooker, 1909, p. 483, contrary to Banks, states

that the type is in the U. S. National Museum.
arenicola Eichwald, 1830, p. 63, PI. II, Fig. 18 (after Gervais). Insufficiently

described ; found on sand, shores of Caspian Sea and in Podolia.
aureolaUis (Fabr.) Fabricius, 1805, p. 355 (Acarus aureolatus Fabricius, 1794,

p. 4i28) = Amblyo7nma aureolatus (Fabr.).
auricidai-ius Conil, 1878, p. 99, PI. IX, Fig. 1 (/. auricularis Conil, in

Neumann, 1899, p. 166). Insufficiently described for identification, although

the author has been to great pains with his measurements, etc.
auriscutellatus Koningsberger, 1900 = Amhly omnia testridinarium Koch, 1835-

1844, Heft XI, 1 ; fide Neumann, 1901, p. 303.
australasiae Fabricius, 1805, p. 355 = ? Amhlyomma.
autumnalis Leach, 1815, p. 398 = hexagonus Leach, 1815.
a visugus Berlese, 1890, fasc. LV, No. b = hrunneiis Koch, 1844.
bengalensis Supino, 1897, p. 15, PI. II, Figs. 6-10 ; 1897, p. 268, PI. XII, Fig. 4

(tarsus). Description insufficient, from Burnin.
herlesei Birula, 1895, p. 359, PI. II, Figs. 1-6. Description and figures

insufficient ; a $ Ixodes from E. Africa.
hibroni Gervais, ] 842, p. xlviii ; Walckenaer and Gervais, 1 844, p. 248 ;

Packard, 1867, p. 68 ; also Neumann, 1899, p. 157. Probably an Ambbjommu ;

from S, America.

Co7idemned Species, etc. 283

hifxtrcatus Neumann, 1899 ( 5 ), not /. hrunneus Koch, as stated in Ncnmann,

1901, p. Ml = Dermaccntor bifurcaius Neumann, 1904, p. 453.
bimaculatus Denny, 1843, p. 312, PI. XVII, Fig. 1 {^) = Amblyomma

hippopotamense (Denny).
bipu7ictaius Risso, 1826, p. 183. Nominal .species, see Neumann, 1899, p. 158.

From S. France.
birmanensis Supino, 1897, p. 12, PI. I, Figs. 6-10 ; 1897, p. 247, PI. XII, Fig. 2

(tarsus 1); see also Neumann, 1897, p. 164. Description insufficient ; from

Burma.
boarum Stoll, 1886-1893, p. 18, PL XIII, Fig. A = Amblyomina dissimile Koch.

See Neumann, 1899, p. 227 (o).
borealis Kramer and Neuman, 1883, p. 518 = 7. puUis (Pickard- Cambridge).
bovis Riley, 1869, p. 168 = Boophilus annulatus (Say 1821) Curtice, 1891.
brevipes Murray, 1877, p. 194. Original description comprised in three lines !

Nominal species.
brevipes (Neumann), 1899 ; see Notes on Doubtful Species, p. 291.
cajeiincnsis (Fabricius) Fabricius, 1805, p. 354. {Acarus cajennensis Fabr.,

1794, IV, p. 427), undeterminable.
calcaratus Birula, (?), y>. IS7 = Boophilus anmdatiis (Say) Curtice, 1891.
californicus Banks, 1904, p. 369, PI. XLI, Fig. 57 ; 1908, p. 24, PI. II, Fig. 12 =

I. ricinus var. californicus (Banks) Nuttall and Warburton, 1911.
camelinus Fischer von Waldheim, 1823, p. 13, PL, Figs. 1 and i = Hyalomma

aegyptium (Linn.).
cenereohcs Lucas, 1840, pp. 47, 48, PL VII, Fig. 10 (after Gervais, 1844, p. 244).

Insufficiently described ; see Neumann, 1899, p. 165; from Canary Islands.
ckelifer Megnin, 1880, p. \Z% = Haemaphysalis concinna Koch, 1844.
chordeilis Packard, 1867, p. 61 — Haemaphysalis leporis palustris (Packard), /o?e

Neumann, 1897, p. 343, but stated to be a distinct species of Haemaphysalis

by Banks, 1908, p. 34.
ductus Fabricius, 1805, p. 356 ; undeterminable.
cinctus Lucas, 1840, pp. 47, 48, PL VII, Fig. 12 (after Gervais, 1844, p. 243).

Insufficiently described ; from Canary Islands.
columbae Fabricius, 1805, p. ZbQ^Argas reflects,
"communis" Marx (label: no date) = /. ricinus var. scapularis (Say) Nuttall

and Warburton, 1911.
cookei Packard, 1869 a, p. 61 = hexagonus var. cookei (Packard) Nuttall and

Warburton, 1911 ; not Haemaphysalis concinna Koch, as stated in Neu-
mann, 1897, p. 338.
cormiger Kolenati, 1857, p. 431, PL VI, Fig. 52 (very poor) = Hyalomma aegyptium

(Linn.) (i^) found on camel, dog, sheep, on the steppes of Kisil-Kum, Sea

of Aral.
coxalis Gervais, 1842, p. 47 ; Walckenaer and Gervais, 1844, p. 249. Judging

from original description may be a Dermacentor ^ ; from New Holland.
crenattts Say, 1821, p. 76; 1859, p. 20. Original description useless.
crenulatus Koch, 1835-1844, H. 39, Figs. 8, 9 ; 1844, p. 232 (listed) ; 1847, p. 22

= 7. hexagonus Leach, 1815, from Germany. "7. crenulatus Koch," as

described by Cane-strini and Fanzago, 1877, p. 116, is undeterminable.

19—2

284 Genus Ixodes

crnciarius Fitch, 1872, p. 366 ( = /. cookei fide Banks, 1908, p. 53) = /. kexa-

gonus var. cookei Nuttall and Warburton, 1911.
decorosus L. Koch, 1867, p. iAl—Aponomma decorosum (L. Koch) Neumann,

1899, p. 194.
distipes Murray, 1877, p. 194. Purely nominal species ; see Neumann, 1899,

p. 161. From Tunis.
dugesii Gervais, 1844, p. 242 (Walckenaer and Gervais), /. plurnbeus Dugfes

renamed = ? Rhipicephalus sanguineus (Latreille), as suggested in Neumann,

1897, p. 385, but undeterminable from description. On dogs in France.
dugesii Gervais, in Mdgnin, 1880, p. 126, Fig. 44 = Boophilus sp.
elegans Guerin, 1829-1843, p. 16, PI. VI, Fig. \ = Amblyomnia variegatum,

(Fabricius), 1794 ; in Neumann, 1899, p. 268.
elegans Neumann, 1910, p. 191, Fig. 1=7. stilesi Neumann, 1911 (original

name preoccupied ; species renamed.)
elephantinus (Linn.) Fabricius, 1805, p. 351 {Ac. elephantinus Linn.) = ?

Amhlyomma elephantinus (Linn.).
erinacei Audouin, 1832, p. 415, PI. XIV, Fig. 32 ( $ ) = /. hexagonus Leach, 1815,

in Neumann, 1899, p. 129.
erinaceus Murray, 1877, p. \QQ = hexagonus Leach, 1815.
erraticus Say, 1821, p. 77 ; 1859, p. 20. Original description useless ; found in

S. United States.
eudyptidis Maskell, 1885. See Notes on Doubtful Species, p. 291, and /.

putus.
eudyptidis Maskell, 1885, in Neumann, 1899, p. 128 = /. neumanni Nuttall and

Warburton, 1911.
eudyptidis var. signata Neumann, 1904, p. 451 = /. signatus Birula, 1895.
exilipes Lucas, 1846, p. 63, PI. I, Fig. 5. Undeterminable ; on Lacerta ocellata,

from Algiers.
fabricii Audouin, 1827, p. 428 (description of plates) = /Tyafomma aegyptium

(Linn.).
fimhriatus Kramer and Neuman, 1883, p. 518 = Ixodes putus (Pickard-

Cambridge, 1878).
fiavidus Koch, 1844, p. 233 ; 1847, pp. 22, 103, PI. XXI, Fig. 77 ( ? ); Neumann,

1899, p. 227, states that it is a o from Rio de J emeiro = Amblyomma

dissimile Koch.
flavipes Koch, 1835-1844, H. 39, Fig. 2 ; 1847, p. 21 {G&YmAny)= I. vesper til ionis

Koch, 1844.
flavomaculatus Lucas, 1846, p. 56, PI. I, Fig. 1 ( ,^ ); 1851, p. 120 ; 1867, p. lxxii =

Aponomma exornatum (Koch) 1844, in Neumann, 1899, p. 186.
fodiens Murray (not Megnin), 1877, p. 191 = i-icitius (Linn.) Latreille, 1806.
forskali Audouin, 1827, p. 430. An undeterminable Argas ; see our Part I, p. 6.
fuscolineatus Lucas, 1847, p.c. Description useless ; from India.
fuscomaculatus Lucas, 1873, p. xxi. Probably an Amhlyomma. See Neumann,

1899, p. 160 ; from S. America.
fuscous {fuscus) Say, 1821, p. 79 ; 1859, p. 22. Original description useless.
fuscus Koch, 1835-1844, H. 39, Figs. 3, 4 ; 1844, p. 232 (listed); 1847, p. 21 =

ricinus (Linn.), in Neumann, 1899, p. 112,

I

Condemned Sijecies, etc. 285

gervaisi Lucas, 1847, p. j.c\\ = Aponomma gervaisi {ljVLca,ii), in Neumann, 1899,

p. 182.
globulosus Supino, 1897, p. 18, PI. Ill, Figs. 11-15 ; 1897, p. 249, PI. XII, Fig. 7,

(tarsus). Description insufficient ; from Burma.
glohuhis Lucas, 1860, p. 538. Insufficiently described, may be an Aponomma.
gracileiitus Lucas, 1845, p. 58, PI. I, Fig. 2 {6) = Hyalomma aegyptium (Linn.),

in Neumann, 1899, p. 286.
granulatus Supino, 1897. See Notes on Doubtful Species, p. 291.
herrerae Dug5s, 1887? p. 487; 1891, PI. VIII, Fig. 5 {^) = Amhlyomma

cajennense Koch, 1844.
hexagonus Leach, in Salmon and Stiles, 1901, p. 467. See synonymy of /.

hexagonus var. cookei.
hexagonus var. inckoatus Neumann, 1901, p. 28S = canisuga Johnston, 1849,

p. 371.
hexagonus var. longispinosus Neumann, 1901, p. 283 = /. hexagonus var. cookei

(Packard) Nuttall and Warburton, 1911.
hippopotamensis Denny, 1843, p. 313, PI. XVII, Fig. 2 { ^ ) = Amblyomma

hippopotamensis (Denny), in Neumann, 1899, p. 256.
hirsutus Birula, 1895, p. 356, PI. I, Figs. 7-9; Neumann, 1899, p. 162.

Description and figures insufficient. From Aleutian Islands and E.

Siberia. Given as synonym of /. putus by Banks, 1908, p. 54.
hispanus (Fabricius) Fabr., 1805, p. 353 {Acarus hispanus Fabricius, 1794,

p. 426) = probably Hyalomma aegyptium (Linn.).
hispanus Kolenati, 1857, p. 431, PL VI, Fig. 52 {'^)'i— Hyalomma aegyptium

(Linn.), in Neumann, 1899, p. 286.
histrio (Fabricius) Fabr., 1805, p. 352 {Acarus histrio Fabricius, 1794, Suppl.,

p. 571). Undeterminable.
holsatus (Fabricius) Fabr. 1805, p. 355 {Acarus holsatus Fabr.); 1794, p. 428.

Undeterminable, probably young of ricinus or hexagonus ; see Neumann,

1899, p. 157.
holsatus Fabricius, in Kolenati, 1857, p. 24, PI. 11 = Dermacentor retieulatus

(Fabricius, 1794) in Neumann, 1897, p. 360. We agree in referring it to

this species after consulting the original description.
huma7ius Koch, 1844, p. 233 ; 1847, p. 104, PI. XXI, Fig. 78 ( ^) = Amblyomma

dissimile Koch ; see Neumann, 1899, p. 227 ; a o, from Brazil.
hydrosauri Denny, 1843, p. 314, PI. XVII, Fig. 4 = Amblyomma hydrosauri

(Denny), in Neumann, 1899, p. 197.
iguanae (Fabr.) Fabricius, 1805, p. 354 {Acarus iguanae Fabr., 1794, iv,

p. 427) = Amblyovima igtianae.
imperfectus Neumann, 1899, p. 118. See Notes on Doubtful Species, p. 292.
indentatus Gamgee, \SQ^ = Boophilus anmdatus (Say), in Neumann, 1897,

p. 408.
indus (Linn.), Syst. nat., ii, 1022 ; Fabricius, 1794, p. 428 (/. indus) ; 1805,

p. 355. Undeterminable.
inermis Neumann, 1901, p. 283. See Notes on Doubtful Species, p. 292.
intermedius Neumann, 1899, p. 132 ; 1904, p. 451=/. neumanni Nuttall and

Warburton, 1911.

286 Genus Ixodes

. j%ivenis Neumann, 1899, p. 124. See Notes on Doubtful Species, p. 292.
kelloggi Nuttall and Warburton, 1907, p. 396 = /. hrunneus Koch, 1844.
, . lacertae Koch, 1835-1844, H. 39, Fig. 11 ; 1847, p. 22 (Germany) =rimiMS

(Linn.), in Neumann, 1899, p. 112. "Zr. lacertae" in Pagenstecher, 1861.

p. 41, Fig. 22 ; may be the same.
laevis Neumann, 1899, p. I'i?) = acutitarsus (Karsch), 1880.
lagotis (rervais, 1849, y>. 49. Insufficiently described ; may be a Haemaphysalis.
Zeac/n'i Audouin, 1827, p. 428. (Savigny, PI. IX, Fig. 9) = Iluemaphysalis leachi

(Audouin).
leporis-palustris Packard, 1867, p. Ql = Haemaphysalisleporis-palustris (Packard).
Imeatus (Fabricius) Fabr., 1805, p. 354. (Acarus lineatus Fabricius, 1794,

p. 428.) Undeterminable.
linnei Audouin, 1827, p. 428 ; Savigny, 1825, PI. IX, Fig. 12 ; Walckenaer and

Gervais, PI. XXXIII, Fig. 1 = Rhipicephahis s.^.1 Undeterminable.
Upsiensis (Fabricius) Fabr., 1805, p. 354. {Acarus lipsiensis Fabricius, 1794,

pp. 427, 428) = young of ricimis or hexagonus ; see Neumann, 1899, p. 157.
lividus Leach, 1824, ii, in Koch, 1844, p. 234 (undeterminable) = young of

ricinus or hexagonus in Neumann, 1899, p. 157.
lividus van Beneden, 1873, XL. A purely nominal species from Vesperugo.

No description ; see Neumann, 1899, p. 160.
longipes Lucas, 1879, p. I's.xiv = vespertib'onis Koch, 1844, p. 232 ; in Neumann,

1899, p. 169.
luteus Koch, 1844, p. 232 ; 1847, p. 102, PI. XX, Fig. 75 ( ? ). Species based by

Koch on one o from S. Africa. Neumann, 1899, p. 146, and 1901, p. 285,

referred a $ to this species, the tick having been taken from a wild dog

in Africa (Paris Museum). Insufficiently described.
marginalis Hahn, 1834, p. 63, PI. LXVI, Fig. 153. Useless description,

undeterminable.
marginalis in Gervais, 1844, p. 242 = ? (Salmon and Stiles, 1901. Synonymy of

/. ricinus).
marginalis Koch, 1835-1844, ii, p. 63, PI. LXVI, Fig. 53 (after Gervais).

Insufficiently described ; see Neumann, 1899, p. 165.
marginatus (Fabricius) Fabr. 1805, p. 354 {Acarus marginatus Fabricius, 1794,

p. 427) = ^r^as reflexus (Fabr.) Latreille, 1796.
mai-ginatus Burmeister, 1837, p. 579 = /. ricinus, fide Oudemans, 1896, p. 191

in Salmon and Stiles, 1901, p. 466.
marmoratus Eisso, 1826, p. \S'^ = Dermacentor reticulatus (Fabricius) 1794, in

Neumann, 1897, p. 360.
maskellii Kirk, 1887, p. 66. Undeterminable, description useless. Found on

Albatross {Diomedia exidans), New Zealand.
megathyreus Leach, 1815, p. 398 = /. 7-icinus (Linn.).
mixtus Moniez, 1896, p. A91 = Amhlyomma mixtum Koch = Amblyomma caj'en-

nense (Fabr.), in Neumann, 1901, p. 348 (Index).
moreliae L. Koch, 1867, p. 241 = Amblyomina moreliae (L. Koch), according to

Neumann, 1899, p. 258. Description insufficient to make it a good species,

however, in our opinion. Found at Bi'isbane on Morelia argus var.
fasciolatu Jan.

Condemned Species, etc. 287

naponensis Packard, 1867, p. Qb = Denncicentor electus Koch in Neumann, 1897,

p. 366 ; but electus = D. variabilis (Say, 1821) /t/e Stiles, 1910, p. 29.
nigrolineatus Packard, 1867, \>\). 66, Ql = Dennaecntor nigrolineahis (Packard)

Banks, 1907, /?o?e Stiles, 1910, p. 51. Insufficiently described by Packard,

who appeared to describe a (^ as a $ ; doubtfully referred to Haemaphy-

salis concinna Koch by Neumann, 1897, p. 338.
nigua (de Geer) Latreille, 1804, p. 52 = Ainblyomma aniericantim (Liini.) Koch,

1844 ijide Salmon and Stiles, 1901, p. 475.
nigua Guivin = AvMyomma americanum (Linn.) Koch, 1844, in Ne'imann,

1901, p. 342 (Index).
ohliquus Koch, 1844, p. 232 ; 1847, p. 99, PI. XX, Fig. 73 (? and its scutum).

Insufficiently described.
obseuriis Fabricius, 1805, p. 355. Undeterminable.
obscurus Neumann, 1899, p. 121 = nigricans Neumann, 1908, p. 75. (Renamed,

name preoccupied.)
ophiophilus Miiller, 1831, p. 233, PI. LXVII = ? Aponomma gervaisi (Lucas),

1847, in Neumann, 1899, p. 182.
orbiculatus Say, 1821, p. 76 ; 1859, p. 20. Original description useless. Found

on Sciurus capistratus, S. United States.
^'' oregonensis" (label), in U. S. National Museum = Dermacentor salmoni Stiles,

1910, pp. 55, 60.
ovatus Neumann, 1899, p. 116, Figs. 2, 3 = in partrjcm?<s (Linn.), says Neumann,

1904, p. 452.
ovatus Neumann, 1899, p. 112, 1904, p. 452 = /. ricinus var. ovatus (Neumann).

Nuttall and Warburton 1911.
pallens Fabricius, 1805, p. 356. Undeterminable.
pallipes {Acarus pallipes Fabr., 1794, p. 426) referred to in Koch, 1835, H. 39,

10 ; 1847, p. 22 (Germany) ==? /. bncnneus Koch, 1844 (larva).
pallipes Lucas, in Webb and Berthelot, 1840, pp. 47, 48, PI. VII, Fig. 9 (after

Gervais, 1844, p. 243). Undeterminable.
pari Leach, 1815, p. 399, on ^' Partis major" = 1 I. brunneus Koch, 1844.
parvirostris Neumann, 1901, p. 284 = 7. signatus Birula, 1895.
perpunctatus Packard, 1867, p. 68. Probably an Amblyomma ; see Neumann,

1899, p. 159. From S. America.
phascolomys Macalister, 1871, p. 163, Fig. A purely nominal species ; see

Neumann, 1899, p. 160.
pictus Gervais, 1844, p. '2Z^ = Dermacentor retimlatus (Fabricius), in Neumann,

1901, p. 265.
pilosus var. howardi Neumann, 1908, p. 125 = /. pilosus Koch.
plumbeus (Panzer) in Fabricius, 1805, p. 353 {Acarus plumbeus Panzer).

Undeterminable.
plumbeus Leach, 1815, p. 397 ; 1824, p. 11. Undeterminable =/. lividus Koch,

in Neumann, 1901, p. 348 ; 1899, p. 157 ; see lividus in this list. On

Hirundo riparia.
plumbeus Dug^s, 1834 c, PI. VII, Figs. 1-\2 = Rliipicephalics sp., as clearly

indicated by the figures ( = / ricinus., according to Neumann's synonymy).

Also figured by Wagner, 1841, Figs. XII, XIII (unrecognizable).
plumbeus in Wheler \%^^ = Ixodes canisuga Johnston, 1849.

288 Genus Ixodes

poortmani Lucas, 1850, p. xli ; Belval, 1861, p. 97, PI. = Amhlyomma hebraeum

Koch, in Neumann, 1899, p. 266.
praecoxalis Neumann, 1899, p. 121=7. neumanni Nuttall and Warburton,

1911.
pulchelhis Lucas, 1845, p. 61, PI. I, Fig. 4 ; 1849, p. Ixxx ; 1849, p. 582 ((J, good

figure of capitulum) = .4m6/yomwa dissimile Koch, in Neumann, 1899,

p. 227.
punctulatus Say, 1821, p. 78 ; 1859, p. 21. Original description useless, said to

resemble " /. variabilis ^^ = Dermacentor variabilis (Say, 1821).
punctulatus Canestrini and Fanzago, 1877-1878, pp. 115, 183 ; Canestrini, 1890,

p. 502. Doubtful if it i.s an Ixodes ; see Neumann, 1899, p. 160. From

Europe.
ptistidarum Lucas, 1866, p. \xn = Ixodes ricinus (Liim.), in Neumann, 1899, p. 113.
pygmaeus Koch, 1844, p. 233 ; 1847, p. 107, PI. XXII, Fig. 81 ( ? ), from Brazil

and Mexico. Types examined by Neumann (1901, p. 289), and found to

be Amblyomma nymphs, of undetermined species.
qu,inquestriatus Fitch, 1870, p. 366 = Dermacentor variegatus Marx and Neumann,

in Neumann, 1901, p. 266.
reduvius Audouin, 1832, nee Linnaeus {^), p. 422, PI. XIV, Fig. 4 = /. hexa-

gonus Leach, 1815.
reduvius Hahn, 1834, p. 62, Fig. 152 = 7. ricinus (Linn.), in Neumann, 1899,

p. 112.
reduvius de Geer, in M^gnin, 1880, p. 126 = 7. ricinus (Linn.).
reduvius {hmw.) =ricimis (Linn.), in Neumann, 1901, p. 348, and numerous

other authors, this incorrect, see p. 143.
reflexus (Latreille) Fabricius, 1805, p. ^^2 = Argas rejlexus (Fabr., 1794, p. 426)

Latreille, 1796. (See our Part I, p. 22.)
reticulatus Latreille, 1804, p. \bl = Dermacentor reticulatus (Fabricius), 1794, in

Neumann, 1897, p. 360.
reticulatus (Fabr.) Fabricius, 1805, p. 355 {Acarus reticulatus Fabr., 1794,

p. 428). Undeterminable.
reticulatus Koch, 1856, p. 413. Insufficiently described ; see Neumann, 1899,

p. 166.
reticulatus Koch, in Rosenhauer, 1856, p. 412. Undeterminable, found on

Lacerta ocelata, in Spain.
rhinocerinus Denny, 1843, p. 313, PI. XVII, Fig. 3 = Dermacentor rhinocerotis {de

Geer), in Neumann, 1897, p. 370.
rhinocerotis Fabricius, 1805, p. 'ib\ r= Dermacentor rhinocerotis (de Geer), 1778.
rhinocerotis Gervais, 1844, p. 246 (Walckenaer and Qi&x\'SA^) = Dermacentor

rhinocerotis (de Geer) 1778, in Neumann, 1897, p. 370.
neinMs ^Megnin, 1880, p. 129 nee Latreille = 7. hexagomis Leach, 1815; see

Neumann, 1899, p. 129.
rostralis Moniez, 1896, p. AQQ = Gonixodes rostralis Dugfes, 1888, p. 129 =

Haemaphysalisle-poris-palustris (Packard) in Neumann, 1901, p. 348 (Index).
rubiginosus (Kolenati), Neumann, 1899 {Dermanyssus rid)iginosus Kolenati,

1857, p. 20, PI. I). Probably an Ixodes, says Neumann, 1899, p. 158.

Found on Plecotus auritus.
rufipes Fabricius, 1805, p. 354. Undeterminable.

Condemned Species, etc. 289

rufus Koch, 1835-1844, H. 39, Fig. 7 ; 1844, p. 232 (listed); 1847, p. 22 ; os,

types examined by Neumann, 1901, p. 282 = /. ricinus (Linn.).
sanguineus Latreille, 1804, p. \bl = Rhipicephalus sanguineus (Latreille) in

Neumann, 1897, p. 385.
sanguisugus (Fabricius) Fabr., 1805, p. 353 {Acarus sanguisugus Fabr., 1794,

p. 426). Undeterminable.
savignyi Gervais, 1844, p. 244. Atla«, PI. XXXII, Fig. 1; XXIII, Fig. 2 =

Hyalomina aegyptium (Linn.) (/. aegyptius Audouin renamed, as he thought,

judging from published figures, that it differed from Cynor. aegyptius

Hermann).
scapularis Say, 1821, p. 78; 1859, p. 21 = /. ricinus var. scapularis (Say)

Nuttall and Warburton, 1911. Original description useless.
scapulatus Megnin, 1880, p. 132. Undeterminable.
tciim Koch, 1835-1844, H. 35, 8 ; 1844, p. 232 (listed); 1847, p. 21 (Germany)

= /. ricinus (Linn.) os, types examined by Neumann, 1901, p. 282.

/. sexpunctatus Koch, 1835-1844, H. 39, 5, 6; 1844, p. 232 (listed); 1847,

p. 22 = /. hexagouus Leach ; see Neumann, 1901, p. 283, who examined

the types (os). Canestrini and Fanzago, 1877, p. 114, described a 9 (?)

as sexpunctatus, but their description is useless.
siculifer Megnin, 1880, p. 132 = /. vespertilionis (Koch) 1844.
spinosus Neumann, 1899, p. 146= fuscipes Koch, 1844.
b-striatus Fitch ; see quinquestriatus.

sturni Pagenstecher, 1861, p. 40=? /. brunnetis Koch, 1844 (larva).
sulcattis Koch, 1844, p. 233; 1847, pp. 22, 108, PI. XXII, Fig. 82 (?) = /

ricinus (Linn.) os. Types examined by Neumann, 1901, p. 282.
sylvaticus Gervais, 1778, p. 162, PI. 38, Fig. 1 = Amhlyomma sylvaticum (de

Geer), in Neumann, 1899, p. 274. On tortoise. Cape of Good Hope. May

attack man.
testudinis Conil, 1877, p. 26 = Amhlyomma testudinis (Conil), see Neumann,

1905, p. 234.
testudinis Supino, 1897, p. l\ = Aniblyomma supinoi Neumann, 1905, p. 234.
testudinis Leydig, 1855, p. 382, PI. XV, Fig. 51, also p. 395; 1857, p. 111 =

Hyalomma syriacum, no doubt.
thoracicus Koch, 1844, p. 232 ; 1847, p. 98, PI. XIX, Fig. 72 ( ? ). May have

been an Amblyomma, as Neumann, 1904, p. 450, rightly suggests.
^'thoracicus Koch " in Neumann, 1899, p. 149 = aurit7dus Neumann, 1904, p. 450.
trabeatus Audouin, 1832, p. 420, PI. XIV, Fig. 3 ($) = / ricinus (Linn.), in

Neumann, 1899, p. 112.
trackysauri Lucas, 1861, p. 125 = Aponomma trachysauri (Lucas).
transversalis Lucas, 1844, and in Murray, 1877, p. IdH = '^ Netcmanniella

transversalis (Neumann)" Lahille, \904 = Aponomma transversale (Lucas)

Neumann, 1899, p. 188.
trianguliceps Birula, 1895, p. 358. See Notes on Doubtful Species, p. 293.
trilineatus Lucas, 1840, pp. 47, 48, PI. VII, Fig. 11. Insufficiently described,

Canary Islands.
trimaculatus Lucas, 1878, p. Ixwn = Aponomma trimaculatum (Lucas), in

Neumann, 1899, p. 187.

290 Genus Ixodes

tristriatus (Panzer), 1795, H. 59, n. 24 ; Koch, 1835-1844, fasc. 59, Fig. 24 ; 1844

p. 234 (listed) = young of ricinus or hexagonus ; see Neumann, 1899, p. 157.
troglodytes Schmidt, 1853 ; see Frauenfeld, 1853, p. 57 = /. vespertilionis Koch.
undatus (Fabricius) Fabr., 1805, p. 353 {Acarus imdatus Fabricius, 1794,

p. 427). Undeterminable.
"unipictits Packard," of Verrill, 1870, p. 118 (lapsus for unipunctata (q.v.) fide

Salmon and Stiles, 1901, p. 476).
unipunctata Packard, 1867, p. 66 = ^1 Hift^j/omma a?Heri'ca?M«?i Koch, in Neumann,

1899, p. 209.
uriae White, 1852. Nominal species, found on Uria troile at Baffin's Bay.

Description and figure inadequate; may=? /. putus.
varanensis Supino, 1897, p. 13, PI. II, Figs. \-b = Aponomma gervaisi (Lucas),

in Neumann, 1899, p. 182.
varanilj. Koch, 1867, p. 'iW^ Aponomma decorosum (L. Koch), in Neumann,

1899, p. 194.
varani Lewis 1892, p. 10, PI. 1 = Aponomma exornatum (Koch) 1844.
variabilis Say, 1821, p. 77 ; 1859, p. 21. Original description useless, said to

resemble erraticus a,nd punctidat us. Listed, however, by Banks, 1895, p. 433,

as occurring at Fort Collms = Bermacentor variabilis (Say) Banks, 1907,^ci?e

Stiles, 1910, p. 29.
variegatus Lucas, 1852, p. \x:&.xvm = Amblyomma albopictum Neumann, in

Neumann, 1899, p. 244.
variegatus (Fabricius) Fabr., 1805, p. 353 {Acarus variegatus Fabricius, 1794,

IV, p. 572) =Amblyomma variegatum (Fabricius).
variolatus Gervais (Walckenaer and Gervais), 1844, p. 249. Probably an

Amblyomma. From Brazil, on a reptile.
vibrans (Fabr.) Fabricius, 1805, p. 356, No. 28 (1794, p. 428, 20 : Acarus

vibrans). Undeterminable.
viperariim Koch, 1844, p. 234 ; 1847, p. 102, PI. XX, Fig. 76 (lax'vae). From

Greece, undeterminable.
vulgaris Fabricius, 1805, p. 352 = /. ricinus or hexagomis.
vulpis Pagenstecher, 1861, lo. 40, PI. I, Figs. 12, \Z = hexagonus Leach, 1815.
walckenaeri Gervais, 1842, p. xlvii ; 1844, p. 246, PI. XXXIV, Fig. 11 = ?

Amblyomma. Undeterminable.
Sarconissus Kolenati, 1856 {Sarconyssus Kolenati, in Murray, 1877, p. 195 ;

Marx, 1892, p. 235, etc., referred to Haemalastor and then to Eschatoce-

phalus by Neumann).
brevipes Kolenati, 1856, p. 21=/. vespertilionis Koch.
exaratus Kolenati, 1856, p. 22 ; 1860, p. 757, PI. I, Fig. 2 = /. vespertilionis Koch,

vide Neumann, viii. 1910, p. 191, who examined type, a nymph.
flavidus Kolenati, 1856, p. 21=/ vespertilionis Koch.
flavipes Kolenati, 1856, p. 21=/. vespertilionis Koch. (Neumann 1910, p. 192,

states the types were not adults.)
hispidiilus Kolenati, 1856, p. 21 = /. vespertilionis Koch.
kochi „ „ p. 21 ; 1860, p. 573, PI. II, Fig. 4 = 1. vespertilionis

Koch, (^ .
nodulipes Kolenati, 1860, p. 576, PI. I, Fig. 3 = probably /. vespertilionis Koch.

Notes on Douhtful Species 291

NOTES ON DOUBTFUL SPECIES OF IXODES.

Ixodes aequalis Jianks, xr. 1909, p. 27G ; 1910, p. 6, PI. Ill, Fig. 23 ($ ciii)itulum
and scutum, sketchy). The author's description and figure are inadequate : the
essential data which we glean therefrom are as follows :

Male : unknown.

Female : Scutum 1 mm. /., nearly tiiamond shaped, antero- and postero-
lateral borders suhequal, many uniform punctations, lateral carinae distinct.
Spiracle circular. CapituUim subtriangular, porose areas rather large, sub-
circular, the interval less than half their diameter. Legs: coxae I-IV with
minute spur at postero-external angle, coxa I with short internal spine ; tarsi
tapering abruptly.

Host : Otospermophilus beecheyi (California ground squirrel), Berkeley,
California (Drs Wellman and Wherry). Banks states that aequalis differs
from angustus in having a shorter and differently shaped scutum, more circular
porose areas, shorter tarsi, etc. According to Hooker, 1909, p. 423, the type is
in Banks' collection, Washington, D.C.

Ixodes brevipes (Neumann), 1899 {Eschatocephalus crassipes Joseph, 1882, p. 16,
renamed owing to crassipes being preoccupied) ; only (^ s found on stalactites in
caves (Ihanska jama, Goba dol, etc., Krain, Austria), the hosts probably being
bats. Insufficiently described ; said by Joseph to have shorter legs than /.
vesjjertilionis.

Ixodes eudyptidis Maskell, 1885, pp. 19, 20, PI. VIII, Figs. 12-14. The only
points given in the author's bad description, and which might be utilized, may
be summarized as follows ; Scutum hairless, glossy ; many fine, shallow puncta-
tions. As the author says, it is "evidently a true tick," and that is all that
we can say about it. It was found " in the gape of the penguin," at Dusky
Sound, New Zealand. See also Synonymy under /. neumanni N. and W.
Judging from the bad figures, it is possible that eudyptidis= I. putus.

Ixodes granulatus Supino, 1897, p. 16, PI. Ill, Figs. 1-10, and 1897, p. 250, PI. XII,
Figs. 5, 6. Also Neumann, 1899, p. 164 ; 1902, p. 125. Supino's description
and figures are insufficient. Neumann found what Supino described as a ^ to
be either a 5 or o. Neumann supplements Supino's description, and regards
the species as allied to 1. minor. The points which may serve for determina-
tion are as follows :

Female: L. 4-5, W. 3'5 mm. Scutum regularly oval (lxO"75 mm.);
cervical grooves scarcely visible ; lateral grooves indicated by a faintly marked
ridge running parallel to the borders ; many fine and uniform punctations ;
emargination very slight. Anal grooves diverging widely behind. Capitulum
long, with dorsal base subtriangulai", porose areas rounded, divergent.
Hypostome long, narrow, lanceolate, 3 | .3, 10-11 sharp teeth per file; digit

292 Genus Ixodes

slender (110 /n ?.), external article 5-cusped ; palps slender, inserted very low
down, article 2 almost twice as long as 3. Legs : coxa I with short spur at
each posterior angle, the inner spur longer ; a tuberosity external to the
others, scarcely visible on coxa IV. Legs slender, relatively long, tarsi
tapering, pad two-thirds the length of claws.

Found on Scmrus yordoni, S. rufigenis, S. striatus and Felis tigris, at
Bhamo, Mooleyis, Tarrawaddy and Terinzo in Upper and Lower Burma,
according to Supino,

Ixodes imperfectus Neumann, 1899, p. 118 (no figure); species founded on 2
nymphs^ having the following characters (condensed from Neumann):

Nymph: Body 2-1 x 1-2 mm. Scutum oval, glossy (0-56 x 0-51 mm.), with
lateral borders almost straight along their anterior half ; cervical grooves well
marked, almost attaining the posterior border ; lateral grooves clearly defined,
straight, divergent behind ; punctations fine, distant. Venter : sexual grooves
divergent, straight, long ; anal grooves parallel behind ; many fine punctations.
Capitulwn (0'33 mm. I.) with base broader than long, narrowed conically
in front ; pointed cornua ; auricula forming a retrograde spine beneath the
postero-dorsal angle ; hypostome narrow, lanceolate, 2 | 2, the external teeth
long, pointed ; palps long, narrow, article 2 longer than 3. Legs long ; coxae
with a spine at the postero-external angle, decreasing in size from pair I to
IV ; a stouter spine at the postero-internal angle of coxa I ; tarsi thick, long
on pairs I and IV, tapering gradually ; pads as long as claws.

Host : Didelphys pusilla, Brazil (Goeldi coll.).

Ixodes inermis Neumann, 1901, p. 283 (no figure). Included under doubtful
species because the description is inadequate.

Male : unknown.

Female : Body 4 mm. long (capitulum included), yellowish. Scutum reddish
brown, light-coloured, glabrous, as broad as long, lozenge- shaped (the lateral
angles toward the middle of the length), no lateral grooves, punctations
numerous ; surface slightly shagreened. Dorsum and venter bearing short
hairs ; anal grooves diverging considerably. Base of capitulum more than
twice as broad as long ; hypostome lanceolate, bearing two marginal rows of
stout teeth and numerous denticles in front ; palps relatively short. Coxae
unarmed ; tarsi fairly long, humped near their extremities.

From Neumann's description, based on 3 $ s and 4 o s collected by Z.
Wagner, locality not given (Berlin Museum). (Two ^s. identical with Ixodes
ricmus were found in the same lot, but difiered too much from the $s to
permit of their being considered as belonging to the same species.)

Ixodes juvenis Neumann, 1899, p. 124 (no figure). Species founded on 6 nymphs
and 1 larva, having the following characters (condensed from Neumann):

Nymph : Body ovoid (2 x ri mm.), dark brown. Scutum rounded, slightly
broader than long, of the same colour as the rest of the body ; punctations few,
very fine ; some short hairs ; cervical grooves attaining the posterior border.
Dorsum bearing long hairs, especially on the borders ; marginal groove

Notes on Doubtful Sjjecies 293

superficial, almost forming the lateral border. Veiite?- glabrous, or almost so ;
sexual grooves straight, divergent, attaining the posterior border ; anal grooves
parallel, united in an arc in front. Spii-aclc large, circular. Capitulvm
0-35 mm. I., base rectangular, broader than long ; hyjiostome elongate, lanceo-
late, 2 I 2, 10-11 teeth per file, the external pointed, the internal blunt. Palps
of medium length, article 3 equal to 2. Legs of medium length, dark brown ;
coxae almost contiguous, glossy, with some very short hairs ; a tuberosity at
each of the two posterior angles, more pronounced on coxa I, almost obsolete on
coxa IV. Tarsi elongate, slightly tapering at their tips ; pad almost as long as
claws.

Larva : similar to o, but shorter (1'2 mm.), hexapod.

Host : Holotropis (Iguana) from New Grenada (Paris Mus.).

Ixodes trianguliceps Birula, 1895, pp. 358, 359, PI. I, Figs. 14, 15 ( $ capitulum, coxa).
This species appears to be closely related to, if not identical with, /. tenuirostris
Neumann. Birula's figures are obviously drawn from a mounted specimen and
consequently it is impossible to reach any conclusions as to its identity. We
wrote to Professor Birula (St Petersburg) for the loan of the (unique) type, but
he feared it might be lost in transit. Should Birula's specimen prove to be
identical with Neumann's type, then trianguliceps would have priority. Birula's
description of the scutum of ti'imiguliceps as being as broad as long may be due
to distortion in the mounted specimen.

294

NOTES ON THE BIOLOGY OF IXODES

By G. H. F. NUTTALL.

PAGE

Introductory 294

Outline of the life-liistory in Ixodes 295

Ixodes ricivus .......... 296

Duration of parasitism upon the host .... 298

Site of attachment upon the host .... 299

The time required for metamorphosis to take place in the

different stages ........ 300

Longevity .......... 301

Time required to complete the life-cycle .... 801

The influence of season and local conditions . . . 303

Copulation .......... 303

Oviposition .......... 308

Relation to disease 312

Eedwater in cattle — Note re Karyolysus . . . 312

Effects of bite on man and animals .... 313

Penetration beneath the skin 314

Ixodes ricinus var. scapularis ....... 315

Ixodes angustus .......... 315

Ixodes pilosus .......... 316

Ixodes canisiKja .......... 316

Ixodes piitus .......... 317

Appendix I. On the process of copulation in Ornithodorus

monbata .......... 318

Appendix II. On the adaptation of Ticks to the habits of

their hosts ... ....... 324

Introductor'y.

Of the 51 species which we recognize as valid, there are but two
about which we are still somewhat in doubt, namely, bicornis, which
may be identical with diver sifossus, and nigricans, which may be but a
variety of ricinus. This leaves 49 well-established species. Of these,
the life-histories of but 8 are known in so far as both sexes, the nymphs
and larvae have been recorded. Only a few of the latter have actually
been raised experimentally through some of their stages. Partial
raising experiments have been carried out in Cambridge with ricinus,

Biology 295

hexagonns and canisuga, the evidence afforded being sufficient to make
it possible to identify the different stages of these species. Lounsbury,
at the Cape, has raised j9i7osws experimentally; and Hadwen, in British
Columbia, has made partial raising experiments on angastus. Of the
remaining species whose various stages are known {tenuirostris, ves-
pertilionis and loricatiis), the close resemblance between the various
stages encountered upon the same host has been accepted as sufficient
evidence for their belonging to the same species. This method of
identification has been applied to nearly all the other species we
recognize in which the life-histories are less completely known. Strictly
speaking, the identification of males and females as belonging to one
species should depend upon their being found in copula; but failing
this, we are obliged to rely upon the evidence afforded by their mor-
phology and their both occurring together upon a host.

In the list of recognised species, given in the table at the end of this
fasciculus, the table serving likewise as an index, the various stages of
each species known to science are indicated by the signs f^, $, o, L,
which stand for male, female, nymph and larva respectively. From this
list it will be seen, if we exclude varieties of some of the species, that
the

(/, $, o and L are known in 8 species.

</•, $ and o „ „ „ 4

(^ and $ „ „ „ 6

o IS ,, ,, 3 ,,

? » „ „ 20 „

$ and O, are „ „ 4 ,,

$, O and L, „ „ „ (j „

It follows that these species are known by 21 J^s, 48 $s, 22 os and 14
larval forms.

Outline of the life-history in Ixodes.

Before proceeding further, it appears expedient to briefly outline the
life-history of an Ixodes as typified by Ixodes ricinus. I shall leave
out of consideration the changes which take place in the internal anatomy
of the ticks whilst attached to the host and subsequently, for these
matters will be considered in a special section of this work. I propose
here to merely outline the main facts in the life-history.

296 Genus Ixodes

Ixodes ricinus.

Commencing with the adult stages : males and females are both
present upon the host, and they are frequently found thereon in copula-
tion. Pairing may, however, take place before the sexes have attached
themselves to a host ; this has as yet only been observed in unfed adults
in captivity. A female may pair with several males in succession.
The males, which are relatively scarce, remain attached to the host only
for a few hours and if they feed they can do so but moderately, their
external structures not permitting more than a limited amount of
distention. The males wander about the host in search of females
during the intervals when they are not feeding^ The females, on the
other hand, after having attached themselves to a host, remain in situ
hanging on by their mouthparts for 7-14 days and possibly longer in
cold weather. During this time the females are sought by the males
and gradually grow in size owing to the distention of their bodies with
blood derived from the host. Shortly before they abandon the host they
swell rapidly and attain their maximum degree of distention. Females
in all degrees of distention are found coupled with males. At times,
one or more additional males may be observed about the female whilst
one male has its mouthparts fixed in the vulva. It would appear as if
the other males were waiting their turn to enter into copulation. We
shall presently describe the process of copulation more fully.

The fertilized and fully gorged female now releases herself from the
host and falls to the grounds I have frequently observed females

1 Although I. ricinus s may attach itself to a host it has not as yet been demon-
strated that it actually sucks blood for there is no record of the intestinal contents having
been examined after a male has detached itself from the host. It is conceivable that the
attachment may be mechanical, but the matter requires further investigation. It appears
worth noting here that we have seen males of Hyalomma syriacum bury their mouthparts
deeply into a cork whilst confined in a bottle after their removal from a tortoise.

2 That the dropping of gorged females from the host may be influenced by temperature
was once observed to a remarkable degree when experimenting with Boophilus decoloratus
which was being raised by me in Cambridge. A cow infested with this tick, many females
being replete, was taken out of the warmed experimental stall into the cool air of a courtyard.
The females immediately began dropping off and " rattled like peas " on the ground. In
nature it is a common matter to pick up ticks along the paths pursued by cattle or game,
and this observation on Boophilus suggests a possible explanation why these paths should
at times be particularly infested, for it is along them that animals like cattle frequently
return from pasture in the cool of the evening. In nature, the female will at most wander
a foot or two from the path pursued by the host from which she fell, and will lay her eggs
at the foot of the vegetation bordering the path. The offspring would, therefore be
literally "put in the way of finding their food."

Biolof/!/ <{f I. rlelinis 297

which have thus dropped from the host mate again with one or more
males placed in a vessel. It is probable that this occurs in nature,
since it has been observed in other species {canisaga, angustus, putus).
Having fallen upon the ground, and being abandoned by the male, the
female proceeds to find shelter beneath the surface vegetation or stones,
or she may burrow into the loose soil. She is fairly active at first and can
climb up smooth surfaces like glass to an astonishing degree considering
the hugely augmented weight, due to the great amount of blood which
she has ingested. If placed in a glass jar covered with gauze and con-
taining earth at the bottom, in the centre of which is placed a tuft of
grass, which is occasionally sprinkled with water, it will be seen that the
females congregate about the grass roots in chinks and crevices in the
earth, and that they lie there very quietly for days or weeks before the
eggs begin to appear. The time when oviposition takes place is
largely influenced by temperature, being greatly retarded by cold.
We shall presently consider the mechanism of oviposition and its
duration, for the latter is also markedly influenced by temperature.
The eggs are next observed in small heaps lying upon the antero-dorsal
surface of the female and their number increases from day to day.
Slightly gorged females lay fewer eggs than do the fully gorged
specimens, and the few eggs they lay usually do not hatch out. From
this it may be gathered that such females are not fertilized or that their
eggs are otherwise imperfect. The eggs are usually very numerous,
numbering hundreds or a thousand or more (see PI. VII, p. 310).

Following upon an interval of varying length, this again depending
upon temperature, the process of hatching commences. From the egg-
shells emerge vast numbers of larvae. If they hatch out in a glass
vessel containing grass the larvae climb up the grass blades and con-
gregate in masses upon the tips of the leaves, or form clusters on the
under side of the gauze which encloses them in the vessel. In nature
they behave in a similar manner, that is, the}- climb to the tips of the
grass blades and the adjacent vegetation.

The larva is hexapod, and possesses neither respiratory nor sexual
organs. After it emerges from the e^g its chitin requires some time to
harden and darken, and during this period it assimilates the yolk-mass
derived from the egg. When the larva has exhausted this reserve food-
supply, it attaches itself to a host and proceeds to suck blood. When
replete, after 4 to 5 days\ it releases its hold, and, falling to the ground,

1 It is possible that the period of attachment to the host may be prolonged in the
various stages by cold weather. Observations in this respect are lacking.

N. I. 20

298 Genus Ixodes

proceeds to undergo its metamorphosis. This process is much influenced
by temperature, being retarded by cold and accelerated by warmth.
At first the replete larva is quite active, but when it has begun its
metamorphosis it lies still. Beneath the larval skin a series of profound
changes takes place. The nymphal exoskeleton developes beneath that
of the larva. The first three pairs of legs are formed within the larval
legs, and a fourth pair appear posterior to these, as can readily be seen
in mounted specimens of the ticks at a period shortly before they emerge
from the larval skin.

The nymph possesses two spiracles, similar to those of the adults in
general structure, from which branching tracheae spread to all parts of
the body. When matured, the nymph withdraws its legs from the
larval skin which encloses the first three pairs, it ruptures the old skin,
emerges, and abandons the exuviae. The nymph differs from the female
in two essential points of external structure : it does not possess an
external sexual orifice, although there may be an indication (Anlage) of
where it will be situated in the adult, and the nymph does not possess
porose areas upon the dorsal surface of the basis capituli. The nymph
takes some ten days to harden and darken after casting the larval skin,
and it is now ready to attack a fresh host upon which it behaves
similarly to the larva, dropping off gorged after three to five days.

The adult develops within the nymphal skin and emerges from it
after a variable interval of time (weeks or months). The sexual organs
in both sexes begin to appear in the nymph but they do not attain their
full development in the female until she has sucked blood. The adults,
having hardened, proceed to attack a fresh host, thus completing the
cycle.

It will be noted that ricinus, in common with most Ixodidae
requires three hosts upon which to feed in the larval, nymphal and
adult stages.

Duration of parasitism upon the host.

Larvae and Nymphs. The following experiments may be cited to
show the behaviour of this tick when raised in the laboratory. On
July 5, 1905, some gorged ?s, taken from cattle in Co. Cavan, Ireland,
were received in Cambridge. The ticks were placed at room tempera-
ture in a receptacle containing sand which was occasionally dampened.
Large masses of eggs had been laid by the end of July from which
innumerable larvae issued in September.

Biology of I. ricinus

299

13.

)?

14.

))

16.

))

18.

I. 1906.

3.

VI. „

7.

5>

8.

))

Experiment II. 18.

X. 1905.

23.

))

24.

n

18.

I. 1906.

Experiment I. 10. x. 1905. Many larvae were placed on a hedgehog in a

tick-proof cage.
68 larvae dropj^ed off' gorged.
365 „

f'O )) » )) „

Nymphs began to emerge from the larval skins.
Thirty of these nyrophs were placed on a hedgehog.
1 0 nymphs dropped off gorged.

Many larvae were placed on a hedgehog.
250 larvae dropped oft' gorged.

Nymphs began to emerge.
1. VIII. 1906. 50 nymphs were placed on a hedgehog.
5. „ 20 „ dropped off gorged.

It follows that the larvae remain attached to the host for 3-6 days, and
the nj'mphs for 4-5 days, when raised on hedgehogs in the laboratory
at a temperature of about l.^*^ C.

These results are in agreement with the statement by Knssel,
Schiitz, Weber and Miessner (1903, p. 40), that larvae remain 3-6 days
and nymphs remain 3—5 days upon the host before they become replete.
These authors raised the ticks on cattle and guinea-pigs, Samson
(1909, p. 186) reports that larvae and nymphs remain attached for four
days to man, and as long as 10-14 days to lizards.

Adults. Bertkau (1881, p. 14-5) reports that he once placed a
hungry (/• and $ upon his arm, with the result that the (/* remained
attached for oidy eight hours whilst the $ lemained upon him for eight
days. Kossel and his colleagues once saw hungry /s readily attack a
guinea-pig upon which they were placed ; they state that the $ remains
attached for a week. According to Samson (1909, p. 216), the $ remains
upon the host for 8-14 days. The fact that the ^ remains attached to
the host but for brief periods explains why it has not as yet been
observed to attack the host under other than experimental conditions.

Site of attachment on the host. Kossel and his colleagues (1903,
p. 39) state that all stages of the tick may be observed on cattle. The
larvae and nymphs attack the beast about the head on the muzzle,
eyelids and ears, and likewise on the udder. When placed experimentally
upon cattle they attack any part of the host's body. Adult ticks attach
themselves chiefly on the flanks, inside the legs, on the neck, udder, and
in the pit about the anus and vulva. Males are frequently found
running about upon the skin, but they have not been found sucking
blood from cattle. They are frec^uently found 171 coitu with attached

20—2

300 Genus Ixodes

females. I have found these ticks attached in similar situations on
deer and roe-deer. On dogs they occur frequently about the head
and neck. They attach themselves to guinea-pigs and rabbits inside
and outside the ears and upon the eyelids and in other situations.
Incidentally, I may mention that I have for several years utilized
hedgehogs for raising experiments with very good results, since the
bristles protect the ticks against injury from the host.

The time required for Metamorphosis.

Hatching of the eggs. A variable period of time elapses between
oviposition and the hatching out of the eggs. According to Chabrier
(1807, p. 366), the larvae issue from the egg after 2-3 weeks, but more
recent observers record a longer period :

Larvae emerge after :
about 6 weeks (Kossel, Schiitz, Weber and Miessner, 1903, p. 40).
„ 7 „ from 17. vi. to 4. viii. (Bertkau, 1881, p. 148).
„ 8 „ „ 25. v. to 19. VII. at 65° F. (Wheler, xii. 1899).
„ 22 „ „ 4. V. to 12. X. (Meek and Smith, cited by Wheler).

„ 36 „ „ IX. to VI. (Ashworth, 1909, p. 133).

As first noted by Gene (1844), the hexapod larva, when it issues from
the egg, is almost transparent. It subsequently grows darker as the
chitin hardens. Having exhausted its reserve food-supply, derived from
the yolk of the egg, it attacks a host and sucks blood, abandoning the
host when replete.

Gorged larvae metamorphose into nymphs after a period which may
be considerably prolonged by low temperature.

Nymphs emerge after :

(in summer) 1 month (Kossel, Schiitz, Weber and Miessner, 1903, p. 40).
(in winter) 5 months ( „ „ „ „ „ „ „ ).

( „ ) 3 „ (in a room ; Nuttall, 1905, v. supra).

As in the case of the larvae, the nymphs darken and harden for
some days after they emerge from the larval skin. They in turn attack
a host and abandon it when replete.

Gorged nymphs metamorphose into adults after a period which like-
wise varies according to temperature :

Adults emerge after :

(in summer) 2 months (Kossel, Schiitz, Weber and Miessner, 1903, pp. 40, 43).

( „ ) 2| „ : 29. V. to 19. vii. (Wheler, xii. 1899).

(in winter) 7 „ (Kossel, Schiitz, Weber and Miessner, 1903, p. 43).

(kept cool) 12 „ : 14. viil. 1899 to 4. viii, 1900 (Wheler, 1902),

I

Biology of 1. ricinus 301

Adults take about 10 days to harden and darken, remaining very quiet
until this is accomplished and they are ready to attack a host (Wheler,
XII. 1899). According to Kossel and his colleagues (1903, p. 40) the
males emerge prior to the females.

Longevity.

The duration of life of ticks, removed from the host, varies gi'eatly
according to the conditions under which they are kept. They are soon
killed by desiccation: thus Wheler, (xii. 1899) found all stages did not
survive over two to three days when placed in a dry empty bottle
They resist cold: Wheler (1899, p. 7) found that females survived after
exposure to "several degrees of frost"; and Kossel and his colleagues
(1903, p. 43) convinced themselves that all stages survived exposure to
the rigours of a German winter. The fact that the different stages
hibernate accounts for the occasional occurrence of piroplasmosis amongst
stall-fed cattle in winter, for bedding containing pathogenic ticks may
at times be brought into cattle-sheds from tick-infested places. All
stages may survive unfed for a considerable length of time in bottles
containing dampened moss, filter paper, sand, or earth, as shown by the
following records.

Larvae (unfed) hatched 4. viii. 1898 were alive and active on 7. vi. 1899 :
101 months (Wheler 1899, p. 50).
survived 19 months (Wheler, 1902).

,, for months (Kossel, Schiitz, Weber and Miessner, 1903, p. 40).

Nymphs (unfed) survived 5 to 7 months (Nuttall, 1905, v. supra).
„ „ „ 18 „ (Wheler, 1902).

Adults (unfed) survived 15 to 27 months, having been derived from nymphs which
took 12 months to undergo metamorphosis into adults (Wheler, 1902).
partially-fed females, taken from a dog, survived one to three months, and
two males placed in a vessel with chei-ries, died in 50 days (Galli-Valerio,
1908, p. 611).

The time required to complete the Life-cycle.

Wheler (xii. 1899) says that this species will, in fine, warm seasons,
probably pass through at least two stages of its life in one year, but
that in cold weather and when there is delay in finding a host, one
stage may occupy a whole season. The term of its whole existence may,
therefore, vary from 1^ to 3 years, and will in the majority of instances

302 Genus Ixodes

last 1^ to 2 years. It seems, however, possible that a whole generation
may be evolved in one season under very favourable conditions.

Allowiug only ten days for a tick in the larval and pupal stages in
which to harden after metamorphosis and to find a host, and 11 weeks
for metamorphosis thereafter, Wheler calculates the time occupied as
follows :

Larva 13 weeks (1^ free, \ on host, 11 gorged).

Nymph 13 „ (U „ I „ 11 „ )•

Adult ? 7^ „ (U „ 2 „ 4 „ ).

In one case the larvae fasted 40 weeks, and allowing 11 weeks for
metamorphosis after an assumed feeding, practically a whole year would
be accounted for. According to Kossel and his colleagues (1903, p. 43)
it takes at least live months for a generation to develop ; our estimate,
based on the following figures, gives the shortest period at about six
months.

On the basis of the observations here recorded, I would calculate
the time it takes for a generation to develop on a warm-blooded host
as follows :

Days

required

Shortest period

Longest period

8

27

42

49-252

10

300-570

3

6

Fecundated ? abandons the host.

¥ begins to oviposit after

Larvae hatch out from the eggs after

Larvae harden and wait to attack host No. 1
after

Larvae remain upon the host for ...

(Larvae, having abandoned the host, undergo
metamorphosis. )

Nymphs issue from the larval skin after ... 28 84-140

Nymphs harden and wait to attack host

No. 2 after 10 200-540

Nymphs remain upon the host for... ... 3 5

(Nymphs, liaving abandoned tlie liost, under-
go metamorphosis.)

Adults issue from the nymphal skiu after 50 210-360

Adults harden and wait to attack host No. 3

after 10 450-810

Adults ( ? s) remain upon the host for ... 8 14

Fecundated ? abandons the host

Total = 178 days. 1345-2724

About 170 days probably represents the shortest possible time
required for the completion of the life-cycle, if we deduct a few days
from the usual time which appears to be necessary for hardening after

Biology of 1. ricinns 303

ecdysis at each stage. It is quite clear that the time required may be
very much prolonged under unfavourable conditions, and it is probable
that in nature it is not infrequently prolonged to two and even three
years. It should be noted that after very prolonged fasting we have
frequently found various species of ticks too feeble to feed upon a host.

Influence of Season and Local Conditions, etc.

According to Kossel, Schutz, Weber and Miessner (1903, p. 39),
ricinus, in Germany, occurs chiefly on damp water-logged land with
rank vegetation comprised of long grass and bushes. Larvae and
nymphs are found in large numbers on cattle, especially in May and
June; few ticks in these stages are found on cattle at other seasons.
The nymphs are the more commonly found throughout the year, and
are more prone to attack man than the other stages. In the spring
the larvae are encountered in masses on the ends of grass or twigs upon
which they have climbed from the ground where the eggs were deposited
by the female. Larvae, nymphs and females may be captured in a free
state at the end of April, proving tiiat all stages may hibernate. They
are readily captured by dragging a woollen clotii over infested grass and
bushes. Samson (1909, p. 185) netted all stages from high grass along
paths in damp woods about Berlin in the spring of the year. Kossel
and his colleagues state that they are readily captured in the woods
during the spring to autumn ; they are so numerous in some localities
that when the cows return from pasture in the evening the farmers
collect the ticks by the litre and feed them to chickens.

Copulation.

A survey of the literature shows that the process of copulation (or
what I shall by preference term coupling^) in /. ricinus was observed
at a very early date. De Geer (1778, p. 104) was the first to describe
and figure the sexes coupled. In de Geer's figure the male is shown
fixed with its venter upon the venter of the female, with its palps widely
separated, and its chelicerae and hypostome deeply implanted in the
female genital orifice, or vulva. Mtiller (1817, p. 278) also observed
coupling in captive specimens, and stated that the sexes might remain
thus united for a week. Gene (1844, p. 771) observed coupling during

1 The introduction of the s mouthparts into the vulva, hereinafter described, con-
stitutes only a stage in the process of copulation ; see pp. 318 et seq.

304 Genus Ixodes

May to October. He noted that the vulva appeared open in some
females, whilst in others it appeared closed " as by a hymen," these
being presumably virgins. He placed males with the latter and
observed them enter into copulation. On turning the males over he
saw that nought but the mouthparts had penetrated the vulva. He
states that he actually saw three males coupling at once with one
female : " fatto del quale non trovasi altri esempio negli annali della
scienza^" Gene (p. 774), on closely observing coupling males, several
times saw " two fusiform bodies, turgid and pearly white, protruding
one on the right the other on the left of the median line close to the
hypostome," When the male was disturbed these organs collapsed and
were retracted into the interior of the mouthparts ; they could be clearly
seen when they were slowly retracted. He concluded that these must
be fecundating organs, but could not explain their mechanism, (We
shall see that Lewis has also observed these organs.) C. L. Koch
(1885-1844, p. 218) also expressed the belief that the male copulating
organs are situated in the mouthparts.

Von Siebold's (1850) statement that two fine ducts lead forward from
the testes to the mouthparts of the male has never been confirmed.
Pagenstecher (1861, p. 38), who described the male sexual organs,
believed that the seminal fluid escapes from the male genital orifice
when the sexes are apposed venter to venter, the genital orifices being
approximated in some way for the purpose. Me'gnin (1878, p. xcii)
affirms that the males possess a penis (!) which is directed toward the
vulva by the male mouthparts. Glaus (1876, p. 574 and 1880, p. 652)
wrongly states that the male capitulum is bent backward when coupling
takes place, the reverse being the case. Bertkau (1881, p. 147) observed
coupling, and he states that he found spermatozoa in all the females
which had coupled under natural conditions or in captivity. Most of
the females, on the other hand, which were found in a free state were
found not to be fertilized. Bertkau holds that coupling constitutes
copulation but does not explain how this can be the case. Wheler
(1899, p. 7) states that he observed two pairs couple in a bottle after
they had been captured on rushes. He also saw a male coupling with
a gorged female after the specimens had been taken from a deer. He
adds (xii. 1899) that coupling takes place between imfed adults and
between replete females and males either upon the host or otherwise.
He considered that the female became impregnated through the male

^ No subsequent author records any similar observations ; two of the males were,
perhaps, only "crowding" the third. (See PI. VII, p. 310.)

Biology of I. ricirms

305

mouth parts. Wheler sent specimens to Lewis (1900, p. 882) to study.
The latter placed males and females together several times and
repeatedly saw coupling occur. The male crept upon the venter of the
female and probed about with its mouthparts, the palps being in their
normal position whilst being passed to and fro across the vulva " until
at length the female, which had remained hitherto entirely passive,
began to show signs of responding : a slight, but very distinct alteration
in colour of the surrounding parts took place, this being shortly followed
by a relaxation of the peculiar crescent-shaped, flap-like opening. The
palpi of the male were now immediately separated as widely as possible,
the extremity of the rostrum was inserted under the flap and gradually
pushed home until the entire organ was buried to its base." This union
lasted some hours. Lewis gives figures of the sexes thus united, and of
the $ and ^ hypostomes in ventral and lateral aspects.

With regard to the cT hypostome he adds : " A little below the
position of the last marginal teeth there are, however, what appear
in a dry specimen to be two reflexed teeth, one on either side of the

Fig. 283. Hypostome of Ixodes ricinus i in ventral and lateral aspects, showing at
A, A^, the " tubular papillae " described by Lewis (1900, p. 385, PI. XXI, Figs. 4 and 5),

median line, these being of larger size, thicker at the base and longer
but not so sharp, and differing also in apparent structure from any
others of their kind. On forcibly separating a male and female and
examining the rostrum of the former immediately after its withdrawal
from the vulva, I saw at once that these supposed teeth had increased
in size and now presented the appearance of flexible semi-transparent
tubular papillae, which conveyed the impression to my mind that here,
possibly, were the organs by means of which the actual impregnation
took place. I killed this tick without loss of time, and removing the
entire capitulum before it had time to dry or contract, mounted it forth-
with in glycerine. As thus mounted, with no preparation and no

306 Genus Ixodes

pressure other than that of the coverglass, these papillae can still be
seen in their distended condition under a ^-inch objective" (see Fig.
283).

Lewis (21. VI. 1901) quotes a letter from Wheler in which the latter
describes more fully his observations (previously referred to) made upon
ticks placed in a bottle and which entered into union in the manner
already described: " After about an hour I killed one pair with chloro-
form and examined the female, without result. A second pair separated
after a couple of hours. In this case plenty of spermatozoa were present
in the female. There seems no doubt that I had separated the first
pair prematurely. The third pair were still attached after 18 hours."

Neumann (1906, p. 195) says correctly that coupling has actually
been observed in but few species of Ixodes, but that in view of the
homogeneity of the group it has been concluded that it is the rule in
Ixodoidea : " the sperm being removed from the genital pore by the (/•
rostrum and introduced into the $ pore." Neumann notes, ns we have
done, that in no other genus of Ixodoidea are specimens received in
which the sexes have died united and remained coupled in the preserva-
tive fluid in which they were placed. He attributes this to the peculiar
sexual dimorphism affecting the hypostome in Ixodes, meaning doubtless
that the hypostomal teeth in the cTs are especially prominent. I
agree that this appears to be the cause of the prolonged union not
observed in any other genera and of many coupled pairs dying in situ.
I may note, however, that in /. putus ^f the hypostome is practically
unarmed, but that nevertheless the sexes couple as do other Ixodes.
The subject certainly needs further study on living ticks (see further
on pp. 317, 334-345).

We have received many specimens of ricinus in alcohol which have
died coupled, and frequently observed the coupled sexes alive on
different hosts. Unfed and partially fed or replete females in captivity
are promptly sought by the males. Under these conditions a male may
be repeatedly driven away from a female and it will repeatedly return.
At times the male is readily separated from the female ; at other times
it dies in situ. Bertkau's observation, cited above, that females
captured in a free state are rarely fertilized is doubtless explained by
what Wheler (1906, p. 426) states regarding his never having found
unfed females and males coupling under natural conditions upon herbage.
On the other hand, when the unfed sexes were placed in a bottle, which
was warmed by being carried in the pocket, coupling usually took place
and lasted for hours. Samson (1909, p. 186) states that unfed ^fs may

I

Biology of I. riciiius 307

copulate and that $s may copulate several times; moreover that $s
will only attach themselves to a host when fertilized, i.e. during or after
copulation. The last statement requires confirmation. This author
(p. 217) denies that the penetration of the (/ mouthparts into the %
genital orifice (what we term coupling) represents coitus. On examining
a $ which had been coupled for an hour with a cT, her goiioduct was
found to be empty and no spermatophore was found in the ^Z* excretory
ducts. That coitus should take place by means of the mouthparts of
the f^ is difficult to understand because {a) the cT mouthparts cannot be
brought in contact with the (/" genital orifice so as to receive the sperm
therefrom ; (6) the structure of the (^ mouthparts shows no adaptation
for this function ; and (c) the ^T mouthparts are immediately introduced
into the vulva when the sexes meet. On one occasion what appeared to
he true copulation was observed : a coupled pair had been observed for
upwards of an hour when the </ was seen to suddenly withdraw his
mouthparts from the vulva and advance his body along that of the % ,
gripping her anterior pairs of legs and body, thus bringing the sexual
orifices into apposition. The (/* remained about two minutes in this
position, then retreated to his original position and pushed his mouth-
parts forward to the anterior border of the vulva " as if he were pushing
something in." The movement was repeated about 20 times, after
which he again introduced his mouthparts for about 15 minutes and then
finally abandoned the female. Samson assumes that the (/ emitted a
spermatophore when the genital orifices were apposed, and that it pushed
it home with its mouthparts so that it should not slip out again.
Coupling, therefore, would only serve to keep the sexes together,
the introduction of the ^ mouthparts serving to enlarge the $ parts
for the reception of the spermatophore. Samson does not state that
the % was subsequently examined for the presence of a spermatophore
which, unfortunately, robs the observation of a considerable part of its
value.

Samson does not refer to the collapsible organs situated near the
base of the (^ hypostome and which were described by Gen^ and
especially by Lewis {vide supra). We shall seize the earliest opportunity
that offers to study these organs in living ricinus in the hope of being
able to determine their function.

It is obvious from the foregoing that the process of copulation in
ticks requires further study. We prefer the term coupling to that of
copulation when referring to the condition when males attach themselves
to females by introducing their mouthparts into the vulva, this in the

308 Genus Ixodes

light of the evidence above presented. It is established that the male
fertilizes the female by means of spermatophores which are received into
the spermatheca, and from this point the spermatozoa pass up into the
ovaries and fertilize the eggs (Bertkau, 1881, p. 147; Samson, 1909,
p. 216, and ourselves). The entrance of the spermatophores would
necessarily be faciHtated by the dilatation of the female sexual orifice
brought about by the entrance into it of the male mouthparts. It is
interesting to mention, in this connection, that we once observed a male
Ixodes tenuirostris with its mouthparts deeply embedded (as in coupling)
in the body of the female to (me side of the anus outside the genital
groove. It is conceivable that the female had received some injury at
this point and that the male had mistaken the wound for the vulva.

Note : We append a description of the process of copulation in
Ornithodorus moubata as recently observed by Nuttall and Merriman,
since it bears directly on this subject. See Appendix I, p. 318.

Oviposition.

Frisch (1724, p. 42, judging from a footnote in Fischer's translation
of Gend (1849, p. 17), wherein Frisch is quoted verbatim) appears to
have been the first to observe oviposition in Ixodes, the " Hunds-Laus '
as he calls it. Frisch wrote , " Das Geburtsglied ist nicht ganz unten
am Bauch, sondern etwas gegen die Mitte desselben. Es lasst das
Weiblein im Eierlegen zugleich einen hellen Saft aus dem Maul (!) womit
es die Eier am Leibe behalten, und bis an das Maul hinaufziehen, ja
theils gar iiber dasselbe auf den Halsschild briugen und in einem
Klumpen beisamen behalten kann, in denen es also steckte und darinnen
starb." Chabrier (1807, p. 367), who observed oviposition in a pinned
$ ricinus, considered that the eggs were extruded through the oral
aperture and it is not difficult to understand how the mistake arose in
view of what actually takes place. He cites (p. 368) a communication
from Kalm, who observed oviposition in "Ixodes amei^icanus" and
oelieved the eggs issued from the edge of the scutum. Chabrier, who
subsequently examined this species, convinced himself that there was
no opening at the edge of the scutum from which eggs could issue.
Miiller (1817, p. 278) likewise observed oviposition in a pinned specimen
of ricinus. The tick began to lay eggs 44 days after it was collected.
He saw the ovipositor protrude eggs.

The process of oviposition in /. ricinus was first clearly observed
and fully described by Gene (1844, pp. 767-777). He described how

Biolof/i/ of /. riclnus

309

the vagina is everted and protruded forward and the Qg^ issuing there-
from is received by a delicate bilobed, bladder-like organ which is pro-
truded over the depressed capitulum from beneath the scutum. Each
heart-shaped lobe possesses a finger-like protrusion anteriorly on the
inner side. This organ, to which Nuttall (1908, p. 398) gave the name
of Genes organ, in honour of its discoverer, completely envelops the egg
as it emerges from the everted vagina. According to Gene it takes

Fig. 284.

crvU::, a ■--CIn'

Fig. 286.

Fig. 287.

»<U^^

Figs. 284-287. Oviposition in Ixodes ricinus (Wbeler, 1899, Fig. 20 ; also 1906, PI. V,
Fig. 3). We reproduce these figures from the author's original blocks as they very well
illustrate the successive stages of the process. In Fig. 284 the capitulum (" head ") is bent
downward to meet the advancing ovipositor. In Fig. 285 the palps are separated ; the
hypostome is hidden behind the ovipositor which has advanced still further and is pro-
truding an egg ; Genu's organ (" membrane") is protruded to receive the egg as it emerges
from the ovipositor. Fig. 286 shows Genu's organ enveloping the egg between its lobes.
In Fig. 287 the ovipositor is being withdrawn and the hypostome appears ; Gene's organ
is collapsing and carrying along the egg toward the basis capituli. These are the only
figures hitherto published adequately illustrating the process of oviposition in ricimis.

310 Genus Ixodes

about 40 seconds for each egg to issue from the female. After the egg
issues, the vagina is retracted within the body and Gene's organ is also
retracted. Gene records that he once observed the process of oviposition
in the case of a female whose capitulum had been torn off when the
tick was removed from a dog. To study the process, he placed the
ticks upon their backs and cut off their legs so as to render them
immovable. The female, according to Gene, at first depresses the
capitulum so that it is applied to the ventral surface of the body. The
palps and chelicerae being divaricated, Gene's organ is then alternately
protruded and retracted "in a hesitating manner," after which it is
rapidly protruded to its full extent, hiding the capitulum whilst it
receives the e^^g from the protruded vagina, or ovipositor. Gene's organ
is quite clearly stated to emit a sticky secretion upon the surface of the
eggs whilst it overlaps them for a period of four to five minutes. If the
organ is pricked with a needle, oviposition ceases for a while, and when
continued the eggs are laid directly from the oviduct ; but eggs laid in
this manner are not glossy and they soon dry up and become shrivelled.
Gene supposed that, in addition to coveting the eggs with the sticky
secretion, which may serve to harden tiie chitinous envelope of the egg,
that the organ may convey spermatozoa in some way from the female's
uterus to the e^g as she is laying. He did not, however, pretend to explain
how fertilization actually takes placed The ovipositor and Gene's organ
having been retracted, the egg is left lying upon the capitulum, which
now resumes its normal position and shovels the egg backward on to
the scutum. This curious process is then repeated as long as the tick
continues to oviposit; the newly-laid eggs push the older ones backward
in a heap upon the dorsum of the tick, the anterior part of the tick thus
gradually disappearing in the heap of eggs.

Bertkau (1881, p. 148) repeated the experiment of pricking Gene's
organ, and confirmed the observation that eggs laid without being covered
by its secretion subsequently shrivel up. This result was also obtained

^ Eailliet (1895, p. 706, Fig. 480) reproduces a figure of an ovipositing ricinus from
Gen6 (1848), and briefly refers to the process and to the everted vagina or oviduct serving as
an ovipositor or " oviscapte "; be refers to Genu's organ as the "bourse seminale de Gen6,"
or " vessie bilobee." Wheler (1899) calls it the " dorsal gland." Samson (1909, p. 223)
calls it the "Subscutalblase." Button and Todd (1905 b, p. 17) describe and figure the same
structure in Orniihodorus moubata as a " pulmonary sac" and Brumpt (1910, p. 523, Fig.
3<i3) calls it the " cephalic gland," following Christophers (1906, p. 95), who gave this name
to the structure in 0. savignyi. None of these names adequately describes Gene's organ.
We give pre-eminence to Genu's observations because they have been generally ignored by
writers on the subject.

TICKS

PLATE VII

Ixodes ricimts. Numerous females ovipositing in a glass dish at room temperature.
4th week ; most of the females appear shrivelled and smaller than the masses of
eggs they have laid. xf. (Original, L. E. Robinson, phot.)

Biologt/ of J. richuis 311

when Geue's organ was not pricked but simply made to retract itself by
being touched with a small rod. The Q^^^ which then appeared had dried
up in 6-8 hours, whilst eggs laid normally remained fresh for weeks ^
Whcler (1899, p. 110, Fig. 20, republished 1906, PI. V, Fig. 3, here
reproduced — see Figs. 284-287) has described and also figured oviposition
in ricinus. He notes that in unfed or partly fed females the capitulum
protrudes forward but that in fully gorged females it is depressed. It
may, in fact, point slightly backward, especially when they arc ovipositing.
Wheler's description agrees with those of Gene and Bertkau which were
unknown to him. The female he observed took three minutes to lay an
eg^, after which there occurred a rest of six minutes before the process
was repeated. Samson (1909, pp. 221-223, Fig. 14) also describes the
process in the same manner as Gene. She states that the egg is turned
to and fro for about two minutes by Gene's organ, and she describes
the internal glandular structures which are connected with the latter.

Regarding the duration of oviposition and the number of eggs laid by
Ixodes 7'icinus, we glean the following notes on the subject from the
literature :

Gen^ (1844, p. 775) states that oviposition lasts 10-30 days, during
which time a female may lay 1000 to 3000 eggs. Oviposition ceases
when the female is disturbed.

Bertkau (1881, p. 147) observed a gorged $, which dropped of her
own accord from a hedgehog on May 28th ; by June 9th she had laid 211
eggs. She then continued laying until June 20tli, when she was killed,
having laid 847 eggs.

Wheler (xii. 1899) observed a $, collected on April loth, which
began to oviposit on May 12th (interval of 27 days). As in other ticks
we have observed, the $ 's body at this time showed yellowish streaks
(due to the increased excrement contained in its malpighian glands).
This $ laid 2050 eggs.

Kossel, Schiitz, Weber and Miessner (1903, p. 40) state that oviposi-
tion commences 8 days after the replete fertilized female has dropped from
the host, and that it lasts 8-14 days, during which period 100 to 1000
or more eggs may be laid.

Galli- Valerie (1908, p. 611, with figure) illustrates, by means of a

^ Schlechtendal (1891, p. 11) gives a very imperfect description of oviposition observed
in the ease of a female of undetermined species, belonging apparently to the Ixodidae,
which was received in a consignment of fossils from Persia. He observed the protrusion
and retraction of Genu's organ and thought it might be a glandular structure.

312 Genus Ixodes

photograph, a number of females which have oviposited in a dish. He
states that oviposition lasts one month.

Samson (1909, p. 221) states that oviposition commences about 14
days after the $ has abandoned the host.

Note: As stated by Nuttall (1908), the process of oviposition in the
other genera of Ixodoidea is essentially similar to that in Ixodes, having
been observed by him in Argas, Ornithodorus, Ixodes, Amblyomma,
Boophilus and especially in Haemaphysalis, the process being illustrated
by a series of sketches in the case of Haemaphysalis punctata. Lewis
(1892) has illustrated the process in Amblyomma, likewise by a series
of sketches. Samson (1909, p. 223) notes the fact that the process is
similar in Rhipicephalus, but this author's description is very imperfect.
As all the other Ixodidae possess a similar anatomical structure to the
foregoing, it is safe to conclude that they also oviposit iv a similar
manner.

Relation to Disease.

Redwater in Cattle : Ixodes ricinus is the only species of Ixodes of
which we have a conclusive demonstration that it plays a part in the
spread of disease. This tick appears to be the chief carrier of redwater
(piroplasmosis) in cattle in northern Europe. Its association with red-
water was first observed in Finland by Kossel and Weber, in 1910, and
subsequently Kossel, Schiitz, Weber and Miessner (1903) demonstrated
experimentally that bovine piroplasmosis could be conveyed through
ricinus. These authors (pp. 50 et seq.) removed engorged and fecundated
ricinus from German cattle suffering from the disease, and collecting
the larvae that had hatched out from the eggs laid by the infected
females, placed them upon susceptible cattle. Kossel and his colleagues
carried out six experiments with larval ricinus, and they record that
the cattle developed redwater 8-14 days after they had been infested
with the larval ticks. In one of these experiments, carried out in
June, 1902, the infective larvae were derived from eggs laid by females
collected from cattle in the previous year (May-June, 1901). The ticks
had been exposed to all the rigours of a German winter. Kossel and
his colleagues also record an experiment in which a cow was successfully
infected through the agency of ricinus nymphs which had fed on infected
cattle in the larval stage.

It is probable that ricinus is the carrier of redwater in Norway, and
it has been found associated with the disease in Great Britain and

Biology of I. rlcinus 313

Ireland. Kossel found ricinus in N. Wales in redwatcr districts, and
specimens have reached me from various redwater localities in Ireland
and England, having been removed, in some instances, from cattle
suffering from piroplasmosis. Stockman (1908) also notes the associa-
tion of this disease with the presence of ricinus in England.

Note: Schaudinn (1904, p. 428) records in three lines that a blood
parasite of lizards {Karyolysus) develops in a mite ("Milbe"). Doflein
(1909, p. 331), Blanchard (1909) and Christophers (1906, 1907) quote
Schaudinn as having observed the development oi Karyolysus lacertarum
in /. ricinus and that the tick transmits the parasite. None of these
authors gives a reference. Prof. Doflein, who referred me to the passage
from Schaudinn above cited, kindly informs me that he believes
Schaudinn told him his "Milbe" was ricinus. In view of Schau-
dinn's death it will be necessary to repeat the investigations he refers
to quite casually in his paper since we can draw no conclusions from
the little he says on the subject.

Effects of the bite of Ixodes ricinus : The following cases of injury
consequent upon the bite of this tick are recorded in the literature
which we have sifted. Perroncito (1901, p. 562) cites earlier authors
like Ercolani, de Geer and Dubini as stating that psoriasis-like eruptions,
or' gangrenous pustules may follow upon the bites of ricinus. It is
possible, however, that I. hexagonus or /. canisuga are also concerned,
as earlier determinations of species are open to doubt. We add the
following instances : Dubreuilh (1838) cites several cases of phlegmonous
inflammation following ricinus bites in man. Raspail (1840), Cosson
(1856), Moquin-Tandon (1862), Despres (1867) and Liegois all record
ill effects of ricinus bites in man. Sangalli (1884) records the case of
a boy who was bitten on the scrotum and soon after suffered violent
pain with swelling, heat and redness of the part. According to
Raymondaud, of Limoges, grave effects may follow the bite. Johannessen
(1885) describes a case in a boy where the tick's body was removed but
the capitulum remained embedded in the skin at the back of the head.
Swelling followed at the point of injury, accompanied by headache,
stiffness and cramp in the muscles of the one side, partial loss of memory,
and polyuria; the pupils became dilated, etc. The boy made a slow
recovery. Blanchard (1891, p. 689) states that accidents of a grave
character occasionally follow ricinus bite, the wound serving as a centre
from which infection may spread to the rest of the body. Mauvezin (cited

21

314 Genus Ixodes

by Railliet, 1895) says the bite may produce gangrenous inflammation
in sheep, whilst in man the bite may be followed by abscess, oedema,
lymphangitis, etc., accompanied with febrile symptoms. Cao (1898) made
a few experiments with ricinus and B. anthracis and Staphylococcus. He
removed these ticks from the ear of a small dog and inoculated their
contents, with negative results, into another dog, as well as into a
rabbit. He then injected a mixed culture of Staphylococcus pyogenes
aureus and alhus into the jugular vein of the same dog and removed the
ticks at intervals of 12-36 hours, making plate cultures of their contents.
He found cocci in the ticks after 12 hours had elapsed since the injection.
After 16 hours they were very numerous ; after 20 hours less so ; after 24
hours they had disappeared from within the ticks. In a second experi-
ment he injected B. anthracis into the jugular vein of a dog and removed
the ticks at intervals of 4-48 hours, plating their contents. His results
in this case were negative, that is, no anthrax bacilli grew upon the
plates. It is evident from the first experiment that bacteria circulating
in the blood may gain access to the body of the tick, but there is nothing
to prove that ticks are capable of communicating a bacterial infection.
M^gnin (1892, pp. 26-28) denied that any ill effects followed the bites
of ricinus. Although this is incorrect, it is certainly the rule that
little injury follows the bite, and a tick may hang on for days without
being perceived.

Penetration of Ixodes ricinus beneath the skin : For reasons which we
do not at present understand the tick in the larval, nymphal or adult
( $ ) stage may occasionally penetrate beneath the skin of the host
and cause local injury. A number of instances are recorded in the
widely scattered literature, as follows :

Dubreuilh (1838, cited by Nuttall, 1899) reported the presence of
ricinus in a pustule in the mastoid region in man. Trillebert (1863,
Rec. veterin., cited by Megnin, 1892) observed a cyst at the end of a
dog's ear which contained a tick supposed to be riciiius. Van Beneden
(1883, p. 142) recorded the penetration of (?) ricinus beneath the skin
in man and regarded it as a normal occurrence. Aurivillius (1886 b,
p. 139) records the presence of ricvius beneath the skin of a fox.
Blanchard (1891, p. 689) describes a much-quoted case of a tumour of
the size of a nut on a man's abdomen. The tumour had the feel of a
sebaceous cyst, there was no external lesion of the skin to be seen
and the tumour had existed some weeks. On opening the tumour a
female ricinus 8 mm. in length and in a living condition was discovered.

Biologji of T. angustus 315

H, Beauregard is quoted as having once observed a similar tumour
beneath a man's clavicle, and Dr Choupe has seen several such cases in
his practice. According to Blanchard nymphs occasionally penetrate
beneath the skin of horses, producing furunculosis. He is doubtless
referring to the case reported by Megnin (1867, Rec. ve'tenw., redescribed
1892, p. 56 ; in the earlier paper he referred the ticks to a new species :
" Ixode penetrant"). In this case a horse at Versailles was suffering
from numerous pustules on the legs. These pustules are stated to have
been due to ricinus nymphs ; a nymph was extracted from beneath the
scab on each pustule after which the horse recovered. Megnin (1892,
p. 62) attributes the penetration of this tick beneath the skin to the
structure of its mouthparts. Kossel, Schiitz, Weber and Miessner
(1903, p. 40) state that the nymphs and larvae occasionally bore them-
selves beneath the skin of cattle.

The penetration of ticks beneath the skin is certainly an unusual
proceeding. It is interesting to note, especially in the case reported by
Blanchard, that a tick may survive being embedded beneath the skin
for a considerable period of time.

Ixodes ricinus var. scapularis.

Hooker (1908 a, p. 43), in the United States, records that he has
observed the sexes of this species coupling both on and off the host.
He writes : " An unengorged, unattached $ , taken in the field from a
hunting dog and placed in a pill-box with unattached ^/'s taken from
the same dog, was shortly after found coupling witli one of the latter.
From this it would appear that it is unnecessary, for the % of this
species at least, to take food prior to fertilization."

Ixodes angustus.

According to personal communications received from Dr S. Hadwen,
this species occurs in all stages upon squirrels at Duncan's, in British
Columbia. The j/s are scarce. He succeeded in raising adults from
nymphs captured on squirrels by transferring them to rabbits in
captivity and allowing them to feed upon the rabbits.

Coupling has been repeatedly observed by Dr Hadwen. In one case
it lasted 15 minutes after the ticks were placed together in a tube.
When the </• withdrew his mouthparts from the vulva the vagina was
seen to be slightly everted. Slight pressure applied subsequently to the

21—2

316 Genus Ixodes

$ caused the vagina to be completely everted. After the </* had freed
himself he immediately attached himself to a second $. He rooted
around "like a pig" for 20 minutes before he succeeded in inserting his
mouthparts into the vulva. This second coupling lasted 10 minutes.

Dr Hadwen only once detected a cT beneath a $ upon a host
but the sexes were not coupled. He has only captured 8 or 9 (/"s
and believes that copulation takes place mainly off the hosts, mostly
in the squirrels' nests, "for the squirrels are infested with almost an
equal number of ticks the year round."

Ixodes pilosus.

Relation to Disease: According to C. W. Mally (ix. 1904), Cape of
Good Hope, the farmers around Carlisle Bridge have no doubt but that
this tick produces " paralysis " in sheep, especially in merinos. Cooper's
dip applied to " paralysis flocks " is stated to check the disease at once.
The tick is, however, frequently found on healthy sheep.

Behaviour of ^ and $ on the host : Mally states that the $ is sought
by the </. The $ prefers to attach herself on sheep "around the
mouth, under the chin and throat, on the bare patches under the legs
and along the edges of the wool, in the wool on the legs, along the belly
and near the udder," The ^ appeared to be ready for the (/• after being
attached to the host for two days, but females can wait indefinitely in
this situation whilst they are gradually becoming replete. After the
replete females dropped from the host they died readily on being sub-
jected to transportation. The ^s are very scarce, and when not
attached to the $ are usually found wandering about. Males were
twice found attached to the skin. Coupling was observed to last from
3^ to 24 hours.

Nymphs and Lar^vae were found engorged in the ears of sheep, but
they were scarce (Mally).

Ixodes canisuga.

In contrast to ricinus, the males of canisuga have never been
recorded as occurring upon the host. Whereas hundreds of females and
nymphs have reached us, we never received a male taken from an
animal. The only male which Mr E. G. Wheler ever saw came from a
dog-kennel at Govvanburn, on the North Tyne, and all the males we
otherwise possess have come from the nests of the sand-martin or bank-

Biology of T. piitiis 317

swallow. Mr Wheler kindly informs us that canisuga swarms on collie
dogs throughout the North Tyne district and probably throughout the
borderland, and adds : " This is partly accounted for by the fact that
the dog-kennel is one of the chief features of a hill farm, and the dogs
all lie together, and there is every opportunity for the ticks to multiply."
Mr Wheler was the first to observe copulation in this species, to which
he provisionally gave the name of/, pluiubeus ? in 1899.

Through the courtesy of Mr. R. F. L. Burton, of Longner Hall,
Shrewsbury, who kindly supplied me witli many living specimens
(including some 16 males), I was able to observe that copulation took
place readily in captivity and that it occurred as in i^icinus. The males
sought the females, the latter being both unfed and in various stages of
repletion. It is obvious that collectors desiring to obtain males will
have to search the habitats of the hosts if they wish to secure them.

Wheler (xii. 1899) kept the unfed larvae of this species alive for
ten months (9. x. 1898-vili. 1899), after they had hatched out from
eggs laid 4. vili. 1898. The larvae were confined in a bottle containing
moss and damp sand.

Ixodes putus.

This species, whilst apparently confined to marine birds as hosts, is
knovn to occasionally attack man and inflict painful bites. The males
have never been found upon the host, and the almost unarmed hypo-
stome of the male indicates that it may perhaps not suck blood, behaving
similarly to Ornitkodorus megnini in this respect. The males have only
been found in the nests of the birds or in their vicinity. Copulation in
this species has been observed both by Wheler (190G, p. 425) and
Hewitt ; it takes place as in ricinus. I am indebted to Mr Wheler for
allowing me to see a letter from Mr W. Hewitt regarding the latter's
observations on putus in June-July, 1902. He found the ticks plenti-
fully on the cliffs at Bempton and Buckton, in Yorkshire. He found
two or three pairs in copula beneath stones, and once saw a male in
copula, whilst four or five near by were seeking to copulate ; this on a
narrow ledge of cliff 320-400 feet high, facing the sea, and frequented
during the nesting season by tens of thousands of birds (guillemots,
razorbills, puffins, jackdaws, with a few herring-gulls and kittiwakes).

318 [39]

APPENDIX I

[Reprinted from Parasitology, Vol. IV, No. 1, pp. 39-44, March, 1911.]

THE PROCESS OF COPULATION IN
ORNITHODORUS MOUBATA.

By GEORGE H. F. NUTTALL, F.R.S. and
GORDON MERRIMAN.

Although the literature on ticks is very extensive, we have failed
to find any satisfactory description therein of the process of copulation.
Most authors state that the male mouthparts are introduced into the
female genital orifice, or vulva, and that this constitutes copulation,
leaving it to the reader's imagination to determine how the seminal
secretion gains access to the female generative organs. The mechanism
of copulation, in other words, remains to be described, and we propose,
in this paper, to give an account of what we have observed in Ornitho-
dorus mouhata. We shall describe the process in Ixodidae in a later
paper, but may state here that, as far as our knowledge goes, it is
essentially the same as in mouhata, that is, impregnation takes place by
means of spermatophores.

We have repeatedly observed the process in mouhata in Cambridge,
and the description which follows is based on the study of several pairs
in copulation. When a male and female mouhata, which are ready to
copulate, are placed in a dish, the male creeps about upon the female
and presently seeks to creep beneath her, usually to one side between
the second and third legs. The posterior part of the female's body is
now raised, and the male advances so as to bring his body into line with
the female's, the ventral surfaces of the pair being apposed. The male
clings with his legs to the basal joints of the female's legs, his pair I in

[40] Co^mlation in Ticks 319

front of pair I of the female, and pair II in front of pair II of the female,
and so on. The male all the while feels about with his mouthparts for
the female's genital orifice.

The process that follows can, with careful handling of the ticks, be
most conveniently observed by placing the pair in a pit made with the
end of a pencil in a ball of modelling-wax (plasticine). The ball may
measure an inch or more in diameter, and can be rolled about to the
desired position, being fixed there at any moment by slight pressure
against the glass dish in which it is contained. The female always

Fig. 288 [1]. Ornithodoriis moubata, ? . Showing the emptied portion of the spermato-
phore with the neck-shaped portion attached within the female sexual orifice. Drawn
with the aid of a camera-lucida (G. M. del.).

seeks to regain her natural position upon a horizontal surface, so it is
necessary to immobilize her sufficiently to prevent this without hamper-
ing the male. This can be done by modelling the wax about the edge
of the pit so that it holds the female in place. Observations on
free pairs shows that this manipulation, if done carefully, in no way
interferes with the process. All of our observations were made with
the aid of a Zeiss binocular microscope which can be inclined at any
desired angle.

The male, having found the female's genital orifice by feeling about
with his mouthparts, immediately introduces all of these (hypostome,

320 ApjMiidix I [41]

chelicerae and palps) together into the orifice. The mouthparts are
pushed in and withdrawn with a sawing motion, the capitulum being
alternately protruded and retracted from his body whilst it is directed
backward almost at a right angle to the long axis of the male's body.
The mouthparts . penetrate more and more deeply but not further
than the basal article of the palps. At each movement of partial
withdrawal the male mouthparts are tilted forward so that friction
against the anterior lip of the vulva takes place. The chelicerae
are all the while being actively protruded and retracted within their
transparent sheaths, the distance which they travel in their movement
to and fro corresponding about to the length of the second palpal
article ; the chelicerae usually move alternately. Presently the male
capitulum is more and more depressed so that the hypostome points
slightly backward whilst the dorsal surface of the basis capituli comes
to be placed on a line traversing the first pair of coxae about midway
along their length when viewed ventrally. (When disturbed at this
stage it is seen that the male and female sexual apertures are in
apposition.) The ventral surface of the female, together with the thick-
lipped vulva, has, up to this, been somewhat prominent, but now the
area about the vulva, and especially posterior to it between the first pair
of coxae, becomes markedly depressed. At the moment when the
female venter becomes depressed the male withdraws his mouthparts
completely from the vulva and at the same time separates his body
somewhat from that of the female to which it has hitherto been closely
applied. The male's capitulum is now directed forward, whilst the palps
and chelicerae are working actively, the digits being moved about
rapidly. (When the sexes are separated at this stage it is found that
the spermatophore is issuing from the male sexual orifice. The
spermatophore, viewed as a whole, is flask-shaped, with a long thin
neck, which is the first part to issue from the orifice ; the tip of the
neck of the flask when it issues adheres to the tip of the male
hypostome. The spermatophore is ejected rapidly and lies along the
venter of the female with the neck directed toward the vulva into
which it is immediately inserted by the hypostome of the male.
A moment later a large amount of clear- watery fluid emerges from
the coxal glands of the male.) When viewed from in front, the ticks
not being disturbed, a large amount of clear fluid now floods the space
between their venters ; the male reintroduces his mouthparts into the
vulva, and the ticks again bring their bodies close together. The male
mouthparts are worked to and fro very much as they were at first, and.

[42] Copulation in Ticks 321

after two or three minutes, they are finally withdrawn and the male
abandons the female.

The ventral surface of the female is found bathed with coxal secre-
tion, the depression about the vulva has disappeared, and the collapsed
spermatophore is found attached to the vulva with its neck pushed into
the vagina. The whole process of copulation lasts about 15 minutes from
the moment that the male inserts his mouthparts for the first time.

The spermatophore, immediately after it has been expelled by the
male, appears as a smooth, glistening, very flexible body completely
filled with slightly milky contents. (On one occasion when the male
was disturbed and the spermatophore was found doubled up on the
venter of the female, the spermatophore literally shot out its long thin
neck when touched with a fine camel's hair brush in the effort to remove
it uninjured.) The end of the neck of the spermatophore appears
closed and rounded, whereas the expanded portion of the spermato-
phore is usually bilobed. When the ticks separate without interference
the spermatophore neck appears shorter and the flask collapsed, and
somewhat spoon-shaped with the concavity on the side where the male's
body had rested upon it whilst in coitu. The neck of the spermatophore
is firmly attached within the vagina and force is required to remove it;
on seizing the spermatophore with a fine forceps the female can he lifted
by it without rupturing the structure. On exerting more force the neck
of the flask tears loose at the point of entrance into the vagina and the
" mouth " of the spermatophore appears spread out and frayed. The full
spermatophore is crowded with " prospermiums " and secretion rich in
granules; when almost collapsed, a few male elements may still be found
therein. The spermatophore, very soon after coitus has taken place,
contracts, flattens and hardens, the edges of the " spoon " remaining
upturned. We possess several females in our collection which have
died or been preserved, with usually one, but at times two or even
three empty spermatophores still adhering. The flask-like covering of
the spermatophore is exceedingly tough, and tested with 10°/o caustic
potash, it is found to be composed of chitin. The freshly expelled
spermatophore, with fully extended neck, measures about 3 to 3"5 mm.
in length and about 1"7 mm. in width at the widest part. When dried
and attached to a female, to whom it may remain attached for some
days, or even for weeks, it measures about V6 mm. in length.

Although there are several points still to be elucidated with regard
to this interesting process, we consider that what we have observed
explains in the main how coitus takes place. The introduction of

322 Appendix I [43]

the male raouthparts necessarily dilates the female sexual orifice and
doubtless excites the female. It is possible that a sticky secretion is given
off by the male mouthparts which serves to attach the neck of the
spermatophore so that its contents are fully voided into the vagina.
The male brings his sexual aperture practically into apposition with that
of the female, and when she is ready for impregnation, he withdraws his
mouthparts and expels the spermatophore which issues with the neck
directed forward so that it adheres to the tip of his hypostome and is
promptly pushed into the vagina. The male removes his venter from
that of the female so as to give space for the spermatophore to pass,
and the female venter at the same time becomes concave for the same
purpose. The flood of secretion from the male coxal glands serves to
prevent the too rapid drying of the chitinous envelope of the spermato-
phore and keeps it highly flexible until its contents have been discharged
— this may be especially necessary in such ticks as 0. mouhata which
live in arid climates. The male reintroduces his mouthparts so as to
" plant " the neck of the spermatophore in the vagina and he appears
to rupture the tip of the spermatophore so that the contents escape
into the spermatheca when the bodies of the ticks are brought together
again closely and press upon the spermatophore, thus pumping out its
contents.

Judging from the fact that we have not as yet observed empty
spermatophores adhering to the external genitalia of female ticks
belonging to other species, we must conclude that in other cases the
spermatophores are introduced in toto into the spermatheca. This may
be the case in 0. savignyi, for Christophers (1906, p. 42) states that he
found the " spermatozoa " (prosperraiums) in this species " included in
cyst-like spermatophores " within the spermatheca of the female. He
figures two such spermatophores of savignyi and also flask-shaped
spermatophores in the spermatheca of Rhipicephalus sanguineus.
Christophers found spermatozoa in the oviducts of 0. savignyi.
Samson (1909, p. 186) has made a suggestive observation on Ixodes
ricinus, which indicates that copulation takes place in an analogous
manner in this species ; she merely states, without giving any particulars,
that the male impregnates the female by means of spermatophores. In
this species, Bertkau (1881, p. 147) and Samson (1909 a, p. 216) have
found that the spermatozoa pass up from the spermatheca into the
ovaries and there fertilize the eggs. We have also observed them in
this situation. Samson (1909 b, p. 495) states that the spermatozoa in
0. mouhata only attain their full development in the ovary.

[44] Copulation in Ticks 323

We have dwelt thus at length upon the process of copulation in
0. moubata, because until now it has never been understood how the
male impregnates the female.

As bearing on the process of copulation in Ixodidae we would add
the following remarks :

We have stated elsewhere that the genus Ixodes (group Prostriata)
is widely separated from the other genera of Ixodidae (group Meta-
striata) in important points of anatomical structure. In Ixodes the anal
grooves surround the anus in front, the sexes show marked dimorphism
in respect to the structure of the hypostome, and in certain species the
sexes are frequently found coupling upon the host. In Ixodes the male
inserts his mouthparts into the female sexual orifice as a part of the
sexual act. In none of the Metastriata has a similar method of coupling
been observed ; here the males remain attached for longer periods to
the host and they do not appear to wander about upon it as do some
species of Ixodes whose males infest the host together with the females.
In Metastriata the females and males are found attached to the host so
that in many cases the venters of the sexes are apposed.

We have discovered that in the Metastriata (including the genera
Haemaphysalis, Dermacentor, Rhipicentor, Rhipicephaliis, Boophilus,
Margaropus, Hyalomma, Amblyomnia and Aponomma) all the males
possess a chitinous apron or flap covering the sexual aperture. The
apron arises from the integument anteriorly to the sexual aperture and
is directed backward over the aperture ; it is frequently serrated along
its rounded posterior border which is free. The apron can be readily
lifted by inserting a fine needle between it and the body surface; it is
very thin at the free margin, whereas it grows gradually thicker toward
its origin. The apron is absent or rudimentary in the females.

Now we find that in Ixodes the males do not show an apron, or it is
very rudimentary in character. On the other hand the apron, though
very thin, may be very well developed in the females ; it is best seen in
young specimens.

The presence of the apron in the males of Metastriata and its absence
in Ixodes is very suggestive, and, taken together with the other dif-
ferences, indicates that copulation may possibly take place in another
manner in the Metastriata to what it does in Ixodes. The mechanism
of copulation in Metastriata still remains to be determined and the
significance of the apron requires to be explained.

324 [46]

APPENDIX II

[Reprinted (with a few slight alterations) from Parasitology, Vol. IV, No. 1,
pp. 46-67, March, 1911.]

ON THE ADAPTATION OF TICKS TO
THE HABITS OF THEIR HOSTS i.

By GEORGE H. F. NUTTALL, F.R.S.

In the course of the extended investigation upon ticks which we
have been conducting for some years, a very large number of specimens
have reached us from all parts of the world, thanks to the generous
aid which we have received from numerous collectors. In addition to
studying this material, the literature on ticks has been sifted for any
information which might throw light upon these parasites. A study of
the data relating to the structure and habits of ticks and of their host
relationships has brought to light certain facts which possess consider-
able interest and consequently appear to me worth recoiding in this
Journal.

The superfamily Ixodoidea is divided into two families, the Argasidae
and Ixodidae, which are distinguished from each other by their external
structure as well as by their habits^

General considerations regar-ding the Biology of the Argasidae

and Ixodidae.

The Argasidae are mostly inhabitants of warm climates. When
they occur in colder parts of the world they seek hosts whose habitats
afford them protection, and insure them possibly a certain degree of

^ Read before the Cambridge Philosophical Society Meeting, 20 February, 1911.
- The main structural differences are described by Nuttall and Warburton (1908) in
Ticks, Part I, p. 1.

[47] Adaptation of Ticks 325

warmth from proximity to animals upon which they feed. Thus,
Argas rejlexus and Aryas vespertilioniH, which occur in colder climates,
are parasitic on pigeons and bats respectively, and they obtain shelter
in pigeon coops and the retreats of bats.

Like all other ticks, they pass through larval and nymphal stages
before attaining maturity. In some Argasidae the larvae suck blood
(as in Ixodidae) ; in others they do not. Where the larvae suck blood
they remain attached to the host for 5-10 days or more (as in Ixodidae).
There are two or more nyiuphal stages in Argasidae (one only in Ixodidae),
and the nymphs, witli one exception, Ornit/iodorus megnini, arc rapid
feeders, that is, they usually feed on blood to repletion within 20-30
minutes (the nymphs of Ixodidae take several days to become gorged).
The adults are not only rapid feeders, but they may feed repeatedly
(not so in $ Ixodidae, though possibly in ^f s).

The nymphs^ and adults of Argasidae are i^ii^ely carried away from
the habitats by their hosts because they feed rapidly. They are mostly
nocturnal feeders, attacking the host when the latter is sleeping or resting.
All species of Argasidae would appear to shun the light, especially when
waiting for a host. When hungry they may leave their hiding places
during the daytime if they become aware of the proximity of a host.
The Argasidae infest the habitat of the host.

The nymphs and adults of Ixodidae behave in a very different
manner. The nymphs attach themselves to the host, and are carried
about for several days, and possibly longer. This likewise holds for the
adults of both sexes in a large number of species. The duration of
their parasitism upon the host may be much prolonged, especially
when the host is a cold-blooded animal; thus, species of Amhlyomma,
Aponomma and Hyalomma, occurring on Reptilia, may remain attached
to the host for many months. On the other hand, we shall see that
there are species of Ixodes in which the males are unknown or in
which they are only recorded as having been found in the habitats
of their hosts.

In the Argasidae the females lay eggs in batches after successive
feeds, and the total number of eggs laid by a female is comparatively
small, say 200. In the Ixodidae, the eggs may number one or more
thousands. The explanation of this difference in tlie number of eggs
laid appears obvious when we consider the life habits of the two
families. The Argasidae are essentially ticks of the habitat in their
nymphaP and adult stages; the females lay their eggs in and about
^ Ornithodorus megnini excepted.

326

Aj)pendix II

[48]

their host's retreat and naturally the offspring have a good chance of
finding the host when their turn for feeding arrives.

With the Ixodidae the case is different : the fertilized and replete
females must frequently drop from the host when it is on its wanderings,
and the offspring may be left stranded in places where they may never
find a host. The loss of life in nature must be enormous in the majority
of species, for in most species the larva, nymph and adult must each seek
a host. The loss of life should be less in those requiring but one host,
as in Boophilus, where the tick attacks the host in the larval stage and
undergoes its metamorphosis up to the adult stage upon a single animal.
The greatest loss of life must of necessity occur in such species as are
parasitic upon hosts having no fixed habitat. In other cases, owing to
the fact that the host returns to a burrow, nest, or the like, it is pro-
bable that there may be less loss of life than in the case where an animal
with wandering habits is chosen as a host. It would be interesting to
make some numerical determinations of the progeny, especially of various
species of Ixodidae, with this point of view in mind.

The large progeny of the Ixodidae secures their survival in nature
in the face of the many difficulties the ticks must encounter in finding
their hosts. In the Argasidae these difficulties are much smaller and
the progeny is proportionately less numerous.

Argasidae.

In the following table I have included all the Argasidae: Argas(A.)
and Ornithodorus (0.) of which we know the host relationships. The
list includes 12 species and 1 variety:

Argasidae
A. persicus

A. reflexus

A. vespertilionis

Hosts and habitat etc. of Tick

Birds : fowls, ducks, geese, turkeys, ostriches,

quail, wild doves, canaries. Infests chicken

coops, etc.
Man : prevalent in Persia where it infests

native dwellings. It also occurs in houses

in Egypt.

Birds : pigeons, fowls.

Man : occasionally entering dwellings from

pigeon coops, or attacking persons having

to do with pigeons.

Bats: several species'; adults found in the
abodes of bats.

Geographical

distribution

(as far as known)

Africa, Asia, Aus-
tralia, N. & S.
America.

Europe, N.Africa.

Europe, N. Africa.

1 See Ticks, Part I, p. 39, where the bats are listed. A variety infests penguins at
Queenstown, Cape Colony.

[49]

Adaptation of Ticks

327

Argasidae
A. hrumpti

0. savigniji
0. moubata

O. turicata

0. talaje

O. talaje var. ca-
pensis

0. pavimentosus
0. tholozani ...
O. lahorensis ...

O. megnini

Hosts and haliitat etc. of Tick

Porcupine: found infestiuf^ dusty burrows of
Hystri.v sp. in Africa.

Man : attacks man when sleeping on the
ground ; hides in dust during the day-time.

Man, camel, horse : attacks them in their rest-
ing places, hides in dust or sand, etc.

Man : infests native dwellings and resting
places along caravan routes.

Domesticated animals : dogs, pigs kept in styes,
goats, sheep.

Man : at times infesting native dwellings ;
attacks at night.

Domesticated animals : pigs in styes, cattle,
llama, horse.

Wild animals : infesting burrows of tortoise
and gopher.

Man : infesting native houses ; attacking at
night.

Birds : found in penguin's and other nests.

Man : resting places infested.

Fowls and camels : infests chicken coops.

Sheep : no particulars as to manner in which
sheep are kept.

Man and domesticated animals : horse, ass,
ox, are all the hosts recorded ; occurring
chietly in the ears.

OeoKraphical

•listribution

(as far as known)

Africa.

Africa, India.

Africa.

N. & S. America.

N. & S. America.

Islands of Cape
Colony, St Paul's
Isl., Siren Isl.

Africa.

Persia.

India.

N. America.

Of the foregoing species the life-histories of only five are known
{A. persicus, A. reflexus, 0. moubata, 0. savignyi, and 0. megnini), the
ticks having been raised experimentally. The various stages of A. ves-
pertilionis are known, and nymphal or adult stages (or both) of some
of the other species are more or less known to science. In A. persicus,
A. reflexus, A. vespertilionis and 0. megnini the larvae attack the host.
In 0. moubata and 0. savignyi, in which the eggs are correspondingly
large, the larva is formed within the &gg, but it is incapable of attacking
a host; the larva in both of these species is inactive, and very soon
after the egg-shell splits and whilst it remains in (moubata) or near
(savignyi) the egg-shell it rapidly undergoes metamorphosis and emerges
from the larval skin as a nymph. It is as a nymph that both the
latter species first attack the host.

328 Ajyj^endix II [50]

The larvae of remain attached to the host for

A.persicxis 5 and 10 days in warm and cool weather respectively (Nuttall).

A. reflexus 7 days (Brumpt, 1910, p. 528).

A. vespertilionis Some days at least (Nuttall) i.

0. megnini 5 days (but they stay on as nymphs upon the host).
The first stage nymphs of

A.persicus lJ-2 hours (Nuttall).

0. mouhata 10 minutes-1 hour (Nuttall).

0. megnini 35-98 days, or more (Hooker) ; abandons the host as a late stage
nymph and does not attack a host as an adult.

The later .stage nymphs
and adults of

A. persicus 5 minutes-2 hours (Nuttall). (Usually less than J hour.)

A. reflexus 20-27 minutes (Alt, Boschulte).

0. mouhata 20 minutes-2 hours (Nuttall).

O. coriaceus 45 minutes-l| hours (Nuttall).

0. turicata Feed more rapidly than the last (Nuttall).

O. tholozani 30 minutes (Megnin).

O. lahoreusi-s 25 minutes-2J hours (Nuttall).

O. savigmji Feed like 0. moubata (Nuttall).

We may divide the ticks above enumerated into three classes,
according to their biology :

Group 1. A. persicus, reflexus, and vespertilionis.

„ 2. 0. mouhata and savignyi.

„ 3. 0. megnini.

Group 1. A. persicus, reflexus and vespertilionis.

In this group the larvae are parasitic upon the host for, say, 5 to
10 days. In persicus, and this probably holds for the two other species,
the nymphs feed about as rapidly as the adults, i.e. in less than
2 hours ; at times in 20-30 minutes. This explains why these stages
are infrequently found upon the host ; they are rapid feeders and
quickly abandon the host when replete. Argas persicus and reflexus
are essentially parasitic on birds. There is no trustworthy record^ of
the larval stages having been found on man or upon other mammalian
hosts. Without denying the possibility of their attacking mammals, it

1 Although I have examined many bats, I have never found A. vespertilionis adults
and nymphs upon them. On the other hand, larvae, in various stages of repletion, were
not infrequently encountered. I conclude from this that the larval stage must remain
attached to the host for some days at least. Doubtless, the duration of parasitism is
influenced by the surrounding temperature.

* It is reported that Starcovici once found reflexus larvae on the horse in Eoumania
{vide Ticks, Part I, p. 27), but I feel warranted in doubting the statement.

[51] Adajytation of Ticks 329

appears reasonable to suppose that when they infest human dwellings
it is because of birds being harboured there. The birds on which the
ticks have been found are nearly all domesticated. The natural hosts
of persicus and refiexus are commonly fowls and pigeons which doubtless
have served to spread the tick amongst other birds in their vicinity.
The thin skin of the bird seems peculiarly adapted for the successful
attack of the small larval tick, and the nesting or roosting habit of the
birds permits the ticks to drop off in situations adapted to the feeding
and other life habits of the nymphs and adults. Both species in their
mature stages are essentially night feeders, that is, they feed when the
birds are in the dark and asleep, and the ticks retreat into crevices and
dark places during the day-time. Where the ticks attack man they do
so at night-time. I assume that 0. talaje var. capensis, which has been
found in penguin's and other birds' nests, will be found to have a
similar life-history to the above ; the larvae should be found on the
birds in localities infested by the tick.

Argas vespertilionis has never been found on other animals than
bats\ Here we have a host which retreats to dark places during the
day-time. The retreats of bats (hollow trees, beneath roofs of houses,
in caves and grottoes) are more or less permanent dwellings where the
ticks find shelter very much as do the Argasids above mentioned which
attack birds. It is clear that to secure the adult stages of this tick the
habitats of bats are the places in which to search for them.

Group 2. 0. mouhata and 0. savignyi.

In this group the larvae are inactive, and the first stage nymph
attacks the host. This nymph feeds as rapidly (in the case of mouhata)
as does the adult, i.e. in 10 minutes to one hour. Ornithodorus mouhata
and savignyi seem to be peculiarly adapted to their hosts and the habits
of their hosts. All of the latter, i.e. man and the domesticated animals
possess thick skins. Man appears to be the chief host, at any rate of
mouhata. The hairless human skin, coupled with man's ability to
remove his ectoparasites, renders prolonged parasitism difficult. Man
herds his animals into a limited space in immediate proximity to his
dwellings or resting places along routes of travel. Both species live in
hot dry climates where there is a largely unclothed human population.
The ground is for long periods of the year covered deeply with a layer
of dry dust or sand both in and out of dwellings.

^ See footnote, p. 326.
N. I. 22

330 A2)2)endix II [52]

Both species, if they had active larvae, would labour under a consider-
able disadvantage under these conditions and an enormous number of
larvae would be lost in the dust or sand, and would die from desiccation.
It may be noted that the females in captivity burrow and lay their
eggs above them on or near the surface of sand. To overcome these
conditions they must be provided with a thick integument to protect
them against desiccation; they must possess mouth-parts capable of pene-
trating a thick skin and of drawing blood rapidly; an increase in size and

12 3 4 5 6 7

Fig. 289 [1-8]. Eggs of Ixodoidea showing their relative size in some species belonging
to different genera. Outlines drawn with a camera-lucida and reduced. The scale
to the right = 1 mm. (Original. G. M. del.) :

1. Ixodes hexagonus 575 x 450 /x. 5. Boophilus decoloratus 625 x 400 /x.

2. Haemaphysalis punctata 570 x 380 fi. 6. Amhlyomma hebraeum 625 x 525 //..

3. Hyalomma aegijptium 550 xi25 fi. 7. Arga.t persicus &25 x 575 fi.

4. Rhipicephalus everts! 550 x 400 fi. 8. Ornithodorus moubata 950 x 750 fi.

strength will render them more capable of dealing with the dust
problem. The larval stage is, therefore, inactive. The females lay
larger eggs (and fewer) than any other ticks, so that they contain
enough reserve upon which the nymphs may develop. The first stage
nymphs are rapid feeders, and very resistant to drought and prolonged
starvation, although somewhat less so than the later stages.

Group 3. 0. megnini.

Ornithodorus megnini is quite aberrant in its habits, and appears
highly specialized in its adaptation to a life of parasitism. As in
Group 1, the larva attacks the host and remains attached thereto for
about five days, by Avhich time it is replete. It moults in situ, and
re-attaches itself to the host for 35-98 days, or more, slowly growing
in size and doubtless moulting as do the nymphs of other Argasidae.
The larvae and nymphs are especially prone to attack the ears of their
hosts. When replete, the nymphs abandon the host, crawl up several
feet on posts, trees, or the like, and hide in chinks and crevices. After
about seven days, in summer, the adults emerge, but they do not seek

[53] Adaptation of Ticks 331

a host. During their long period of parasitism as nymphs they have
doubtless undergone internal changes leading to maturity, otherwise
they would not emerge so rapidly as adults from the nymphal skin.
The adults, without further feeding, mate ; the females oviposit in the
chinks and crevices, and the larvae which emerge from the eggs await
the coming of a host. Hooker, to whom we are indebted for most of
what is known of this curious life-history, very naturally regards the
habits of this tick as correlated with its peculiar form of parasitism in
the ears of its hosts. He does not, however, discuss the subject.

Animals rubbing their ears or heads against infested posts or trees
(" rubbing places ") would necessarily pick up the waiting larvae.
I would add that the structure of the mouth-parts, and especially
the presence of an unarmed hypostome in the adults, indicate that
they can scarcely be blood-suckers. It would be a matter of consider-
able interest, it seems to me, to examine the internal anatomy of the
adults of this species to see in how far it is modified, especially with
regard to the pharyngeal pump, salivary glands and digestive organs.

The life-history of 0. megnini is that of an animal highly specialized
to a life of parasitism chiefly in the ears of its hosts. The larvae issue
from eggs laid on rubbing posts or trees, and the like, some feet above
the surface of the ground, so that they may readily gain access to the
heads of their hosts. The larvae enter the ear unperceived. It would
be a matter of increasing difficulty for succeeding stages to ''cast anchor
in such a small harbour " a second time for, apart from their increasing
size, which would militate against their re-entering the ear, the number
of ticks of later stages which survive even from a large batch of larvae
(in all species of ticks) is very small and grows progressively smaller
toward the period when maturity is reached. The tick, having entered
the ear where it is safe for an unlimited time, takes advantage of the
protection afforded and only leaves it once when it abandons it as
a large nymph almost ready to cast its skin and emerge as an adult.
The period of parasitism as a larva is normal, but that of the nymphal
stages inordinately prolonged so that the tick only needs to go in and
out once through the small aperture of the ear. It is owing to the
choice of the ear for its place of feeding upon the host that the life-
history of this tick has become so peculiarly modified. Its spiny
integument, to which it owes its name of " spinose ear tick," may be
of use in connexion with its parasitism in the ear in lessening its
chances of adhering with a large body-surface to the cerumen within
the ear.

22—2

332

Appendix II

[54]

Of the other species of Argasidae which have been enumerated in
the list on pp. 326, 327, too little is known to permit of much comment.
It would seem to me as if A. br^umpti, 0. turicata and 0. pavunentosus
may prove, on investigation, to have a life-history similar to that of
0. moubata and 0. savignyi ; 0. talaje and 0. tholozani may possess
life-histories like A. persicus, and this appears highly probable for
0. talaje var. capensis. It is to be hoped that the life-history of
0. Iahore7isis will be worked out in India. We are studying 0. turicata
and 0. talaje var. capensis.

Note on the structure of the Hypostomes in Argasidae.

The dentition of the hypostome is in accord with what I have
stated regarding the feeding habits of the Argasidae. This is illus-
trated by typical examples in the accompanying figures. Thus, in
A. persicus, rejiexus and vespertilionis, where the larvae attach them-
selves to the host for some days, or longer, the larval hypostome is

7 ^Wv)

Fig. 2901.

Fig. 291.

Fig. 292.

Fig. 293.

Fig. 294.

Fig. 290. A. persicus, larva (similar in larva of A. rejiexus)

Fig. 291. A. vespertilionis, larva.

Fig. 292. A. persicus, adult.

Fig. 293. A. rejiexus, adult.

Fig. 294. A. vespertilionis, nymph.

Figs, 290-297 are reproduced from Ticks, Part I,

[55]

Adaptation of Ticks

333

Fig. 295. Fig. 296. Fig. 297.

Fig. 295. 0. mvigmji, adult. Fig. 296. 0. moubata, adult.

Fig. 297. O. moubata, 1st stage nymph.

Fig. 298'.
Fig. 298. 0. megnini, larva.

Fig. 2991.
Fig. 299. 0. megnini, nymph.

1 Figs. 298, 299 are reproduced from Salmon and Stiles, 1901, Figs. 99 and 102
(Vlth Ann. Hep., V.A.I., U. S. Dept. Agricult. and from Ticks, Part I).

334 Ai^pendix II [56]

armed with long pointed teeth running down to its base (Figs. 290, 291).
The adults or nymphs of these species (Figs, 292, 293, 294) show a con-
siderable reduction in the armature of their hypostorae. In 0. savignyi
and 0. mouhata, where the larvae are inactive and the nymphs and
adults are both rapid feeders, the dentition of the hypostome is
reduced (Figs. 295, 296, 297). In 0. megnini the larval hypostome
(Fig. 298) is powerfully armed, and the same is the case with that
of the nymph (Fig. 299), which we have seen remains attached for
a long time to the host. The hypostome of the adults, which do
not suck blood, is unarmed, and the capitulum, as a whole, is quite
exceptionally small. The figures are reproduced from Ticks, Part I,
without regard to the relative magnifications employed.

Ixodes.

When we come to consider the biology of Ixodes, which are clearly
marked off from other Ixodidae structurally, there are two points
which are very striking in respect to the adult stages : (1) the con-
siderable number of species in which the females only are known, and
(2) the fact that the males of some species are frequently found in
copula on their hosts (in some species the sexes remain in copula even
when they are dropped into spirit). In no other genus of Ixodoidea
is this the case. On the other hand, in those species in which both
sexes are known, some are found in copula upon the host, whilst others
are not. I have sought a reason for these peculiarities in Ixodes, and
believe I have found an explanation of these differences. I shall begin
by discussing the host relationships in Ixodes.

Of the 51 species listed we have no data regarding the hosts of nine,
namely, nigiHcans, fossidatus, aciditarsus, gigas (^), japonensis, per-
cavatus, cordifer (</), tasmani and coxaefurcatus ((/*). We nevertheless
include two of these {percavatus and tasmani) in the subjoined lists
because the presumption appears justified that they came from birds :
percavatus came from isolated islands in the Pacific inhabited by birds,
and we possess a variety of this species, taken from a puffin ; tasmani
was collected on the Island of St Pierre, and by the celebrated ornitho-
logist, Jules Verreaux, in Tasmania. We therefore are able to list
44 species in relation to their hosts, there being but two of these
{percavatus and tasmani) with regard to which the evidence is not
quite satisfactory.

[57] Adaptation of Ticks 335

In the appended lists I have grouped the various species of Ixodes
as follows :

I. Species of which both the (^ and $ are recorded as occurring
upon the host.

(a) Species in which the sexes have been found in copula upon
the host,

(b) Species in which the sexes have been found near together
upon the host.

IT. Species of which the females only have been found upon the
host.

III. Species of which the females only are known.

The immature stages of these ticks are left out of consideration in
these lists. Alongside the name of the hosts are placed certain signs :

+ denotes a host possessing wandering habits.

0 denotes a host with fixed habitat for more or less prolonged
periods ; the habitats being burrows, nests, caves, hollow trees, etc.

+ is used in the case of Canis familiaris, as in some places it may
be fittingly described as a wandering host, in others as a host with fixed
habitat (the kennel).

A consideration of the data tabulated in the following pages brings
out certain striking facts. The ticks included under Group 1 (a) are, in
most cases, encountered upon (+) wandering hosts. In the case of
ricinus, our records show that it has been found 63 times on wandering
hosts and only 10 times on hosts which burrow. Being a very pre-
valent and hardy species, it is natural that it should occasionally occur
on a variety of hosts. The tick may at times infest dog kennels, thus
taking advantage of the fixed habitat of the host. The three varieties
of ricinus (var. scapularis, ovatus and californicus) appear to follow the
same rule, and in the case of pilosus we appear to have a striking
example of a tick occurring upon wandering hosts.

Group 1 (6). The species included in this group occur, in some cases,
on wandering hosts, in others not. Thus, angustus and tenuirostris, ac-
cording to my notes, occur solely on nesting and burrowing animals. In
angustus we have a form in which the males occur very rarely upon the
host, and in which the male hypostome^ is poorly armed; angustus may,
therefore, be regarded as an intermediate form in respect to its adapta-
tion. In tenuirostris we have a tick which, as far as we know, only

1 The variations in this structure in male Ixoclea will be discussed presently.

336

Appendix II

[58]

I. Species of Ixodes of which both </ and $ are recorded as
occurring upon the host.

(a) Species in which the sexes occur in copula upon the host.

I

Species

Host's
habits

Host

Number of Geographical

times found distribution

thereon of tick

I

ricinus ...

+

Bos taurus

... 13^

Europe, N. Africa,

+

Ovis aries

... 13

W.Asia, W.Ame-

±

Canis familiaris ...

... 13

rica.

+

Cervus elaphus ...

5

+

,, capreolus

4

+

, , dama

1

^63

+

Capra hircus

2

+

Equus caballus ...
Homo sapiens

6
2

+

Felis concolor

1

+

Genetta sp.

1

+

Lepus europaeus...

2J

0

Erinaceus europaeus

2^

0

Mustela putorius

1

0

,, erminea...

1

0

Males taxus

1

0

Mus decumanus ...

1

^10

0

"Mouse"

1

0

Myoxus sp.

1

0

Talpa europaea . . .

1

0

Ground squirrel ...

Larvae and nymphs
lizards (10) and birds
of different species.

IJ

on
(5)

I.

ricinus var. scapu-

+

Cattle ...

N. America, Costa

laris ( c? ¥ known)

+

Deer
Dog
Man

Kica.

I.

ricinus var. ovatus

+

Horse ...

Japan.

(only 2 ? s known)

=t

Dog

I.

ricinus var. califor-

+

Black-tail deer ...

N. America.

n icus

0

Grey fox

I.

pilosus ...

+

Capra hircus

3

S. Africa.

+

Bos taurus

2

+

Equus caballus ...

3

+

Sus scrofa domestica

2

+

Ovis aries

1

+

Duikerbok

1

+

Tragelaphus sylvaticus

1

db

Canis familiaris . . .

1

[59]

Adaptation of Ticks

337

Species
I. pilosus .

I. pilosus var. howardi
( i ? known)

/. boliviensis

I. schillingsi
I. rasus

I. ugcmdamis

I. ugajtdamis var. dja-
roneiisis ( s ? known)

Host's
habits

Host

+

Felis domesticus...

+

Viverra civetta ...

+

Felis paidus

Homo sapiens

+

Cat

0

Erinaceus frontalis

0

Rbinolophus sp.

?

Icticyon (Speothos) venati-

cus (Ganidae)

+

Colobus caudatus (monkey)

+

Bos taurus

+

Felis pardus

±

Canis familiaris

0

Herpestes ichneumon

0

Hyrax sp.

Homo sapiens

-f

Ovis aries

0

Aulacodus sp.

' ' Large rodent "(? Aulacodus)

?

Genetta suahelica

0

Procavia brucei

Number of

times found

thereon

1

1

1

1

Oeograiiliical

distribution

of tick

S. Africa.

S. Africa.

Bolivia.

E. Africa.

E, & W. Africa.

E. & W. Africa.

E. Africa.

(b) Species in which the sexes have been found together upon the host.

I. anguslus

0

Sciurus hudsonius douglasi

N. America.

0

Tamias townsendi (Sciuridae)

?

Lepus dalli

0

Neotoma occidentalis (Muridae)

0

" Mouse "

I. tenuirostris

0

Evotomys glareolus

3 Europe.

0

Arvicola amphibius

2

0

,, pratensis

1

0

Microtus agrestis

2

0

,, arvalis ...

0

Sorex vulgaris

0

,, miuutus ...

0

Mus miniatus

I. minor

0

Hesperomys sp. (Muridae)

1 Guatemala.

I. cavipalpus

+

Cynocephalus babuin

1 Africa (Rhodesia

Homo sapiens

I & Angola).

I. 1-ubicundus

+

Ovis aries

1 S. Africa.

+

Capra hircus

22—5

338

Appendix II

[60]

Species
J. ruhicundus var. Urn-
batus ( ? only known)

I. holocyclus

I. loricatiis

Host's
habits

+

+

+
+
±
0
0

+

0
0
0
0

Host

Number of

times found

thereon

Sheep ...
Goats

Bos taurus
Macropus sp.
Canis familiaris
Sciurus variabilis
Phascogale penicillata (Mar
supial tree shrew)

Ateles melanochoerus
Didelphys aurita
,, opossum
sp.
Microdidelphys sorex (o & l^)

Geographical

distribution

of ticlc

C. Africa (Congo).
Australia, India.

S. America.

occurs as a parasite on small mammals which burrow ; these burrows
are, however, near the surface of the ground and liable to he destroyed
or flooded, consequently, for the maintenance of the species, it is es-
sential that both sexes should be carried about upon the host as in
the case of ticks occurring on hosts with wandering habits. The other
species follow the same general lines as 1 (a) with regard to their host
relationships ; the majority of the hosts have wandering habits. In
minor (only found once) the sexes may occur together on the host for
the reasons stated above for tenuirostris ; loricatus appears to be an
exception.

If the reader will run his eye down the column of + and 0 signs in the
lists I (pp. 336-338) and II (pp. 339-340) of the hosts of the different
species of Ixodes, he cannot but be struck by the differences. The
hosts enumerated in II, under " Species of which the females only have
been found upon the host, the males being known," are nearly all
animals which burrow or nest. Thus, we record the finding of hexagonus
no less than 40 times on such hosts, once each on three wandering hosts
(if we include man), and three times on the dog. The list of hosts of
hexagonus var. cookei (taken from Banks) falls in with that of hexagonus.
In the case of canisuga, the dog is the most prominent host, simply
because nearly all of our specimens from the dog came from the north
of England where the shepherd dogs are largely confined in kennels.
The dog in these regions is a host with a fixed habitat, as with most
of the other hosts ; in only two instances out of 23 has this species
been recovered from a wandering host.

^ o and L signify that nympbal and larval stages only were found.

[61]

Adaptation of Ticks

339

II. Species of which the females only have been found upon the
host, the males being known.

Host's
Species habits

I. he.vagonus ... 0

0

Host
Erinaceus europaeus
Mustela errainea . . .

Number of Geographical

times found distribution

thereon of tick

9^ Europe, N.Africa,

8 N. & S. America.

0

,, vulgaris ...

7

0

,, putorius...

6

0

,, furo

2

0

Lutra vulgaris ...

1

>^40

0

Meles taxus

1

0

Lepus cuniculus . . .

1

0

Myopotamus coypu

1

0

Sciurus sp.

1

0

Canis vulpes

3'

=L

Canis familiaris . . .

3

+

Ovis aries

1

+

Sus scrofa

1

Homo sapiens

1

I. hexagonus var. 0

Lutra ...

N. America.

cookei ( (? ? known) 0

Mustela vison

0

Spermoplnlus

0

Weasel

0

Porcupine

0

Marmot

0

Pocket gopher

±

Dog

+

Cat

+

Sheep

I. canisuga ... ±

Canis familiaris . . .

... 12

^ Europe, N. Am-

0

,, vulpes

2

erica.

0

Cotileriparia (bank-swallow) 3

0

Mustela furo

;2i

0

Meles taxus

0

Sciurus ...

0

Talpa europaea (larva)

J

+

Equus caballus ...

1-2

+

Ovis aries

)

I.putus. On marine birds only ( c? only found in nests or

beneath rocks and soil near nests)

0

Phalacrocorax verrucosus ... Europe, N. & S

0

Pygosceles taeniatus

America, Asia,

0

Spheniscus magellanicus Australasia.

0

Fratercula arctica

3

0

Penguin sp.

2

0

Uria troile

1

0

Sea-gull sp.

1

0

Sula bassana

1

0

Fulmarus glacialis

1

0

Cormorant sp.

1

+

Man (occasionally, see

text, p. 34(

))

340 Ap2^e7idix II [62]

Species

Host's
habits

Number of Geographical
times found distribution
Host thereon of tick

^iy(c?

found in caves etc. inhabited by bats).

0
0
0
0
0
0

Plecotus auritus ... ... Europe, Africa,

Ehinolophus hipposideros Australia.

,, ferrum equinum

,, euryale

,, hippocrepis

,, blasius.

0

,, clivosus

0
0

Vesperugo pipistrellus
,, tricolor.

Ixodes putiis and vespertilionis are solely parasites of birds and bats,
although putus may occasionally attack man when he approaches the
infested birds' nests.

With the exception of /. hexago7iiis\ including var. cookei, of which
we have no particulars, the males of these species have never, apparently,
been captured upon the host. On the other hand, the males of the other
species have been found in the habitats of their hosts : canisuga in the
nests of sand-martins (bank-swallows) and in a dog kennel, vespertilionis
in caves inhabited by hats, putus in and about the nests of marine birds.
In such localities the males would have ample opportunities of feeding
upon their hosts for brief periods after the manner of male Argasidae,
assuming that they do feed. It is conceivable, however, that such
habitat-infesting males may not be blood-suckers at all. We have seen
that Ornithodorus megnini J" does not feed, and that it nevertheless
fertilizes the female. The same may hold for certain Ixodidae </s.
The matter requires further investigation. The sexes of canisuga and
jmtus have often been observed in copula apart from the host, and it
appears probable that this is the usual occurrence in nature^

List III (p. 341) is likewise very striking. It includes all the
species (26) of which the males are still unknown, With the exception
of three species (bicornis, stilesi and australiensis), all of the females
have been found solely on birds, burrowing or nesting hosts, and bats.
Of /. bicornis only three females are known ; they were found on
as many hosts. Ixodes stilesi has only been found once, and all the
specimens (14 females) were found upon a single host. Ixodes austra-
liensis was found once on a dog ( + ), and once on the marsupial rat-
kangaroo (0). It is true that many of the species are represented only
by one or two females, or by few specimens, and consequently there
may be an error in certain cases in attaching much importance to the
character of the type host — nevertheless, there is a very remarkable
co-relation between the absence of males on the host and the life habits

of the host.

1 See note on p. 345.

[63]

Adaptation of Ticks

341

III. Species of which the females only are known^.

Species

Host's
habits

I. acuminatus

0

I. dentatus

0

I. diversifossus

0

I. fuscipes

0

I. sculptus

0

I. spinicoxalis

0

I. marxi

0

I. rubidtis

0

I. nitens

0

I. bicornis

+

I. auritulus

0

I. brunneus

0

I. caledonicus

0

I. simplex

0

I. texanus

0

I. stilesi

+

I. neumanni

0

I. percavatus

0

I. percavatus var.
rothschildi

0

I. lunatus

0

I. ornithorhynchi

0

I. tasmani

0

I. fecialis

0

I. fecialis var.

0

aegrifossus

I. australiensis

±

0

I. vestitus

0

I. signatus
I. tmicavatus

Host
Mus agrarius (only 2 9s known)
Rabbit (1 ? known)
Procyon lotor (2 ? s recorded)
Dasyprocta aguti (? Felis pardalis)
" Rock squirrel "
Mustela flavigula (4 ? s known)
Red squirrel and fox (few ? s known)...
Bassaris astuta (1 ? known)
Mus macleari (2 ? s known) ...

Felis concolor, F. onca, Homo sapiens
(may = diversifossus, only 3 d s known)

Trupialis militaris and another bird.sp.?

Birds: many species

Domestic pigeons ...
Rhinolophusferrumequinum,Vespertilio

Grey squirrel, Procyon lotor (few ? s

known)
Padua humilis (Cervidae) ; (14 ? s

found once)
Mai-iue birds (10 ? s known)...
Marine birds (presumably)

Marine bird (puflfin)

Hallomys audeberti (1 ? known)
Ornithorhynchus (found 5 times)
Probably birds (Verreaux coll.)
Dasyurus geoffroyi (1 ? known)
Opossum (3 ? s); Paramelesobesula(l ? )

Canis familiaris (found once)

Bettongia lusueuri (marsupial rat-kan-
garoo), (found once)

Myrmecobius fasciatus (Dasyuridae),
(found once)

Diemenia superciliosa (snake), (found

once)
Phalacrocorax
Phalacrocorax graculus and P. carbo. ...

Geographical

distribution

of tick

Italy.

N. America.

N. America.

C. & S. America.

N. America.

Sumatra.

N. America.

Mexico.

Christmas Island,
Pacific Ocean.

Mexico.

S. America.

Europe, N. Ame-
rica.

Scotland.

China.

N. America.

Chili.

New Zealand.

Tristan d'Acunha
Island.

No locality given.

Madagascar.
Tasmania.

Australia.

Asia, N. America.
Great Britain.

^ Nymphal and larval stages are known in many of these species.

842 Appendix II [64]

On the structure of the hypostome in ^ Ixodes.

There are 18 species of Ixodes of which the males and females are
known. About one of these {pratti, Banks) we have no information
such as we require for this discussion. Of the 17 remaining species,
the sexes have been found together upon the host in 13 (I (a) and {b)),
and in 4 (II) only the females have been found upon the host:

I (a). In six the sexes have been found in copula upon the host,
namely, ricinus, rasus, ugandanus, schillingsi, jyilosus and hoUviensis.
The hypostomes in the males of the first five species are characterised
by prominent basal teeth ; hoUviensis has a well-armed hypostome.

I {h). In seven species (cavipalpus, rubicundus, minor, loricatus,
tenuirostris, angustus and holocyclus) both sexes have been found upon
the host, but there is no specific mention of their having been found
in copula. The hypostomes of the first two species possess prominent
basal teeth ; minor, loricatus and tenuirostris possess hypostomes which
are well armed with pointed teeth ; in angustus and holocyclus the
hypostome is only moderately armed.

II. In four species (hexagonus, canisuga, imtus and vespertilionis)
there is no record of the males having been found upon the host. In
the first two the hypostome is but moderately armed, whereas in putus
and vespertilionis it is almost unarmed.

The structure of the male mouthparts, therefore, bears a direct
relation to the presence or absence of the male upon the host and the
occurrence of the sexes in copula upon the host. Of the six species
(I (a)) which have been found in copula upon the host no less than
five possess hypostomes with prominent basal teeth. It is probable that
some of the species placed in the following division (I (b)) will yet be
found in copula when they have been more carefully observed. In
Group I (a) and (h) there are 13 species enumerated in only two of which
the hypostome may be described as moderately armed. In Group II,
regarding which there is no record of the males having been found upon
the host, the male hypostome is but slightly armed in two and prac-
tically unarmed in two species.

[65]

Adaptafio7i of Ticks

343

Fig7ires illustrating the hypostomes of Ixodes.

The accompanying figures show the marked sexual dimorphism which is observable in
some species of Ixodes, and also types of male hypostomes which show considerable
modifications of structure.

300 301

302

303

304 305

306

307

Figs. 300, 301 represent the hypostomes of I. ricinus s and ? . That of the i may be
taken as the type of a hypostome with large basal teeth as observed also in the males
of rasiis, 2iganda7ms, schillingsi, rubicundus, cavipalpus and pilosus, the basal teeth
being less prominent in the last two species mentioned. In boliviensis <? the basal
tooth, whilst prominent, is much less developed.

Fig. 302 represents the hypostome of /. minor i , with well-developed outer files of teeth.
(The hypostome of tenuirostris and loricatus approximate to the type of structure
fornd in the females.)

It is poorly but relatively better

Fig. 303 represents the hypostome of I. holocychis <? .
armed than that of canisuga which follows.

Fig. 304 represents the hypostome of 1. canisuga <? , poorly armed, and much emarginated
distally. That of hexagonus is similar but not emarginated.

Fig. 305 represents the hypostome of I. imtits <? , practically unarmed, emarginated, and
so reduced in length as to be useless for attaching the tick to a host. This structure
alone suggests that the male of this species is probably incapable of sucking blood.

Figs. 306, 307 represent the hypostomes of I. vespertilionis <? and ? ; that of the s practi-
cally unarmed ; that of the ? illustrating the dimorphism existing between the sexes.
The figures are chosen from the series illustrating this work, without regard
to the magnifications employed.

N.B.— It is a striking characteristic of hexagonus, canisuga, piitus and vespcrtiUons males
that the capitulum is 7niich sinaller compared to the size of the body than in any other
male Ixodes that are known. The size of the capitulum apparently bears a direct
relation to the dentition of the hypostome and to the habits of the male with regard
to copulation and parasitism on the host.

344 Appendix II [66]

Summary.

A consideration of the facts presented in the foregoing pages
appears to warrant certain conclusions which must, however, be regarded
in part as provisional and subject to revision when our knowledge of
the various species of ticks has become more extended. The views
here expressed may prove of practical use in the study of the Ixodidae.

The Argasidae represent the relatively primitive type of ticks
because they are less constantly parasitic than are the Ixodidae. Their
nymphs and adults are rapid feeders and chiefly infest the habitat of
their hosts. In certain Argasidae (0. moubata and 0. savignyi) the
disadvantage of their possessing an " active " larval stage has resulted
in the development of an " inactive " larva, i.e. the young nymph being
the first to suck blood. In 0. megnini we have a considerable adaptation
brought about by the difficulty there must be in the tick entering the
small aperture of the ear more than once. Owing to the Argasidae
infesting the habitats of their hosts, their resistance to prolonged
starvation and their rapid feeding habits, they do not need to bring
forth a large progeny, because there is less loss of life in the various
stages, as compared to Ixodidae, prior to their attaining maturity.

The Ixodidae are more highly specialized parasites than the pre-
ceding. The majority are parasitic on hosts having no fixed habitat
and consequently all stages, as a rule, occur upon the host. In the
genus Ixodes we find an adaptation of certain species according to the
habits of the hosts upon which they are parasitic. In the species which
usually occur upon wandering hosts both sexes are found upon the
host, whereas in other species which occur on hosts possessing more or
less fixed habitats the males are rarely or never found upon the host.
The males of species of Ixodes, both sexes of which occur upon the host,
are characterized, as a rule, by the possession of hypostomes with
prominent teeth, the reverse being the case in forms wherein the males
do not occur upon the host. Where species occur upon a wandering
host it is essential, for their propagation in nature, that both sexes
should be carried about upon the host.

It appears to follow that the copulation of ticks upon the host is an
indication of a higher degree of specialization to a life of parasitism.
When copulation does not take place on the host, we have merely the
retention of a primitive character as found in Argasidae. From the

[67] Adaptation of Ticks 345

fact that many species of Ixodes are only known to science in their
female and immature stages, it would appear probable, if my theory
holds, that this is merely due to the males not having as yet been sought
for in the habitats of their hosts \

A great many males belonging to Haemapliysalis, Deiinacentor,
Rhipiceplialus, Amhlyomma (and Aponomma) are known, these genera
of Ixodidae being rich in species. The males of BoopJdlus, Hyaloinma,
Rhipicentor and Margaropus, of which there are few species, are likewise
known. In all of these genera the males are found close to the females
and attached to the host. These genera at'e chiefiy parasitic on animals
with wandering habits, and the males of these genera do not exhibit
hypostomes which are markedly dissimilar to those of the females;
the hypostomes are well armed in both sexes. It is only in the genus
Ixodes that we find well-marked sexual dimorphism in respect to the
structure of the hypostome.

1 As a further confirmation of the views here expressed, I would state that we
have recently (March, 1911) discovered a male of Ixodes hexagoniis in the nest of a
hedgehog. This is the first male we have captured, whereas we possess hundreds of
females, nymphs and larvae found on various hosts. (The rarity of the male is rendered
evident by the fact that there is not a single specimen of a male in the Museums in
London, Paris and Berlin.) When the male was placed in a receptacle with an engorged
female from the same nest, copulation occurred after a few minutes and was repeated
several times.

346

Genus Ixodes

INDEX TO VALID SPECIES OF IXODES

Together with a List of the Collections in which
the Types are to be found.

stages of each

Names of Species,
Authors and Date

acuminatus Neumann, 1901

acutitarsus (Karsch, 1880)

*angustus Neumann, 1899

auritulus Neumann, 1904

*australiensis Neumann, 1904

bicornis Neumann, 1906
boliviensis Neumann, 1904
*hrunneus Koch, 1844
*caledoiiicus Nuttall, 1910

*canisuga Johnston, 1849

*cavipalpus Nuttall and War-
burton, 1907
cordifer Neumann, 1908
coxaefurcatus Neumann, 1899
dentatus Marx, 1899, iu Neu-
mann, 1899
diversifossus Neumann, 1899
*fecialis Warburton and Nuttall,

1909
*fecialis var. aegrifossus War-
burton and Nuttall, 1909
fossulatus Neumann, 1899
fuscipes Koch, 1844
*gigas Warburton, 1910

*hexagonus Leach, 1815
*hexagonus var. cookei (Packard,

1869)
*holocyclus Neumann, 1899

japonensis Neumann, 1904

^ Determined by Karsch, but not the type, which is neither in Berlin nor Munich, and
cannot be traced.

* Species contained in the Cambridge collection ; types, and others which have been
compared with the types.

Collections which include

species which

the Types

are known

Page

Toulouse, 2 ? s

?

161, 341

Berlin \ 1 ?

?

202

Washington (a), 1 ? ;

(? ? O L

195, 315, 334 et seq.

Cambridge, <? o l

Paris (a), 3 ? s;

?

187, 341

Toulouse, 1 ? , co-type

Cambridge, 2 ? s ;

?

250, 341

Toulouse, 1 ? , co-type

Toulouse, 3 ? s

¥

186, 341

Toulouse, 1 ,? , 1 ? , 1 o

(? ? o

166, 334 et seq.

Berlin, 1 ? (in balsam)

¥ O L

189, 341

Cambridge, 1 ? , o l

¥ O L

198, 341

Edinburgh, 1 ? , o l

Lost, but substitutes in

<^ ¥ O L

209, 316, 334 et seq.

Cambridge : c? ? o l ;

Toulouse

Cambridge, <? s, ? s

S ¥

192, 334 et seq.

Leyden, 1 c?

c?

233

Berlin, 1 c?

<?

270

Washington (b), 1 ?

¥

162, 341

Washington (e), 2 ? s

¥

163, 341

Cambridge, ¥

¥

248, 341

Cambridge, ?

¥

250, 341

Berlin, 1 ?

?

201

Berlin, ? ; Toulouse, o l

¥ O L

169, 341

Calcutta, 1 <? ;

<?

203

Cambridge, 1 cf

London, ?

<? ¥ O L

177, 330, 334 et seq,

Harvard, ?

¥

183, 334: et seq.

London, Paris (a).

<? ¥ o

235, 334 et seq.

Toulouse, Berlin, ? s,

Hamburg

Paris (a), 1 ?

?

208

Index to Valid Species

347

stages of each

Collections which include

species which

the Types

are known

Page

Toulouse, London, <? ?

<r 9

O L

266, 334 e« seq.

Cambridge, %

?

269

Leyden, 1 ?

?

226, 341

Washington (f), ?

?

173, 341

Toulouse, 1 <? , 1 ?

c? ?

164, 384 et seq.

Toulouse, 7 ? s, 12 o s

? o

217, 341

Hamburg, 1 ?

?

159

Cambridge, 2 ? s (o lost)

?

185, 341

Paris (a), ? o ; (^) ?

? o

242, 341

London, 4 ? s

?

220, 341

Cambridge, 2 ? s, 3 o s

? o

221, 341

Berlin, 1 ? ; Toulouse,

<? ?

O L

221, 316, 334 e«

seq.

<f ? L

Toulouse, i ?

• i ?

226, 334 et seq.

Washington {d), i ?

i ?

174

Paris (a), i ? ; Pickard-

rf ?

O

256, 317, 334 et

seq.

Cambridge, o

Toulouse, (J ? ; Berlin

c? ?

228, 334 et seq.

Losti

i ?

0 L

143, 296, 334 et

seq.

Washington (c)

■? ?

159, 334 et seq.

Hamburg, 1 ? ;

?

158, 334 et seq.

Toulouse, 1 ?

Lost

S ?

156, 315, 334 et

seq.

Toulouse, c? ? ;

(? ?

204, 334 et seq.

Pretoria, rf ?

Katanga, ?

s

207, 834 et seq.

Toulouse, 1 ? , 3 o s

? o

175, 341

Paris (rt), i ; Berlin, ? s

c? ?

O

238, 334 et seq.

o s ; Cambridge, ? o ;

co-types

Washington (6), 1 ?

?

171, 341

St Petersburg, ? ; Hamburg,

? o

L

261, 341

o ; Cambridge, o l

London, ? o

? o

207, 841

London, 4 ? s

?

171, 341

Paris (a), 14 ? s

?

216, 341

Paris (a), ? s ; Berlin, 1 ?

?

244, 341

London, j ; Berlin, ? o ;

c? ?

O L

246, 834 et seq.

Cambridge, ? o l

Washington {d), ?

?

214, 841

London, 1 j , 1 ?

i ?

230, 384 et seq.

Sjostedt, s ?

c? ?

288, 834 et seq.

Names of Species,
Authors and Date

*loncatus Neumann, 1899
*loricatus var. spinosus Nuttall,
1910

lunatus Neumann, 1907

marxi Banks, 1908

mirior Neumann, 1902

neumanni Nuttall and War-
burton, 1911

nigricans Neumann, 1908
*mtens Neumann, 1904

ornithorhynchi Lucas, 1845

percavatus Neumann, 1906
*percavatus var. rothschildi Nuttall

and Warburton, 1911
*pilosus Koch, 1844

pilosus var. howardi Neumann,

1908
*pmtti Banks, 1908
*putus (Pickard-Cambridge, 1876)

*ras!/.s' Neumann, 1899
*i-icinus (Linnaeus) Latreille, 1804
ricinus var. californicus (Banks,

1904)
ricinus var. ovatus (Neumann,
1899)
*ricinus v&r.scapidaris (Say, 1821)
*ruhicundus Neumann, 1904

rubicundiis var. Umbatus Neu-
mann, 1908
ruhidus Neumann, 1901
*schilliiigsi Neumann, 1901

sculptus Neumann, 1904
*signatus Birula, 1895

simplex Neumann, 1906
spinicoxalis Neumann, 1899
stilesi Neumann, 1911
tasmani Neumann, 1899
*tenuirostris Neumann, 1901

*texanus Banks, 1909
*ugandanus Neumann, 1906
ugandamts subsp. djaronensis
Neumann, 1907

1 Not in Upsala: only Hyalomma aegijptium still in the Linneau Collections, 1911.
* Species contained in the Cambridge collection ; types, and others which have been
compared with the types.

Collections which include
the Types

stages of each

species which

are known

Page

Edinburgh, ? o l;
Cambridge, l

? O L

264,

341

Berlin, 1 ?

cT ? O L

271

334 et seq

London, 1 5 ; Cam-
bridge, O L

? O L

252,

341

348 Genus Ixodes

Names of Species,
Authors and Date

*unicavatus Neumann, 1908

*vespertilionis Koch, 1844
vestitus Neumann, 1908

* Species contained in the Cambridge collection; types, and others which have been
compared with the types.

The important collections, judged by the number of types they contain, are to be
found in the following cities (the figures in parentheses indicate the number of types in the
cities enumerated): Toulouse (16), Cambridge (15), Berlin (11), Washington (10, in five
collections), London (9), Paris (8), Hamburg (4). Under types are included the different
stages of each species which have been first described from the various collections.

Note : In the second column of the foregoing list the whereabouts of the collections is
but briefly indicated :

Berlin signifies Entomologische Abteilung, Zoologisches Museum, Berlin.

Calcutta ,, Entomological Department, Indian Museum, Calcutta.

Cambridge ,, Collection of Professor G. H. F. Nuttall, F.R.S., Quick Laboratory,

University of Cambridge.
Edinburgh ,, Collection of Mr W. Evans, F.R.S.E., Edinburgh.
Hamburg ,, Naturhistorisches Museum, Hamburg.

Harvard ,, Museum of Comparative Zoology, Cambridge, Mass., U.S.A.

Katanga ,, Special Committee's Collection, Katanga, Congo Free State.

Leyden ,, Natural History Museum, Leyden, Holland.

London ,, Entomological Department, British Museum (Natural History),

Cromwell Road, London.
Paris (a) ,, Musee d'histoire naturelle, rue Buffon, Paris.

,, (6) ,, Laboratoire de Parasitologic, Ecole de Medecine, Paris.

Pickard-Cambridge signifies Collection of Rev. 0. Pickard-Cambridge, F.R.S., Wareham,

Dorset, England.
Pretoria signifies Collection of Mr C. P. Lounsbury, Acting Union Entomologist,
Union Department of Agriculture, Pretoria, Transvaal, S. Africa
(late Government Entomologist, Cape Town).
St Petersburg ,, Collection of Professor A. Birula, K. Akad. Wiss., St Petersburg.
Sjostedt ,, Collection of the Sjostedt Zoological Expedition.

Toulouse ,, Collection of Professor L. G. Neumann, Laboratoire d'Histoire

naturelle, Ecole Nationale V^terinaire, Toulouse, France.
Washington (a) signifies Bureau of Animal Industry, U.S. Department of Agriculture,
Washington, D.C.
,, (b) ,, National Museum, Washington, D.C.

,, (c) ,, Collection of Mr N. Banks, of the Bureau of Entomology,

U. S. Department of Agriculture.

,, (d) ,, Bureau of Entomology, U.S. Department of Agriculture.

,, (e) ,, Collection of Dr A. Hassall, of the Bureau of Animal Industry,

U.S. Department of Agriculture.

(The distribution of types in the collections in Washington is given by

Hooker, 1909, pp. 423-425.)

PARASITOLOGY

A SUPPLEMENT TO THE
JOURNAL OF HYGIENE

Edited by George H. F. Ndttall, RR.S., Quick Professor of Biology in
the University of Cambridge, and A. E. Shipley, F.R.S., Master
of Christ's College.

When the Journal of Hygiene was founded it was announced that papers
on Parswitology " in relation to hygiene and preventive medicine " would be
published in its pages. It has however been felt for some time that the
Journal was becoming undidy burdened with papers dealing with the anatomy
of mosquitoes, fleas, protozoa and other parasites — of great importance in
themselves — but having only an indirect relation to hygiene and preventive
medicine.

The remarkable development of parasitology in recent years, and the
increase in our knowledge of the part played by parasites in human and
animal diseases, demand a means of publication, in the English language, of
original papers dealing with the subject in its widest sense. It is proposed in
fU^re to relegate all such papers to Parasitology.

Papers on the subjects we have mentioned are now scattered in journals of
widely different character, into some of which they ar6 but grudgingly admitted.
We trust that Pa/rasitohgy will fulfil the purpose the editors have in view,
of encouraging the study of parasitology, especially in relation to disease, by
providing a means for the publication of papers relating to pathogenic and
disease-transmitting parasites.

It is therefore hoped that this publication will appeal not only to
medical men at home' and abroad, but to veterinarians, zoologists and
agriculturalists.

Parasitology i though a supplement to the Jov/rnal of Hygiene, will be
issued in the form of a separate yearly volume of four to five hundred pages.
The parts will be issued when sufficient material has accumulated. The
first three volumes are now ready. The subscription price is £,\. Is. per
volume (post-free), payable in advance. Subscribers to the Journal of
Hygiene may obtain single numbers of Parasitology at the reduced price of
5s. net, or may become subscribers at the reduced rate of 1 5^. Subscriptions
may be sent to any Bookseller, or to Mr C. F. CLAY, Makagbb, Cambridge
University Press, Fetter Lane, London^ E.C. (The Contents of the last
number of Parasitology is given on the next page)

PABASITOLOGY (coyitinued)
CONTENTS OF VOL. IV, NO. 1 (March 1911)

GiLRUTH, J. A., Sweet, Georgina, and Dodd, Sydney. Observations

on the occurrence in the blood of various animals (chiefly mono-

tremes and raarsupials) of bodies apparently identical with

Anaplasma marginale, Theiler, 1910. (With Plate I)
Huffman, O. V. and Wherry, Wm B. A description of four Filaria

loa from the same patient. (With Plate II)
Drew, Harold G. A note on the application of Giemsa's Romanowsky

stain to the blood and tissues of marine invertebrates.
Porter, Annie. Some remarks on the genera Orithidia, Herpetomonas

and Trypanosoma.
HiNDLE, Edward. The passage of Trypanosoma gamhiense through

mucous membranes and skin.
MACKINNON, Doris L. On some more protozoan parasites from

Trichoptera. (Plate III and 8 Text-Figures)
NuTTALL, George H. F. and Merriman, G. The process of copulation

in Ornitkodorus moubata. (With one Text-Figure)
NuTTALL, George H. F, On the adaptation of ticks to the habits of

their hosts. (With 26 Text-Figures)
Bacot, a. W. The persistence of Bacillus pyocyaneus in pupae and

imagines of Musca domestica raised from larvae experimentally

infected with the bacillus.
Huffman, Otto V. The embryos of Filaria loa. (With Plate IV)

TICKS

A Monograph of the Ixodoidea

Now ready. PART I. THE ARGA8IDAE. With 116 text
figures and 8 plates. Royal Svo. hs. net.

This book will deal with the Classification, Structure and Biology
of Ticks, the study of the group having occupied the authors for several
years. Practically all that has been published on the subject has
received adequate consideration.

The book will be issued in four or more parts, which will be
complete in themselves. A complete bibliography to the work will
appear as a separate Fasciculus simultaneously with Part II.

CAMBRIDGE ; PRINTED BY JOHN CLAT, M.A. AT THE TJNIVEBSITT JKEBS.

Part III, Ixodidae]

[October, 1915

TICKS

A MONOGRAPH OF THE

IXODOIDEA

By

GEORGE H. F. NUTTALL, M.A., M.D., Ph.D., Sc.D., F.R.S., Fellow of
Magdalene College, Quick Professor of Biology, University of Cambridge

CECIL WARBURTON, M.A., F.Z.S., Christ's College, Zoologist to the
Royal Agricultural Society

W. F. COOPER, B.A., F.Z.S., F.L.S., and L. E. ROBINSON, A.R.C.Sc. (Lond.),
of the Cooper Laboratory for Economic Research, Watford

PART III

THE OENUS HAEMAPHYSALXS

GEORGE

H.

F. NUTTALL, F.R.S.,

AND

C.

WARBURTON

CAMBRIDGE
AT THE UNIVERSITY PRESS

LONDON: FETTER LANE. C. F. CLAY, Manager
AND H. K, LEWIS, GOWER STREET

EDINBURGH : lOO, PRINCES STREET

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MACMILLAN

COMPANY

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JOURNAL OF HYGIENE

EDITED BY

GEORGE H. F. NUTTALL, M.D., Ph.D., Sc.D., F.R.S.

QUICK PR0FE8S0B OF BIOLOGY IN THE UNIVEKSITY OF CAMBRIDGE
IN CONJUNCTION WITH

JOHN S. HALDANE, M.D., LL.D., F.R.S. ARTHUR NEWSHOLME, C.B., M.D.

LATE READRB IN PHYSIOLOGY IN THE MEDICAL OFFICER TO THE

UNIVERSITY OP OXFORD LOCAL GOVERNMENT BOARD

CHARLES J. MARTIN, M.B., D.Sc, F.R.S.

DIEBCTOB OP THE USTEE INSTITUTE, LONDON

J. C. G. LEDINGHAM, M.B., D.Sc. G. S. GRAHAM-SMITH, M.D.

BACTERI0L0OI8T-IN- CHIEF, LISTER INSTITUTE, UNIVEESITY LECTURER IN HYGIENE,

LONDON CAMBRIDGE

M. GREENWOOD, Junr., M.R.C.S.

STATISTICIAN, LISTER INSTITUTE, LONDON

The Journal of Hygiene ia issued as material accumtdates. A volume containing
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Volumes I, II, and III (1901-3) complete. In Four Parts, paper covers
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The subscription price is £1. Is. per volume (post-free), payable in advance
Subscriptions may be sent to any Bookseller, or to Mr C. F. CLAY, Manager
Cambridge University Press, Fetter Lane, London, E.C.

CONTENTS OF VOL. XV, NO. 1 (July 1915)

Hewlett, R. T. and Revis, C. On a Complement-stimulating Sub-
stance in Cow's Milk.

Brownlee, J. Studies in the Meaning and Relationships of Birth and
Death Rates.

II. Density of Population and Death Rate (Fan's Law).

III. The Constitution of a Death Rate. (With one Chart.)

IV. On the Range of Instances in which Geometrical Pro-
gressions describe numerically processes of life, i.e. those
processes which might be explained by a monomolecular
reaction.

V. On the difficulty that in applying the laws of physical
chemistry to life processes, indices occur which suggest
the actions of fractions of a molecule.

Radcliffe, J. A. D. Complement Fixation in Pulmonary Tuberculosis.

Mackintosh, J., Pennington, S. and Williams, R. S. The Supply of
Non-tuberculous Dairy Stock.

Haffkine, W. M. Concerning Inoculation against Plague and Pneu-
monia and the Experimental Study of Curative Methods.

Fletcher, W. The Wassermann and Luetin Reactions in Leprosy.

EwART, R. J. The Influence of Age of the Grandparent at the Birth
of the Parent on the Number of Children born and their Sex.
(With one Text-figure.)

Taylor, F. E. The Sterilisation of Vaccines ; and the Influence of the
Various Methods Employed on their Antigenic Properties. (With
one Chart.)

TICKS

A MONOGRAPH OF THE

IXODOIDEA

PART III

CAMBRIDGE UNIVERSITY PRESS

C. F. CLAY, Manager

ILonton: FETTEK LANE, E.G.

GEDinburgl): 100 PRINCES STREET

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rcikun: THE MARUZEN-KATUISHIKI-KAISHA

All rights reserved

Prof. L.-G. NEUMANN of Toulouse

To whom we dedicate this fasciculus in grateful acknowledgment
of the help he has given us

TICKS

A MONOGRAPH OF THE

IXODOIDEA

By

GEORGE H. F. NUTTALL, M.A., M.D., Ph.D., Sc.D., F.R.S., Fdlow of
Magdalene College, Quick Professor of Biology, University of Cambridge

CECIL WARBURTON, M.A., F.Z.S., Christ's College, Zoologist to the
Royal Agricultural Society

W. F. COOPER, B.A., F.Z.S., F.L.S., of the Cooper Laboratory for
Economic Research, Watford

L. E. ROBINSON, A.R.C.Sc. (Lond.), of the Cooper Laboratory for
Economic Research, Watford

PART HI

Cambridge :

at the University Press

1915

IXODIDAE

Part III. The Genus Haemaphysalis

by

GEORGE H. F. NUTTALL

and

CECIL WARBURTON

PREFACE TO PART III

WE have little to add to what has been stated in the Preface to
Part II. The present fasciculus follows in order and deals with
the genus Haemaphysalis of which we recognize 50 species and varieties ;
three species (spinulosa, ohtusa and numidiana) being included although
we are somewhat doubtful as to their validity. Many species hitherto
regarded as valid are noted in our synonymic lists.

All of our descriptions are original ; they include three new species
and three new varieties, abeiTant forms and numerous hitherto un-
described stages of known species.

The terms and signs used are the same as in Part II, pp. 127-182,
to which the reader is referred.

Illusti^itions.

We again lay stress upon the illustrations. Those appearing in
the six plates are republished from papers by Nuttall, Cooper- and
Robinson, and Nuttall respectively, and comprise 29 figures in addition
to 144 figures in the text, making a total of 173 figures illustrating
this Part. Of these figures, 24 are taken from other authors, the
impressions being made from the original blocks. The majority of the
new text-figures were drawn under our direction by Mr N. Cunliffe.

The genus Haemaphi/salis has been very poorly illustrated hitherto
and we have sought to make good this deficiency, many species,
including their immature stages, being now figured for the first time.

Of the 72 new text-figures 45 were drawn by N. Cunliffe

24 „ ,, G. H. P. Nuttall

2 „ „ N. Cunlitfe and E. Wilson

1 was drawn by C. Warburton

Of the 72 previously J 21 are from publications by C. Warburton
published text-figures^ 21 ,, ,, L. G. Neumann

17 „ „ C. Warburton and G. H. F.

Nuttall
9 ,, ,, G. H. F. Nuttall

2 ,, ,, W. Donitz

1 is from a publication by L. E. Robinson
1 ,, ,, Nuttall, Cooper and Robinson

144

Most of the figures were drawn to scale, others, not so drawn, are
described as " sketches."

vi P^'eface

A chno wledf) men ts.

We gratefully acknowledge the generous gift of £100 from Mr and
Mrs P. A. Molteno towards the expenses of publishing this work.

We are especially indebted to the following gentlemen and institu-
tions for access to material : Professor L.-G. Neumann (lately of
Toulouse) has kindly lent us all of his types and has presented us with
co-types wherever possible in exchange for similar specimens from our
collection. We have to thank Dr L. Lavarra (Rome), Professor Gestro
(Genoa Museum), Professor A. Brauer (Berlin Museum), Professor S.
Schenkling (Deutsches Entomologisches Museum, Berlin-Dahlem) and
Mr S. Hirst (British Museum) for the gift of co-types. We have
received numerous specimens from the Imperial Bureau of Entomology
(Mr Guy A. K. Marshall, Secretary) and from the many gentlemen
whose names are mentioned as collectors in the text.

The bracketed numbers accompanying the " records of specimens
from different countries relate to catalogues in particular collections
severally indicated by a letter or the name of the collection before the
number. Thus all numbers preceded by " N " relate to specimens in
our collection ; those preceded by " E " relate to specimens received for
identification from the Imperial Bureau of Entomology, London. Other
collections we have examined and determined are as follows : (Ash-
worth) = Dr J. H. Ashworth, Zoological Department, University of
Edinburgh ; (Berlin Mus.) = Zoologischcs Museum, Berlin ; (D.E.M.)
= Deutsches Entomologisches Museum, Berlin-Dahlem ; (Genoa Mus.)
= Museo Civico di Storia Naturale, Genoa ; (Gowdey) = Mr C. C. Gowdey,
Government Entomologist, Entebbe, Uganda ; (Ind. Mus.) = Indian
Museum, Calcutta; (K.) = Mr N. B. Kinnear, Bombay Natural History
Society; (Knuth)= Professor Knuth, Tierarztliche Hochschule, Berlin;
(Liverpool) = Liverpool School of Tropical Medicine ; (Schtiffner) = Dr
W. Schtitfner, Deli, Sumatra ; (Sweet) — Dr Georgina Sweet, University
of Melbourne, Australia. Where two numbers are connected by =
(thus E. 692 = N. 3153) the sign indicates that the specimens have
been divided between the two collections E. and N.

As stated in Part II, we shall be grateful to readers who may draw
our attention to any errors and omissions.

G. H. F. N.
C. W.

Cambridge,

Septembe); 1915.

CONTENTS

OF
PART III

II. THE GENUS UAKMAPHYSALIS C. L. KOCH, 1814.

Synonymy and Literature relating to the Genus ......

The Generic Characters defined .........

Keys for the determination of species of Hacmaphjsalia :

Males

Females

Nymi)hs .

Larvae

Note to terms used ...........

PAGE

349
351

Specific descriptions of valid s[)ecics and of their varieties

N.B. The signs J^, 5 j o, L alongside of the specific names, indicate
the various developmental stages (male, female, nymph, larva)
which are known to science

1. inermis c^, ^, o, L .

„ var. aponommoidcH 5

2. warhiirtoni J , $

3. cinnaharina ^ ^ $ , o, L .

„ var. p^mctata ^ , ? , o.

4. leporis-palmtris (^ , 9 ? o> L

5. inontgomeryi (^ , ? .

6. kinneari $

7. uhurensis $

8. fonnoseiisis S ■> 9 ' •

9. japonica ^

,, var. douglasi (^

10. papiiana ^ , 9 > O) L

11. Jlava (^,9

12. turturis ^

13. aciculifer ^, 9

14. kochi (^, $

15. birmaniae ,^

16. silacca $ .

1 7. parmata ^ , 9 ? o, L

353
356
359
360
361
362

362
367
369
372
378
387
395
397
398
400
402
403
404
408
410
411
413
415
416
418

Vlll

Contents

18. hystricis c?) ? •

19. bispinosa ^, $ , o, L

„ var. intermedia ^, $

20. parva (^ , $ , o, L .

21. cuspidata (^, $ , o, L

22. aculeata (^, $

23. calcarata (J , ? , o .

„ var. houyi ^,

24. calvus cJ, $

25. spinigera ^ , '^

„ var. novae-guineae c , 9 ,

26. concinna (^ , $ , o, L

27. simplex ^ , o .

28. ^eofcAi (J , $ , o, L

29. spinulosa $

30. ohtusa $ .

31. numidiana cJ, $

32. tvellingtoni ^ , $ , o, L

33. doenitzi cf, ?.

34. /iooo?j (^ , $ , o, L

„ var. orientalis ^ , 9

35. hancrofti ^ , 9 » O) L

36. campanidata $, 9

37. howletti cj, ? •

38. vidita ^ .

39. humerosa (^ , ? , o

40. elongata ^ , ?

41. cornigera ^, 9

,, var. anomala (^

42. dentipalpis ^ .
Geographical distribution and host« of th
List of condemned and doubtful species

synonymy and literature, by G. H
Notes on the Biology of Haemaphysalis by G. H
Haemaphysalis cinnaharina var. punctata

„ cinnabanna

„ leporis-palustris

„ leachi

„ coticinna .

„ inermis

Index to valid species of Haemapkys^

e Genus Haemaphysalis
of Ha,emaphysalis, including
F. Nuttall .

Nuttall

their

alis^ together with a list of the collections

in which the types and co-types are to be found

PAGE

422

426

433

435

438

440

442

444

445

447

449

452

458

460

476

477

478

479

482

483

486

487

491

493

495

496

498

500

504

505

507

512
518
519
528
530
536
542
545

548

TICKS

PLATE VIII

oLors&l aspect

^^s.-

ventraL aspect

E Wilson, Cambridge

HAEMAPHYSALIS PUNCTATA Can 5c Fan,
The Larva

TICKS

N^mph

Female (unfed)

dorsal aspect

Female (replete)

0 If

ventral/ aspect

HAEMAPHYSAL

PLATE IX

Male (replete)

Male (unfed)

E.Wilson.del et Uth Cambridge

NGTATA Can. b. Fan.

TICKS

s.oh''

Fig.1.

d.ch^ 9-

dLOCGO^V -

Fig.2.

L.E.Iiancl JTorddel.

Fig.3.

s.ch-

^0

..-d/ ch

ch'.

■I :#

''s.d.

bloc, cav

Fig.4.

PLATE X

d.ch sh. \ I

s.chr ■

'.>b.c.

Fig.5.

■ s ch.

F.g.6.

d.ch

d ch.

3s,2.

> 3 if.

••:b.c.

Fig.7.

Morgan J. Kidd Richmond CoUoty

Fig.8.

TICKS

PLATE XI

a, nip. . - -

'.ch.

Fig.

sh..

Fig.4.

s.ch

kd.

pr

Fig. 2.

sh.

LERand J, Ford. del.

Fig.5.

Fig. 6.

Morgan 4Kidd Riohmond Collotype.

TICKS

PLATE XII

W^

1

Fig. I

Fig-. 2

1#

Fig-- 3

Fig:- 4

Fig. 5

TICKS

PLATE XIII

Fig. I

Fig. 2

LIST OF ILLUSTEATIONS

Frontispiece : Portrait of Professor L.-G. Neumann . . facing Title-page

PLATES'

Facing p. viii
Plate VIII. Haemaphysalis cinnaharina var. punctata

Larva : dorsal and ventral aspeets, x 62.
Drawn from living specimen by J. Ford.

Plate IX. Haemaphysalis cinnabariaa \ar. punctata

Nymph in dorsal aspect, x 30.
Female unfed, in dorsal jispect, x 20.

„ gorged, in dorsal aspect, x 5 detail of siu-face more highly

magnified to left.
„ gorged, ventral aspect, x 5. Spiracle to right.
Male unfed, dorsal aspect, x 15.
„ replete, ventral aspect, x -20.

,, unfed. Posterior portion of ventral surface, showing appearance
of festoons for comparison with those of the replete male, x 20.
Drawn from living specimens by E. Wilson.

Plate X. Haemaphysalis cinnaharina var. punctata

Fig. 1. Capitulum of Female, dorsal aspect, x 85.

Fig. 2. „ „ „ ventral aspect, x 85.

Fig. 3. ,, „ Male, dorsal aspect, xllO.

Fig. 4. ,, „ „ median portion with left palp, in ventral

aspect, X 125.

Fig. 5. „ ,, Nymph, dorsal aspect, x 220.

Fig. 6. „ „ „ ventral aspect, x 220.

Fig. 7. „ „ Larva, dorsal aspect, x400.

Fig. 8. „ „ „ ventral aspect, x400.

' N.B. Plates VIII — XI are reprinted from Nuttall, Cooper and Robinson, vi. 1908
(Pis. VIII and XI); Plates XII and XIII are reprinted from Nuttall, iv. 1904 (Pis.
XII and XIII).

List of llhistratlons

Plate XI. Haeindphijsalis cinnaharina var. punctata

Fig. 1. Female : tactile hairs on infra-internal margin of palji, x 250.
Fig. 2. Male : ventral surface, showing details of external structure,

x30.
Fig. 3. Larva : chelicerae, seen from the ventral surface, x 1000.
Fig. 4. Female : right chelicera, seen from the ventral surface ; the

sheath removed, x 250.
Fig. 5. Male : left chelicera, seen from the ventral surface, x 250.
Fig. 6. Male : light chelicera, lateral aspect ; sheath removed, x 250.

Plate XII. tlueinaphysalis leuchi

Fig. 1. Female ovipositing, ca. x 2^.

Fig. 2. Larvae as they appear on emerging from the egg ; dorsal aspects,
x20.

Fig. 3. Nymph (unfed) ; ventral aspect. Balsam mount, x 20.

Fig. 4. Showing three gorged nymphs in the centre, an unfed male to
the left, an unfed female to the right. The male appears
glossier than the female because of the scutum which covers
its whole dorsum. The adults ai'e about the same size as the
gorged nymphs from which they emerge.

Fig. 5. Male in ventral aspect. Balsam mount, x 20.

Plate XIII. Ilaemaphysalis leachi

Fig. 1. Female in ventral asi)ect ; the scutum being seen by trans-
parency. The body appears broader than in the male, x 20.
Fig. 2. Male capitulum in ventral aspect, x 150.

IN THE TEXT

FIOrRE

308. Illustrating llaem<iplijjsalis : H. wellinytoni ^ dorsum and venter,

^ dorsum ; with names of parts indicated in accordance with the
generic description in the text .....

309. Haemaphysalis inermis ^, dorsum .....

310. ,, „ 9 , essential parts

311. „ „ o, „ „ . . . .

312. „ ,, Jj, „ ,,....

313. „ inermis var. apotiommoides 9 > essential parts

314. „ ivarburtoni (^ , dorsum and essential parts

315. ,, ,, $ , essential parts

316. „ cinnabarina ^, dorsum and essential parts

317. „ „ $ , essential parts .

318. „ „ ? , caijitulum and scutum

319. ,, „ o, essential parts

320. „ „ L, „ „ . . .

321. „ cinnabarina var. punctata :

(J, dorsum and essential parts.

322. 9 , essential parts ....

351
363
364
365
366
368
370
371
373
374
374
375
376

381

382

List of IlhistrationH

XI

323. Haemaphysalis

324.

325.

326.

327.

328.

329. „

330.

331.

332.

333.

334.

335.

336.

337.

338.

339.
340.
341.

342.
343.
344.
345.
346.
347.

348.

349.

350. „

351.

352. „

353. „
354.

355.

356. „

357.

358.

359. „

360. „
361.

362. „

362a. „

363.

364.

365.

cinnabarina var. pidictata :

9, capituluiu, scutmii, spiracle and tarsus IV
o, dorsum and essential parts .
leporis-palvMris ^, dorsum and essential parts
„ ,, 9 > essential parts

„ „ 9 > palps, tarsi I and IV .

„ „ o, essential parts

montfiompryi (^ , doi'suni and essential parts

„ 9 ) essential parts .

kinneari 9 ? essential i)arts ....
ahorensis 9 •
formoaends ^, dorsum and essential paiis

„ $ , essential parts

japonica ^, dorsum .....

„ (^, venter, spiracle

papuana ^, dorsum, venter, spiracle

9 , capitulum, scutum, spiracle .
» c?> atypical form : dorsum and essential parts

„ o, essential parts ....

flava (^, dorsum, coxae and spiracle .

„ 5 1 essential 2>arts ....
ttirturis ^ , dorsum and essential parts
admlifer (J, „ „ „ „

,, 9 J essential parts

loc/n (^, dorsum and essential i)arts

5 , capitulum and scutum .
hirmaniae ^, dorsum and essential parts .
silacea 9 ■, dorsum and essential parts
parmata (J, „ „ „ „

„ 9 ? essential paits ....

■>■> '^i 11 11 • • • •

11 '-'i 11 11 • • • •

hystricia ^, dorsum and essential i)arts
„ ^, dorsum, venter and spiracle .

„ 9 ? essential parts ....

„ 9 1 dorsum and essential i)arts

bispinosa ^, dorsum, venter, spiracle, tarsus I\
,, (^, dorsiun, part of venter, etc. .

„ 9 ) essential parts

„ 9 > P^ip ^iid tarsus IV

,, o, essent'al parts

» ^1 11 11 ...

bispinosa var. intermedia ^, capitulum

„ „ „ 9 ) capitulum, scutum, etc

parva (^, dorsum .......

PAGE

382
383
389
390
390
391
392
395
396
398
399
400
401
403
403

404
405
406
407
409
409
411
412
413

414
41G
417
419
419
420
420
423
423
424
424
427
428
429
429
430
430
434
434
436

Xll

List of Illustrations

FIGURK

366.

Haemaphysalu

367.

n

368.

»

369.

M

370.

»

371.

»

372.

»

373.

J>

374.

55

375.

55

376.

55

377.

55

378.

55

379.

55

380.

55

381.

55

382.

55

383.

55

384.

55

385.

55

386.

55

387.

55

388.

55

389.

55

390.

55

391.

5*

392.

5)

393.

)5

394.

55

395.

55

396.

55

397.

55

398.

55

399.

55

400.

55

401.

)5

402.

55

403.

55

404.

55

405.

55

406.

55

407.

55

408.

55

409.

55

parva ^, capituliim in ventral aspect, coxae I
cuspidata (^ , dorsum, etc. ....

„ $ , capituluin, scutum, etc.

„ o, essential parts

55 ^^i )5 )» ...

aculeata (^ , dorsum, etc

55 ? ) capitulum, scutum, etc. .
calcarata (J, dorsimi, venter, spiracle and tarsus

IV

„ c^, palp

„ ^, coxae and trochanters .

„ $ , essential parts ....

caltms (J, dorsum, venter, etc

„ 5 , essential parts .....
spinigera (J, capitulum in dorsal and ventral aspect:

„ 1^, coxae, palp, digit ....

„ 9 5 scutimi, capitulum, tarsus IV

„ 5 , coxae and spiracle ....

spinigera var. novae-guineae (J, dorsum, venter, etc.

„ ,, „ „ $ , essential parts

„ „ „ „ o, dorsum, venter, etc.

eoncinna ^, dorsum, part of venter, etc.

„ (^, capitulum

)j 6^ hypostome

6, digit ....

„ $ ) essential parts

11 ^1 51 )5 • •

)) -Lis )) )) • •

simplex ^, dorsum, part of venter, etc.

„ (^ , digit and palp .

„ (J, coxae and trochanters

„ ^, tar.sus IV .
leachi (J, dorsum, etc.
„ (^, showing variation in size
„ (J, capitulum ....
,, (J, coxae .....
„ ^ and $ , digits
„ 9 , dorsum and part of venter .
„ o, dorsum, venter, etc.
„ L, capitulum in dorsal and ventral aspects
,, L, scutum .....

„ L, venter

„ 9 , " var. indica" essential parts
,, (^ , " I'oningshergeri" dorsum etc
„ 9 5 )) scutum, capitulum, spir

acle, tarsus IV .

PAGE

436
438
438
439
439
441
441

442
442
443
443
444
445
446
447
448
448
448
450
451
451
453
453
454
454
455
455
456
459
459
459
459
462
462
463
463
463
463
465
465
465
466
468
468

468

List of Illustrations

Xlll

FIGURE

411. Haemaphysali

412.

))

413.

>»

414.

»

415.

»

416.

»j

417.

>»

418.

5»

419.

)>

420.

»

421.

>>

422.

»

423.

)5

424.

5>

425.

»

426.

)>

427.

5)

428.

»>

429.

»

430.

)>

431.

»

432.

5)

433.

)5

434.

)i

435.

)?

436.

))

437.

))

438.

>7

439.

)5

440.

))

441.

)?

442.

>5

443.

J)

444.

)»

445.

»

446.

>5

447.

U

448.

)>

449.

»

450.

t spinulosa 9 > scutum aiitl capital um .

„ 9 > ventral aspect of capituluni, coxae I

obtusa (^ , capitulum in dorsal ami ventral aspects,

coxae I .
numidiana ^, dorsum part of venter, etc.

„ $ , scutum, capitulum, .spiracle

wellingtoni ^, dorsum and venter

„ 9 > dorsiun

„ 9 ) hypostome and digit .

„ o, essential parts

doenitzi ^, dorsum ....

„ 5 5 scutum, capitulum, spiracle, tarsus IV
hoodi (^, dorsum, etc. ....
„ 9 , scutum, capitulum, etc.
„ o, essential parts

>? '-'■) 11 )) • • •

bancrofti ^, dorsum and essential parts

„ 9 , essential parts

» o, » „ • •

campanulata ^, dorsum, coxae, spiracle

„ $ , scutum, capitulum, spiracle, tarsus 1\

howletti (^ , dorsum and essential parts

„ $ , essential parts .
vidua ^, dorsum, etc.
hnmerosa (^ , dorsum, etc. .

„ $ , scutum and capitulum .

elongata ^, dorsum and essential parts
„ (^, pal]), part of venter

„ 9 , dorsum part of venter, spiracle, tarsus I\

cornigera ^, „ „ „ „

„ (^, capitulum in dorsal and ventral aspects
coxae ......

(?, palp

„ (J, coxae and trochanters

„ $ , essential parts .
„ var. anomcda ^J, capitulum in dorsal and
ventral aspects .
dentipalpis ^ , capitulum in dorsal and ventral aspects
^'^ longicomis" Nn. $> palp and coxa I
cinnabarina var. punctata, longitudinal section of capi
tulum
„ „ „ process of oviposition .

PAGE

476
476

477
478
478
480
480
480
480
481
482
482
484
484
485
485
488
488
489
489
491
491
493
494
495
496
496
499
499
500
501

502

502
502
503

504
505
513

525
527

349

Genus II. HAEMAPHYSALIS C. L. Koch, 1844.
Synonymy and Literature.

The following references only relate to publications which deal with the genus
or cognate matters.

Ixodes 1827. Audouin, p. 428 (/. lcachii=H. leachi). 1849. Gervais, p. 49

(/. lagotis= Haemaphysalis n]).). 1869. Packard, p. 67 (/. chordeilis=H. cinna-

harina Koch ; /. leporis-palustris = U. leporis-palustris). 1880. Megnin, p. 132

(7. chelifer=H. concinna Koch ^). See further under our list of condemned
species.

Haemapliysalis 1844. Koch, p. 237 (genus first defined. Four species : 1 H. rosea,
2 cinnabarina, 3 sanguinolenta, and 4 concinna, of which dnnaharina and

concinna are good species). 1847. Koch, p. 25. 1877. Canestrini and

Fanzago, p. 110 ; Murray, p. 199. 1890. Canestrini, pp. 483, 493. 1892.

Canestrini, p. 581 ; Marx (a), p. 233, (6), p. 283 ; Trouessart, p. 47. 1895.

Railliet, p. 714 (nothing original). 1897. Neumann, pp. 326, 358 (key for

determination) ; Supino, pp. 241, 243. 1900. Ward (a), p. 200 (nothing

original). 1901. Salmon and Stiles, pp. 457-459, figs. 219-221 (from Neumann,

1897 ; they give short generic diagnosis, synonymy, bibliography and transla-
tion of Neumann's key). 1905. Labille, pp. 14, 15, 44, 154. 1906. Buy,

p. 118 (generic characters tabulated) ; Wheler, p. 420. 1907. Donitz, pp. 15,

67, 111 ; Hunter and Hooker, p. 52 (incomplete generic diagnosis) ; Pocock,
p. 190 (key to genera of ticks) ; Warburton (a), p. 92, pi. VII, fig. 2 (classifica-
tion briefly treatedl 1908. Banks, pp. 13, 20, 32 ; Bonnet, p. 259 (condensed

from Neumann), figs. 28-30 (poor) ; Nuttall (VII), p. 16 (oviposition described
and figured) ; Nuttall, Cooper and Robinson, p. 155 ; Warburton (IV), p. 508

(diflficulties of classification discussed). 1909. Blanchard, pp. 145-147 (generic

characters defined, key to species following Neumann) ; Hunter (VI), p. 255
(thinks ground birds disseminate immature stages) ; Rohr, p. 141 (nothing

original, quotes Neumann and Aragao). 1911. Neumann (cr), p. 105 (gives

key to 13 species recognised as valid together with brief notes on 13 doubtful
species ; four species comprise nine subspecies) ; Nuttall, p. 67 (habits of
ticks in relation to hosts ; behaviour of male considered) ; Nuttall and
Merriman, p. 44 (males of this genus, in common with other Metastriate
ticks, have a ehitinous flap over the sexual orifice) ; Nuttall and Warburton,

p. 119 (of this book ; generic characters fre.shly defined and figured). 1912.

Hooker, Bishopp and Wood, p. 89 (nothing original) ; Warburton (VII), p. 122

(generic charactei-s discussed). 1913. Nuttall, p. 195 (structure of hypostome

and variability observed in the genus) ; Patton and Cragg, p. 627 (genus

defined and key for determination taken from Neumann, 1911 a).

N. I. 23

350 Genus Haemaphysalis

Wrongly spelt Haemophijsalis by Canestrini, 1884, p. 113, and by a few other
authors.

Haemaphysalis concinna Koch, designated as the type of the genus by
Neumann, 1901, p. 340.

Rhipistoma 1844. Koch, p. 239 (two species : 1 R. leachii, 2 R. ellipticum). 1847.

Koch, p. 27 (but on p. 135 R. ellipticum is transferred to the genus Rhipi-
cephalus ; both ellipticum and leachii, however, = Haemaphysalis leachi

(Audouin, 1827). 1861. Furstenberg, p. 208. 1878. Karsch, p. 337

("ii. leachii"). 1888. Dugfes, p. 129. 1890. Canestrini, p. 483.^ 1892.

Marx (a), p. 233 ; (6), p 283. 1896. Osborn, p. 261 {R. leporis which = //aema-

physalis leporis-palustris (Packard, 1869)). 1897. Neumann, p. 326 (given as

a synonym of Haemaphysalis). 1901. Salmon and Stiles, p. 457 (follow

Neumann as do also subsequent authors).

Wrongly spelt Rhipidostoma by Karsch, 1878 ; Duges, 1888 ; also misspelt
Rhiphistoma by Osborn, 1896, and the authors who quote him and the spelling of
the foregoing writers.

Rhipistoma leachii Koch, may well be designated as the type of the genus
although R. ellipticum might serve as well ; since they are synonymous.
Salmon and Stiles, 1901, p. 457, cite Marx as if he had designated '' H. leachii"
as the type, but he did not do so.

Rhipicephalus 1847. Koch, p. 135 {R. llipticus = H. leachi). 1877. L. Koch (6),

p. 196 {R. e.vpositicius = moat probably H. cinnabarina \a.v. punctata (Canestrini
and Fanzago, 1877).

Gonixodes 1888. Dugfes, p. 129. 1897. Neumann, p. 343 (as a synonym of

HaemapJiysalis ; subsequent authors agree with Neumann. Gonixodes rostralis
Dugfes, the only species = partly Haemaphysalis leporis-palustris (Packard,
1869)).

Herpetobia 1890. Canestrini, pp. 486, 493, 527. 1891. Canestrini, p. 719.

1897. Neumann, pp. 327, 329 (as a synonym of Haemaphysalis ; we agree
with Neumann. Herpetobia sidcata Canestrini, the only species = probably
Haemaphysalis cinnabarina var. punctata (Canestrini and Fanzago, 1877) ;
immature stages which Canestrini, 1891, p. 719, supposed might prove to
belong to one of the already established genera).
Opisthodon 1897. Canestrini, p. 468 {0. cuscobius the type = H. cuscobia, a nominal
species). Supino (o) and (6), p. 252, gives the following 3 species : 1 0. asiaticus,
which Neumann regards as a nominal species : we have examined the type and
find it to be H. leachi; 2 0. canestrinii, and 3 0. gestroi Supino = likewise
H. leachi. Neumann, 1897, p. 326, gives Opisthodon as a synonym of Haema-
physalis, with which we agree.

Wrongly spelt Opistodon by Wheler, 1906, p. 420, and either authors.
Prosopodon 1897. Canestrini (a), p. 417 (name proposed as a substitute for
Opisthodon which the author found to be preoccupied ; one species, P. cziscobiics
= merely a nominal species). Merely listed by Blanchard, 1909, p. 150, and
Neumann, 1911a, p. 105.
Not Pseudixodes Haller, 1882, p. 311, as stated by Canestrini, 1890, pp. 485-526.

Generic Characters

351

Generic Characters : Metastriata, i.e., with anal grooves embracing
the i(7ius poster io)iy. Usually of small size. Scutum inornate and
without eyes, and in the female, without lateral grooves. Capitulum
With base sub-rectangular, and with palps normally short and conical,
broadest near the posterior end of article 2, which {except in rare cases)
projects laterally beyond the base. Sexual dimorphism slight, the male
possessing no ventral plates or shields. Coxa I never bifid ; trochanter I
with a blade-like dorsal retrograde process.

The chief difficulties presented by the genus Haemaphysalis arise
from the absence of characteristics — such as eyes, colour-markings, anal
armature in the ^, etc. — which are of great specific value in other
genera. The prevailing colour is yellow, in ungorged specimens, and
the general integument is rather highly chitinised, so that those
structures which in most ticks are conspicuous by their comparative

<,aJ/:a^ '^'^^^■^

i^/'o^'i*:/!

Fig. 308. HdemitjiJnjmlis loellimitoni s in dorsal and ventral aspects, ? in dorsal
aspect. (Reprinted from Pai't II, Fig. 120.) To show the generic characters and
nomenclature of parts.

hardness stand out in less salient relief. Thus, the scutum of an unfed
nymph is sometimes only visible after rather close examination. The
capitulum is in most cases much alike in the cf and % so that in many
species identification is almost equally easy from either sex. There
are exceptional cases, however (e.g. H. concinna, H. cornigera, etc.), in
which the ^ palps have striking peculiarities not found in those of
the % . The sexes are more frequently dimorphic as regards the coxal
armature. Wherever the coxal spines are strongly developed in the
</ their peculiarities are only very faintly echoed in the $ . There is

23—2

352 Genus Hae^iiaphysalis

what may be considered a normal coxal armature to which a large
number of species of Haemaphysalis conform in both sexes (Fig. 360).
It consists of a moderate internal spur on coxa I (which is sub-
triangular), a slight spur at the middle of the posterior borders of coxae
II and III, and at the internal angle of coxa IV. If a $ specimen
possesses anything more pronounced than this in the way of coxal
armature it is pretty certain that the (/* coxae will be unusually
spinose.

All species of Haemaphysalis have a blade-like dorsal retrograde
spur on trochanter I. Some species have ventral retrograde spurs on all
or some of the trochanters, and these are excellent specific characters
usually to be found both in the </ and in the % . Many forms have a
hardly perceptible prominence in that position, but we shall only
speak of the trochanters as " spurred " if this characteristic is readily
recognizable.

Species belonging to the genus Haemaphysalis are less readily
identified than Ixodes by means of a dichotomic key, because, as
already stated, there are few salient features which serve to
differentiate them. The diagnosis has frequently to be based on
numerous minor points which, in the aggregate, permit of the identifi-
cation of a species, due regard being paid to individual differences
which may be fairly marked. Stress must be laid for this reason upon
accurate figures to accompany the text.

Although Neumann (1901, p. 340) designated H. concinna Koch
as the type of the genus, the (/" of this species is atypical in that its
palps are unlike those of any other member of the genus. Of the
four species originally included by Koch, who founded the genus
Haemaphysalis, only a second species, H. cinnabarina, remains which
would take priority over H. concinna and might, with better right, be
designated as the type of the genus.

We have attempted a linear arrangement of the species, beginning
with atypical forms like H. iner^nis, with the palps simple and non-
salient laterally, and proceeding by degrees to species like H. dentipalpis
with complex, strongly salient palps. No such arrangement can be
more than partially successful, but it is hoped that nearly allied species
will in most cases be found in tolerably close juxtaposition.

Ken f^ species

353

Key for the determination of species of Haemaphysalis'.

Some varieties are included under tlie type, others are directly
determinable by the key. The species which include varieties are
indicated by {v) after the specific name. There are 45 species and
varieties whose j/s are known.

Males.

4.
5.
6.

I Palps not salient laterally, i.e. no wider than the
basis capituli ......
Palps more or less salient laterally

{No lateral grooves or cornua, dentition 2 | 2
(Europe, Transcaucasia)
Lateral grooves and cornua, dentition 4 ] 4 (Asia)

jNo lateral grooves ......

(Lateral grooves present .....

'Palpal article 2 with marked lateral salience
curving forwards (Malaya) ....
] Palpal article 2 only slightly salient and scarcely
\ larger than article 3 .

[Palpal article 3 with dorsal retrogi-ade spur (Asia)
[ „ „ 3 without spur (Burma)

jCoxal spurs normal and inconspicuous
(One or more coxae strongly spurred .

[All the coxae with strong subequal spurs
(Coxae II and III only feebly spurred

{All trochanters with ventral spurs
Only trochanters I and II spurred ; tarsi
humped (Africa)

Talpal article 2 much longer than 3 ; body pro-
truding behind scutum (Madagascar) .

Palpal articles 2 and 3 sub-equal, each with a
ventral retrograde spur ....

1
2

inerinis
warburtoni

32
5

vidua

hystrids
hirmaniae

16
6

7
10

8

362
369

495

422
415

calcarata (v) 442, 444

elmgata
9

498

1 For explanations of terms and signs used see pp. 127-132 {Ticks Part 11). and p. 361.
^ See II. obtusa, from K^union, which is leachi-hke in other particulars.

354

Genus Haemaphysalls

9.
10.
11.
12.

13.

14.

15.

16.
17.
18.

19.
20.

/Palpal article 2 with external spur, on lateral

--; salience (Borneo) ..... calviis

I Palpal article 2 with spur at internal angle (India) montgonieryi

Coxa IV with 2 long spurs close together (Asia) cornigera (v)
„ „ „ 1 long spur . . . . .11

C Coxae I and IV with long sj)urs (India, Ceylon,
J. Judea) ......

lOnly coxa IV with a long spur .

[Lateral grooves long .....
[ „ „ short .....

["Palps rounded laterally, very short (Europe, N
Africa, Asia Minor, Transcaucasia)

spinigera {v)
12

13
14

cinnaharina
var. punctata
Palps salient laterally, curving forwards

(America) ....... cinnaharina

Palpal article 2 with ventral spur (New Guinea,

Queensland) . . . spinigera var. novae-guineae
Palpal article 2 without spur . . . .15

Coxa IV with needle-like spur ; palpal article 3

as broad as 2 (Africa) ..... aciculifer

Coxa IV with stouter and more tapering spur ;

palpal article 2 broader than 3 (Asia) . flava

Palpal article 3 with dorsal spur, more or less erect 17
„ „ 3 with no such spur

(Lateral grooves very short (Africa)
I „ „ fairly long

Cornua very strong, as long as basis capituli
moderate .....

21

parmata
18

19
20

Palpal article 2 very long, much longer than 3
(India, Ceylon) . . . . .

Palpal articles 2 and 3 sub-equal (Ceylon)

I^Scutum long-oval, finely punctate (Asia, E.

! Africa, N. S. Wales)

I Scutum somewhat truncate posteriorly, coarsely
1^ punctate, a small species (India, Ceylon)

aculeata
cuspidata

PAGE

445

395

501

447

380
373

449

411

408

418

440
438

bispinosa (v) 427, 433
parva 435

Key to species 355

f Palps pincer-like, article 3 being long and

21. j incurved (Europe) .....
^ Palps otherwise ......

/Palpal article 2 with several processes at pos-

22. \ terior margin (Malaya) ....
^Palpal article 2 otherwise .....

(■Palps only slightly salient laterally, articles 2

23. j and 3 sub-equal ......

^Palps strongly salient at the base of article 2

/Trochantal spurs strong; lateral grooves very

24 J short (Ceylon)

( Trochantal spurs absent or inconspicuous .

rScutuui with surface irregular, very coarsely

25. - punctate^ (Asia) ......

'.Scutum otherwise ......

(-Lateral grooves very short ; palpal article 3 with

26. \ very strong ventral spine (S. America)
lOtherwise ........

/Hypostome GjG; trochantal spurs very slight

27. J (Formosa, Burma) „

[Hypostome 515; trochantal spurs absent (Japan) japonica (v)

28.

29
30

PAGE

concinna

453

22

dentipalpis

505

23

24

28

turturis

410

25

papuana

404

26

kochi

413

27

formosensis

400

japonica (v)

402, 403

["Basis capituli with ventral cornua; hypostome

j 3 j 3 (N. and S. America) .... leporis-palustris 388

Basis capituli without ventral cornua ; hypostome

with more teeth 29

Scutum markedly elongate
short-oval

(Each palp long and narrow (Australia)

I „ „ conical, article 2 being broad basally .

("Lateral grooves very short ; tarsi humped (Mada-
gascar) .......

^^' I Lateral grooves long ; tarsi tapering (Africa,
{ Asia, N. S. Wales)

1 Regarding au atypical form from Malaya which is devoid of coarse punctations, see
p. 405.

2 See also H. numidiana, from Algeria, p. 478.

30

32

linmerosa

496

31

simplex

458

leachi -

462

356

Genus Haemcqihy salts

32.

33.

34.

35.

rPalpal article 3 recurved to a point at postei'o-
\ internal angle ......

I Palpal article 3 straight along posterior border .

'Lateral contour of palp much curved (Africa)

or (Australia)

Lateral contour of palp nearly straight (a very

small species, Asia) .....

/Lateral salience of palp projecting almost at a
J right angle (small species, Singapore)

Lateral salience a gentle curve, capitulum bell-
^ shaped .......

[Hypostome 6 | 6 ; tarsi tapering (India)
[ „ 4 I 4 ; tarsi humped (Asia)

33

34

Itoodi (v)

483

hancrofti^

487

wellingtoni

479

doenitzi

482

35

howletti

493

campanulata

491

2.

3.

6.

Females.

There are 41 species and varieties of which the $ s are known.

/"Palps not salient laterally, no wider than basis
\ capituli ....... 1

I Palps more or less salient laterally . . .3

rTarsi humped ; coxal spurs rather strong and
\ blunt (Asia) ...... luarhurtoni

ITarsi not humped ; coxal spurs small and pointed 2

(Hypostome 3|3 (Transcaucasia, Caucasia, France) inermis {v)
[ „ 4 1 4 (Formosa, Burma) . . . for^mosensis

4
8

(Trochantal spurs distinct .

( „ „ feeble or absent

[Trochanters all spurred

(Distinct spurs on some trochanters only

(Scutum broader than long (Borneo) .
I „ longer than broad .

rPalps strongly salient ; hypostome 3 [ 3 (Mada-

1

5

7

calvus
6

elongata

gascar) ......

Palps very slightly salient; hypostome 5 1 5 (India) montgomeryi

370

364, 367
401

446

500
396

^ These two species should be differentiated by refereuce to the descriptions.

Key to sjjecies

357

10.

11.

12.

13.

14.

15.

16.

'Trochanters : only I and II bear conspicuous
spurs (Africa)

Trochanters: only II and IV bear conspicuous
spurs (India)

I Palpal article 3 with distinct dorsal spine, more
or less erect ......
Palpal article 3 without dorsal spine, but pos-
terior border may protrude

rPalpal article 2 much longer than 3 (India,

Ceylon)

I Palpal articles 2 and 3 sub-equal

{Lateral salience of palpal article 2 with ventral
retrograde process (India, Ceylon, Judea) .
Lateral salience of palpal article 2 without such
a process .......

I Dorsal spine on palpal article 3 very strong;
cornua very strong (Ceylon)
Dorsal spine on palpal article 3 moderate ; cornua
moderate .......

rScutum longer than broad (chiefly Asia) .

I „ at least as broad as long

Basis capituli thrice as broad as long, broadest
posteriorly (Africa) .....
] Basis capituli twice as broad as long, rectangular
V (Asia)

[Scutum much longer than broad

( „ only slightly or not longer than broad .

Palp with lateral salience protruding sharply

almost at right angles to axis (Singapore) .

„ with lateral contour straight or only slightly

curved .......

rPalp much longer than broad ....
A „ short cone-shaped, as broad as long (Africa,
[ Asia, N. S. Wales)

PAGE

calcarata (v)

443

howletti

494

9

14

aculeata

441

10

spinigera (v)

448

11

cuspidata

. 12

. bispinosa {v)
or (India, Ceylon) ^Jarva^
. 13

parmata

hysti^icis

1.5

19

doenitzi

16
17

leachi ^

438

428, 433
436

419

424

483

464

1 Not at present distinguishable in the ? .

^ See also H. spinulosa, from Uganda (p. 476) and 7f. numidiana, from Algeria (p. 478).

358

Genus Haemapliymlis

17.

18.

19.

20.

21.

23.

24.

25.

26.

27.

28.

/Capitulum base with ventral cornua ; hypostome
J 3 I 3 (N. and S. America) .... leporis-palastris

Capitulum base normal ; hypostome with more

teeth 18

^Lateral grooves very long; hypostome 5 5

J (Australia) ....... humerosa

I Lateral grooves half the scutal length ; hypo-

\ stome 4 j 4 (Asia) ..... wellingtoni

{ Palpal article 3 as broad or broader than article 2 20
( „ ,,3 distinctly narrower than article 2 21

rScutum broader than long ; tarsi very long

\ (Abor Country)

IScutum cordate ; tai'si moderate (Africa) .

rScutum broadest in front ; porose areas large
, ,, „ middle ; porose areas mode-

ahorensis
aciculifer

22

1

rate

23

/'Palps rounded laterally (Europe, N. Africa, Asia
„^ J Minor, Transcaucasia) . . cmnabarina var. punctata

Palps with more pronounced lateral salience pro-
, curved (America) ....

(Hypostome 6 \ 6 (Europe) ....
[ „ less than 6 | 6 .

r Palpal article 3 with distinct ventral spur .

^ „ „ 3 „ no spur or at most a slight

I point .......

(Palpal article 2 widest distally (India)
1 „ „ 2 „ basally .

Scutum cordate, narrowing posteriorly ; tarsi
humped (Asia) .....

Scutum sub-circular or with sides sub-parallel
tarsi tapering .....

Coxa I with long sharp spur (Asia) .
„ I ,, slight spur ....

'Basis capituli thrice as broad as long; porose
areas far apart ......

Basis capituli twice as broad as long; porose
areas reniform, closer together (Brazil)

cinnaharina

concinna
24

25

30

kinneari
26

cainpanulata

27

cornigera
28

29
kochi

PAGE

390

497
479

398
412

381

374
454

397

492

503

414

29.

30.

Keji to species 359

/Scutiini finely punctate ; cervical grooves page

) shallow (Africa) silacea 416

Scntnm coarsely punctate ; cervical grooves

deep (Asia) papuana 406

I Palps rounded laterally, widest at middle of
article 2 (Asia) flava 409

Palps with strong lateral salience at base of
article 2 31

I Palpal article 2 with posterior border straight
and horizontal (Africa) .... hoodi (v) 484

Palpal article 2 with posterior border contour
broken (Australia) bancrofti 489

Nymphs.

There are 19 species and varieties of which the os are known.

r Palps narrow basally, not salient laterally

} (Europe, Transcaucasia) .... inermis 365

IPalps salient laterally beyond the basis capituli . 1

rBody very elongate and narrow ; palpal article 2

•> produced backward (Australia) . . . humerosa 497

lOtherwise ........ 2

(Basis capituli with ventral cornua . . .3
i „ „ without ventral cornua . . 5

jBasis capituli rectangular (N. and S. America) . leporis-palusti^is 391
1 „ „ with pointed projecting sides . 4

/-Occurring in Europe, N. Africa, Asia Minor,

\ Transcaucasia .... cinnabarina var. punctata 382

lOccurring in N. and S. America . . . cinnabarina 375

j'Palp with lateral salience slight, articles 2 and 3

sub-equal ....... 6

.10

6.

iPalps with lateral salience marked
Cornua veiy strong (Ceylon)

inconspicuous

^ (Palpal article 3 longer than 2 (Asia)
I „ „ 2 at least as long as 3

cuspidata 439

7

ivellingtoni 481

8

360 Genus Haemaphy sails

PAGE

[Scutum broader than long (Africa) . . . parmata 420

I „ sub-circular . . . . . .9

Basis capituli shorter than palps ; lateral palpal

salience rounded (Asia) .... papuana 406

9. i Basis capituli as long as palps, lateral salience
pointed (Asia, E. Africa, New South Wales) hispinosa 430

or (India, Ceylon) parva 437

rPalp with lateral salience bearing a ventral

10. \ retrograde point 11

I „ without such a point . . . .13

rCoxal spurs distinct and sharp (Africa) . . calcarata 443

11- i or c, var. houyi^ 445

I „ „ weak and inconspicuous . . .12

-^2 (Refer to Figure (Africa, Asia, N. S. Wales) . leachi 465

i „ „ „ (New Guinea, Queensland)

spinigera var. novae-guineae 450
13 JTrochantal spurs present (Queensland) . , hancrofti 490

1 „ „ absent 14

rBasis capituli with sides rounded; no cornua

14. \ (Europe) concinna 455

IBasis capituli rectangular ; with distinct cornua . 15

[Tarsi humped (Madagascar) .... simplex 458

1 „ otherwise (Africa) hoodi 485

Larvae.

We know the larvae of 14 species and varieties. All known larvae
have hypostomes with dentition 2 j 2.

(Basis capituli with ventral cornua (America) leporis-palustris 391
(Basis capituli otherwise 1

rPalps, when joined, forming an obtuse angle
1. \ (Africa, Asia, N. S. Wales) .... leachi 465

IPalps otherwise ...... 2

{Palpal article 2 not protruding laterally (Europe,
Transcaucasia) inermis 365

Palpal article 2 protruding .... 3

^ H. calcarata var. hoinji has a slight ventral spur on trochanter I which is not
discoverable in the type o.

Key to sjyecies

361

3.

5.

G.

Basis capituli without distinct cornua
„ „ otherwise

(Basis capituli without projecting lateral angles
1 „ „ with projecting lateral angles

(Europe, N. Africa, Asia Minor,

Transcaucasia) . cinnabarina var. punctata

or (America) .... cinnabarina

.Basis capituli with posterior border straight
(Europe) ......

(Asia, E. Africa, N. S. Wales) .

{ (Africa)

(India, Ceylon) .....

Basis capituli with posterior border concave

(Asia)

jCoxa I spinose ......

( „ I with blunt tuberosity (Africa)

Coxae II and III unarmed (Ceylon) .
„ II and III slightly armed

Basis capituli broader than long (Queensland) . bancrofti
„ „ narrow (Asia) .... papiuma

concinna
bispinosa
parmata
parva

wellingtoni

7
hoodi

cuspidata

383

375

456
431
420

437

481

485
439

490
407

I^ote to Terms and Signs.

Ventral corruia is a new term whereby we describe processes protruding from
the postero-lateral angles of the ventral surface of the basis capituli. So far we
have only observed them in H. leporis-palustris (all stages ; see Figs. 325, .326, 328,
329), and in H. cinnabarina (o and L ; see Figs. 319, 320), as well as in its var.
punctata (o and L, Fig. 324).

Spiracle. The letters A and D, in connection with the figures of the spiracle,
indicate its orientation with regard to the body of the tick : A = anterior,
D = dorsal.

The explanation of the other terms and signs will be found in Part II.
pp. 127-132.

362

SPECIFIC DESCRIPTIONS

OF VALID SPECIES OF HAEMAPHYSALIS AND OF THEIR

VARIETIES

1. HAEMAPHYSALIS INERMIS Birula, 1895.

Figs. 309-312.

Lit., Synon. and Icon. :

Haemaphysalis inermis Birula, 1895, pp. 360, 361, PI. II, Figs. 7-9 ( 5 capitiilum,
showing porose areas, anus, coxa I ; evidently drawn from balsam-mounted
si)ecimen, therefore misleading) ; Latin description of $ only. Neumann,
1901, p. 264 ; gives French translation of Birula's description. Neumann,
1911 o, p. 116; listed as a doubtful species.

Haemaphysalis amhigua Neumann, 1901, pp. 262, 263 (not figured) ; the descrip-
tion given by the author agrees in all essentials ; he only refers to the 9 j
the description being based on 6 $ probably of French origin, from
E. Simon's collection. Neumann, 1906, p. 217 ; describes a more highly
chitinized 5 in which he thought he perceived a trace of eyes (specimen
in British Museum). Bomiet, 1908, p. 260 ; describes the 9 i" two line.s
and states that it probably does not occur in France, being Asiatic ; his
Fig! 30, of the ? capitulum, is bad. Blanchard, 1909, p. 148, Fig. 178 ;
species merely listed and figure reproduced from Bonnet. Neumann, 1911 a,
p. 109, states the species occurs in France.

Male (Fig. 309): Scutum 27xl-9 to l-74xl-24 mm.i, fairly glossy,
convex, dark, with very many small discrete j^unctations, rather linearly

' The scutums of 10 c? , collected by Brumpt in France, measured in mm. :

2-4 xl-6 2-2 xl-6

2 -38 X 1-7 2-08 X 1-3.5

2-3 xl-6 2-0 X 1-4.5

2-25 X 1-6 2-0 X 1-4

2-23x1-55 1-74x1-24

H. inermis

363

arranged towards the posterior end, and leaving a few non-punctate
islands, the largest being median and longitudinal ; no cervical or
lateral grooves ; festoons short and ill-defined. Capitulum very small,
and departing widely from the form normal to Haeniaphysalis; base
rectangular, with rounded angles, broader than long, deeply pitted on
its dorsal surface ; palps clavate ; article 1 easily visible dorsally ;
articles 2 and 3 not clearly separated, not salient at the base, broadest

Fig. 309. //. inermis i (N. 739 d). Dorsum, and more highly magnified detail of
capitulum and legs. Specimen found on fox at Surnabad, Transcaucasia, 1903, by
Dr E. Dschunkowsky. Original, N.C. del.

towards the distal end ; hypostome 2 [ 2. Venter: genital orifice between
coxae II ; spiracle very large, elongate, tapering only slightly. Legs
fairly long and strong ; all the coxae with a single short pointed spur,
internal on coxae I and IV, median on coxae II and III ; trochanter I
very large, with broad, blunt dorsal retrograde spur ; tarsus IV swollen
in the middle and tapering gradually ; pad short.

364

Greniis Haeniaphysalis

Female (Fig. 310) : Scutwn broader than long or the reverse,
1"24 to 0"9 mm. long\ with rather sinuous contour; punctations
numerous, uniform, discrete ; cervical grooves far apart, shallow and
inconspicuous. Dorsal surface of body closely and finely punctate ;
festoons distinct when unfed. Capitulum departing widely, as in the
J', from the normal Haemaphysalis form ; the base sub-rectangular but
rather salient postero-laterally ; no cornua ; porose areas very large,
deep, circular, separated by their diameter, a triangular pit in the
posterior part of their interval. Palps as in </ , but longer. Hypostome
3 j 3. Venter : vulva between coxae III ; spiracles large, short comma-
shaped; anal grooves ogival. Legs as in j/*.

"

:;C,\.

'.'.J

/ • 1 -'

• " /

.'»■

°« • ° -

\ A

"

-:?'••.

<!\

,■ ' '

Fig. 310. H. inermig ? (N. 788). Scutum and capitulum in dorsal and ventral aspects,
coxae with trochanters, spiracle and tarsus IV. Same origin as <? in Fig. 309.
Original, N. C. del.

The scutums of 19
1-24 X 10
12 xl-2
1-18 X 0-9
1.13x1-10
1-12 X 0-96
1-1 xl-1
1-1 xl-05
1-07 X 1-06
1 •07x0-95
1-0 xl-08

? , collected by Bnimpt in France, measured in mm.

0-98x1-44
0-97x1-15*
0-96x1-33
0-95x1-12*
0-94x1-16
0-93x0-95
0-9 xl-16*
0-9 xl-15
0-9 x 1-09

The bodies of 3 unfed ? (marked * in
the adjoining scutum-measurements)
measured respectively in mm. :

2-32x1-8

2-5 xl-85

2-57x1-8

H.

mermts

365

Nymph (Fig. 311): Body dark brown, broad, with large deep
punctations, uniformly distributed ; sharply defined median groove and
festoons ; about 16 x 13 mm., when unfed, attaining about 25 x 2"1 mm.
when replete. Scutum about 0"44 x 0"65 mm.S cordate, with slight
emargination, strongly shagreened in texture ; cervical grooves well-
marked to the posterior border. Gapitulmn : base fairly long, without
cornua but projecting laterally at its posterior border in rounded

Fig. 311. H. inermis o (N. 2762). Scutum, capitulum in dorsal and ventral aspects,
coxae, spiracle and tarsus IV. Specimen from France, received from Dr Brumpt, 1911.
Original, N. C. del.

prominences. Palps massive, rounded externally, widest at the base of
article 3. Hypostome relatively long and narrow, lanceolate, dentition
2 I 2. Veiitei^ : spiracle ovate, the narrow end directed dorsally. Legs :
coxae with short blunt spurs in the normal position ; the dorsal retro-
grade spur on trochanter I short and rounded ; tarsus IV tapering
gradually ; pad medium.

Larva (Fig. 312) : Bodi/ broad, with a few scattered punctations
and long, sharply defined festoons ; about 0"9 x 0'8 mm., when unfed ;

d bodies of 5 o.

collected by Brumpt, measured in mm

Scutum

Body

0-47 X 0-65

2-5 x2-l gorged

0-45 X 0-66

1-3 X 1-12 unfed

0-44 X 0-65

1-45 X 1-2

0-43 X 0-66

1-6 xl-36 „

0-43 X 0-60

1-6 xl-3

24

366

Genus Haemaphysalis

attaining about 1'8 x 1-3 mm. when replete. Scutum about 0'3 x 0"45 mm.\
like that of the o, but with shallow, more parallel cervical grooves.
Capitulum : base without cornua or lateral projections, its antero-lateral
edges rounded. Palps as in the o, except that articles 2 and 3 appear
fused. Hypostome and Legs as in the o.

Our description of this species is based on the examination of many
specimens chiefly derived from France (vide infra). The type of the
? is in Birula's collection, Petrograd, those of the {/•, o and L are

Fig. 312. H. inermis larva (N. 2757 a). Scutum, capitulum in dorsal and ventral aspects,
coxae and tarsus III. Specimen raised by Dr Brumpt. Original, N. C. del.

in Cambridge (N. 789, 2762, 2757 a), the latter stages not having
hitherto been described. The species is so remarkable that Neumann
(1901, p. 263) stated that he would have referred it to Aponomma were
ti not for the resemblance to Haemaphysalis in the leg structure.

Geographical Distribution and Hosts.

Caucasia : We are indebted to Prof A. Birula for (N. 1226) a $
received xi. 1910. Birula (1895, p. 360) founded the species on
1 $ supposedly derived from the Caucasus, the host not being recorded.
Transcaucasia : We have examined many specimens collected at
Surnabad by Dr E. Dschunkowsky who kindly allowed us to retain
(N. 781) 6 $ found on cattle, and (N. 788, 789) 3 cT 1 ? found on focc in

1 The scutums and bodies of 4 L, collected by Brumpt measured in mm. :
Scutum Body

0-3 xO-47 09 xO-8 unfed

0-3 xO-45 0-86 X 0-8

0-3 X 0-43 1-8 X 1-3 gorged

0-28 X 0-42 0-85 X 0-73 unfed

H. inermis 367

company with H. cinnaharina var. 'punctata and Boophilus. France :
Neumann (1901, p. 262) appears to have been the first to record the
species from France, describing it under the name of H. ambigua, of
which 6 $ were collected in 1894 by E. Simon. Neumann has pre-
sented us with a co-type (N. 2881). Dr E. Brumpt has sent us his
unpublished records relating to the occurrence of the tick on deer, on
which he found adults on 13 occasions (9 times with H. concinna and
once with H. cinnaharina var. punctata) at Fontainebleau, D^pt. Seine-
et-Marne, and in the Depts. Vienne and Indre in Western and Central
France respectively. We have examined all of Brumpt's material,
including specimens which he raised in the laboratory, and he has
allowed us to retain numerous examples of all stages (N. 1525-1528,
2752, 2753, 2756-2759, 2762); further particulars regarding them will
be found in the Section on Biology. [Japan: Neumann (1906, p. 217)
records 1 $ from Naemorhaedus crispus Temm., found in company with
Ixodes and Haemaphysalis spp.^]

See further under Section on Biology (p. 545). The biology of the
species is remarkable.

Haemaphysalis inermis var. aponommoides
Warburton, 1913.

Fig. 313.

Lit. and Icon. ; Warburton, vii. 1913, pp. 128-130, Fig. 8 (reproduced).

Male : unknown.

Female (Fig. 313): Scutum: broader than long, 1 x 1"4 mm.,'
broadest at the anterior third, and much more narrowed posteriorly
than in the type ; glossy, but with numerous discrete, medium-sized
punctations ; cervical grooves broad sub-parallel tracts without any
initial pits ; eniargination slight. Capitulum : even more ApononiJiia-
like than in the type ; base bluntly salient at the sides, almost
destitute of cornua ; porose areas large sub-circular, bounded ex-
ternally and anteriorly by sub-rectilinear ridges, the interval about
equal to their diameter; palps long and narrow, their outer border
straight and not convex as in the type ; article 1 well visible ; articles
2 and 3 not distinctly separated, article 2 corrugated dorsally, and
without any trace of lateral salience. Hypostome spatulate, dentition

^ Probably //. iiietinis var. aponommoides, q. v.

24—2

368

Genus Haemaphysalis

3 I 3. Venter: spiracle short comma-shaped, the narrow end postero-dorsal ;
anal grooves semicircular. Legs : as in the type ; coxa I without definite
spur, but with its internal end truncated and the corners slightly

Fig. 313. H. inermis var. aponommoides ? . Capitulum and scutum, profile of right palp,
coxae, spiracle and anus with anal grooves. (N. C. del.) Warburton, 1913, Fig. 8.

protuberant ; coxae II-IV with a slight protuberance in the middle
of the posterior border ; tarsus IV rather long and tapering (as in
H. hoiuletti $, see Fig. 434).

Described from (N. 1566) 23 $ taken from a Himalayan Zebu
(Bibos sp.) at Belgachia, Calcutta, India, ill. 1912, in company with
Boophilus australis, by Col. F. Raymond, F.R.C.V.S. We have since
determined (Berlin Mus. 173) 1 $ found on a horse at Fukoka, Japan.

Types in Cambridge ; we have presented co-types to the Berlin
Museum and to Prof Neumann's collection in Toulouse.

H. inermis Birula, 1895, has hitherto been considered the most
aberrant form of Haemaphysalis, and its attribution to that genus
would be extremely doubtful were it not for such intermediate forms
as H. warburtoni Nuttall, 1912. The present variety still further
departs from the normal Haemaphysalis type, and still more closely
approaches Aponomma. The genus Haemaphysalis and sub-genus
Aponomma have normally little in common, except the negative
characteristics of the absence of eyes and anal plates, and we should

H. inermis 369

not have expected any question to arise as to which of them a newly
discovered form belonged. Yet a grave question does arise in the
present case, and though wc judge the tick here described to be a
Haemaphy salts, its resemblances to an Aponovima are more than
superficial. It possesses one Haemaphysalis characteristic never found
in Aponomma — the blade-like dorsal retrograde spur on trochanter

1. Moreover, it is found on a mammal, whereas Aponomma is
essentially parasitic on reptiles. The long palps need not trouble
us much, as we know of several species of Haemaphysalis almost
equally aberrant in this respect^ ; and, though the palps are long
and very Apoiiomma-Mke when viewed dorsally, they do not recall that
genus when viewed in profile (see Fig. 313). The Aponomma-\ike
shape of the scutum, also, is of little importance, for there is a great
range of scutal design in Haemaphysalis. It must be admitted,
however, that not only the dentition (3 | 3) but the whole appearance
of the hypostome is more like that found in Aponomma than in
Haemaphysalis, and that coxa I, though not distinctly bifid, displays a
tendency in that direction.

2. HAEMAPHYSALIS WARBURTONI NuttaU, 1912.

Figs. 314, 315.
Lit and Icon. : NuttaU, 1912, pp. 55-57, Figs. 5, 6 (reproduced).

Male (Fig. 314) : Scutum 25 x 1*8 to 2*3 x 1*6 mm.-, narrow in
front, broadest on a line with the spiracles ; cervical grooves short, con-
vergent pits ; lateral grooves include two festoons and extend forwards
to I the body-length ; posteriorly a median groove and two lateral
depressions, two longitudinal grooves anterior to the latter extending
forward to ^ the length ; festoons short ; punctations few, incon-
spicuous. Capitulum : 0'4 to 0"5 mm, long, base sub-rectangular, with
concave dorsal ridge connecting stout, somewhat convergent cornua

^ See H. loarburtoni (Fig. 314), leporis-palustris (Fig. 326), aculeata (Fig. 371),
loelUngtoni (Fig. 417), humerosa (Fig. 436).

2 Five (? gave the following measurements in mm. :

Capitulum
Scutum (length, measured dorsally)

2-5 X 1-8 0-5

2-4 X 1-7 0-5

2-4 X 1-7 0-5

2-4 X 1-6 0-5

2-3 X 1-6 0-4

370

Genus Haemaphysalis

having rounded points ; the base bulges ventrally ; palps longer than
broad, being broadest at the distal end of article 2, which is about ^
longer than article 3 ; hypostome broad, 4 j 4, with corona followed
by 8 distinct teeth per file. Venter : genital orifice between coxae II ;
spiracle longer than broad, with recurved dorsal margin. Legs short.

Fig. 314. Haemaphysalis ivarhiirtoni, s . Capitulum in dorsal and ventral aspects,
scutum, spiracle, trochanter I seen from in front (and, attached to body, from above),
coxae, tarsus IV. (G. H. F. N. and E. W. del.) Nuttall, 1912, Fig. 5.

stout ; coxae I-IV each bearing a stout spur, longest on coxa IV and
concave externally ; trochanter I with a very large dorsal blade ; tarsi
remarkable, the distal portion bulging dorsally and ventrally, tapering
rapidly, bearing a spur ; pad half as long as the claws.

Female (Fig. 315) : Body, unfed, 2'3 mm. long' with marginal groove
including the second festoon and almost attaining the scutum. Scutum :

1 None of the ? were fully gorged, the most swollen specimen measured 6*1 mm.
in length.

Capitulum

Scutum (length, measured dorsally)

1-6 X 1-4 0-9

1'5 X 1-6 0-8

1-5 X 1-6 0-9

1-2 X 1-6 0-8

H. warhurtoni

371

1*6 X 1-2 to 1-2 X 16 mm., cordiform, cervical grooves not attaining the
posterior border, postero-lateral border almost straight, few incon-
spicuous punctations. Gajntulum : 08 to 0-9 mm. long, base broader
than long, with sides angular, the an tero- lateral borders converging,
dorsal ridge concave, wavy, connecting short stout cornua ; porose areas
long-oval, converging anteriorly, far apart, in some specimens separated
by a median depression ; viewed ventrally the base bulges markedly.
Palps atypical, very long, with article 1 distinctly visible dorsally, article
2 about twice as long as 3, the palps being broadest where articles
2--3 join. Hypostome broadly spatulate, 4 | 4 or 5 | 5, with emarginate

Fig. 315. Haemaphysalis warburtoni ? . Capitulum in dorsal and ventral aspects,
scutum, coxae, spiracle and tarsus IV. (G. H.F.N, and E.W. del.) Nuttall, 1912,
Fig. 6.

corona, about 10 distinct teeth per file. Venter : vulva between
coxae II when unfed, facing second intercoxal spaces when replete ;
spiracle as long as broad, somewhat angular, with slightly marked
postero-dorsal angle. Legs resembling those of the (^, the spur on
coxa IV less developed.

Described from (N. 1400) 5 ^^ and 9 ? taken from Seroiu goat,
at Wen-chwan-hsien, near Si-ho-hsien, China, and purchased, vii. 1911,
from Mr T. V. Sherrin, Taxidermist, Hampton. We have since

372 Genus Haemaphysalis

received (N. 2914 c), 1 (/ taken from cattle at Taihoku, Formosa,
16. VII. 1909, coll. Dr M. Miyajima, and have determined (Berlin
Mus. 160) 1 % from Tscholesmantal, Altai Mts., Western Siberia,
10. IX. 1907, coll. C. Wache and Dr Biedermann.

Types in Cambridge ; we have presented co- types to Neumann's
collection, Toulouse.

3. HAEMAPHYSALIS CINNABARINA Koch, 1844.

Figs. 316-320.

Lit., Icon, and Synon. :

Haemaphi/salis cinnabar ina C. L. Koch, 1844, p. 237 ; 1847, p. 123, PL XXVI,
Fig. 97, $ . When in Berhn in 1909, one of us (G. H. F. N.) examined and
drew the unique type from Brazil, and, after consultation with the late
Prof. Donitz, concluded that it was closely allied if not identical with
H. punctata Canestrini and Fanzago. We have since been able, however, to
identify it with the American form known as H. chordeilis (Packard) which
therefore falls into synonymy. Koch sjjelt the name correctly at first (1844),
but later (1847) misspelt it cinnaberina, doubtless having in mind the German
word " Zinnober " ; the original spelling has not been followed by other
authors hitherto ^ The body of the type specimen is still red, like cinnabar.
Koch's original (coloured) figure is scarcely recognisable as that of a Haema-
physalis.

Haemaphysalis sanguinolenta C. L. Koch, 1844, p. 237 ; 1847, p. 124, PI. XXVII,
Fig. 98, 9 • The coloured figure is poor, the scutum too elongated. Nuttall
examined the unique type in the Berlin Museum in 1909, and identified
it with H. cinnabanna ; the scutum is slightly more elongate than in the
type. The specimen came likewise from Brazil.

Ixodes chordeilis Packard, 1 869, p. 67.

Haemaphysalis punctata var. ciyinaberina (Koch) Neumann, 1905, p. 237.

Haemaphysalis punctata cinnaberina (Koch) Neumann, 1911 a, p. 108.

Haemaphysalis chordeilis (Packard) in Banks, 1908, p. 34, PI. IV, Fig. 11
(desci'ibes the $ ; figures capitulum and scutum, coxae I and IV, tarsus IV,
spiracle ; he studied the types). Banks, 1908, p. 54 (listed). Hunter and
Hooker, 1907, p. 53 (casual mention). Hooker, 1909, p. 423, brief reference
to distribution.

'•'■Haemaphysalis chordeilis Banks, 1908," only listed by Blanchard, 1909, p. 148.

Haemaphysalis chordeilis (Packard) listed as a doubtful species by Neumann,
1911a, p. 115.

'■'■Haemaphysalis chordeilis Packard," Hadley, 1909, p. 606 ; recorded as killing
turkeys in Vermont. Hunter and Bishopp, 1911, p. 229. Bishojip, 1911,
pp. 207-208, describes $ (not figured) and raised the species. Hooker,
Bishopp, and Wood, 1912, pp. 97-102, PI. VII, Figs. 7-10 {$ dorsum

1 Neumann, 1897, p. 331 and other authors.

H. cinnaharhia

373

and venter ; o replete, dorsum and venter ; all .stages described, hosts

distribution, biology).
Haemaphysalis punctata Cane.strini and Fanzago, in I lad wen, 1912, p. 98.
Not Haemaphysalis leporis-palustris (Packard), as stated by Neumann, 1897,

p. 343.

Male (Fig. ^\Q): Scutum: l-.Q x 1-3 to 2-7 x 1-5 mm., elongate,
strongly and irregularly punctate, the punctations being coarse and
often confluent in the middle region ; cervical grooves very short, well
marked ; lateral grooves very long and deep, including three festoons ;
festoons short ; dorsal, median and postero-lateral grooves are present,

Fig. 316. H. ciiinabarina s . Dorsum, ventral aspect of capitulum, coxae with tro-
chanters, spiracle and tarsus IV. (Specimen, N. 2452, raised in Cambridge, parents
from cattle, Winnipeg, Canada.) Original, N. C. and E.W. del.

recalling those of Rhipicephalus. Capitulum : base not much broader
than long, punctate, with strong cornua ; palps massive ; article 2 slightly
salient laterally ; no dorsal spines ; a slight ventral retrograde point
under article 3 at its inner angle ; hypostome : dentition -t | 4 to 5 | 5 ;
very small sharp teeth without any median interval. Vente?- : spiracle
large, short, comma-shaped. Legs : coxal spurs strong, longest on coxa
IV ; tarsus IV short, tapering rather abruptly.

374

Grenus Haemaphysalis

Female (Figs. 317, 318): Scutum: 0-9 x 09 to 16 x 1-2 mm.,
usually distinctly longer than broad, broadest near the anterior border,
punctations coarse ; cervical grooves deep and converging to the middle

Fig. 317. H. cinndbarina ¥ . Scutum, capitulum in dorsal and ventral aspects, coxae
with trochanters, spiracle and tarsus IV. (Same source as s in Fig. 316.) Original,
N.C. del.

Fig. 318. H. cinnabarina ? . Scutum and capitulum in dorsal aspect (N. 2833, received
from Mr F. C. Bishopp, Victoria, Texas). Original, N.C. del.

of the scutum, then shallower and diverging. Capitulum : base twice
as broad as long ; cornua very slight and blunt ; porose areas large,
almost reaching the posterior border, not always definite, except where

H. cimidbarma

375

bounded by the lateral 7'idge ; but in some specimens distinctly oval,
converging in front ; palps and hypostome as in the </. Venter:
spiracle large, sub-circular, with distinct dorsal process. Legs : as in
the (^ , except that the spur on coxa IV is short. (Replete specimens
may attain 9 x 6'6 mm., according to Hooker, Bishopp and Wood.)

Nymph (Fig. 319) : Scutum: 0-62 x 06 to 0*45 x 0-o5 mm., variable,
generally cordate, with few punctations and well-marked cervical
grooves. Capitulum : base liexagonal, with lateral angles, and with
ventral cornua ; palps recalling those of the adult ; dentition of

Fig. 319. H. cinnabarina, nymph. Capitulum in dorsal and ventral aspects, scutum
(variations in shape indicated by second contour), coxae with trochanters, spiracle,
tarsus IV. (Same source as j in Fig. 316.) Original, N.C. del.

hypostome 2 | 2, about 8 teeth per file. Venter : spiracle ovoid, with
distinct dorsal process. Legs : as in the $ , with coxal spurs unusually
well-marked for a nymph.

Larva (Fig. 820) : Scutum : 0-24 x 0-3 to 0-21 x 0-26 mm., cordate
broader than long ; cervical grooves fairly distinct, parallel. Capitulum:
base with lateral angles and with distinct ventral cornua ; palps with
slight lateral salience ; hypostome : dentition 2 | 2, about 7 teeth per
file. Legs : coxa I with slight spur, coxa II flanged, coxa III unarmed ;
tarsus III tapering.

Our description is from specimens reared in Cambridge, being
the progeny of (N. 2452) 2$s, from cattle, Winnipeg, Canada, collected
by Mr J. R. N. Harrison. It is evident, however, from other specimens
since received from Mr F. C. Bishopp (N. 2833, 2834, 2 (/■ 2 $, from
Sturnella magna, Victoria, Texas, 1909), that well-developed and

376

Genus HaemapUy sails

strongly chitinised individuals may depart rather widely from the
description here given, the tendency in such specimens being towards
elongation. The larger measurements we give relate to the specimens

Fig. 320. R. cinnabarina, larva. Capitulum in dorsal and ventral aspects, scutum,
coxae, tarsus III. (Same source as c? in Fig. 316.) Original, N. C. del.

received from Mr Bishopp, see Fig. 318 showing the long scutum of
the ?.

Type, a dried $ , from Brazil, in the Berlin Museum. The types
(2 $ ) of Packard's H. chordeilis are in the Museum of Comparative
Zoology, Harvard University, Cambridge, Mass., where they were
examined by N. Banks; J' in Bishopp 's collection, o and L in
our collection.

Geographical Distribution and Hosts.

H. cinnabarina appears to be^ widely distributed on the continent of
North America. The specimens we raised in Cambridge (N. 2452)
were derived from replete females found on cattle in Winnipeg,
Canada, ix. 1913, by J. R. N. Harrison; similar specimens (N. 934)
were found by Dr S. Hadwen, xi. 1909 (same host and place). We
have furthermore received (N. 3023) c/s ?s from cattle, Ashern,'
Manitoba, 17. ix. 1914, J. R N. Harrison coll.; (N. 3026) 2^/2$ from
cattle, Stonewall, Manitoba, 14. ix. 1911, J. D. Ross coll. and (N. 3027)
a J" found attached to the arm of a man, at Kamloops, British Columbia,
IX. 1914, both lots having been presented by Dr S. Hadwen. Packard's
types (two replete $s) were found on the night-hawk {Chordeiles
popetue), at Milton, Massachusetts. Banks (1908, p. 34) records 1 %
from a turkey, Taftsville, Vermont. Hadley (1909, p. 606) states that
in June, 1909, this tick was observed on two farms at Norwich, Vermont,

H. cmnabarina 377

in May, where it killed off 40 out of 46 young turkeys before the ticks
were finally destroyed by hand-picking. Many immature and adult
ticks, the latter to the number of 70-80 per bird, were found attached
chiefly to the birds' necks ; the ticks were determined by Banks.
Hooker, Bishopp and Wood (1912, pp. 97-102) state that the tick
chiefly infests ground-inhabiting birds, the immature stages usually
occurring on the top of the birds' heads, around the eyes and ears,
and at times beneath the bill. It is often found together with
H. leporis-palustris. Other hosts are the meadow-lark, jackdaw, red-
winged blackbird, marsh-hawk. In Texas the meadow-lark is chiefly
infested ; Hunter (vi. 1909, p. 252), with J. D. Mitchell, found what
appeared to be the O and L on a quail {Colinus virginianus) at
Brownsville, in the autumn of 1907, and o and L (also L skins)
were found on the heads of quails and larks (Stmmella magna neglecta)
at D'Hannis (an adult which developed from a replete o was identified
a,sH. chordeilishy N. Banks). From 12 quails dropped into bags after
•being shot in Victoria County in December, 1907, Mitchell reports that
over 500 ticks were collected: "that there were 1000 ticks upon the 12
quail would be a conservative estimate." The author adds that some
of these ticks may have been H. lejyoris-palustris. We have received
specimens (N. 2833, 2884) from the meadow-lark (Sturnella magna),
Victoria, Texas, 1909, presented by Mr F. C. Bishopp. Bishopp
(VI. 1911, p. 208) states that J. D. Mitchell found the tick three
times on meadow-larks in Victoria County, Texas : 1 </ on 23. ill.
1910; 1 c/ 4 ? on 12. xi. 1911; 5 cf 4 o 23 Z on 25. xi. 1911.
In the autumn, nearly all the ground-feeding birds are infested with
the O and L as observed by Mitchell in Victoria County, and at Grand
Cane, Louisiana and in Quincy and Hawthorne, Florida. Hooker
(1909, p. 423) states that the tick occurs also in the State of New
York; we have received (N. 2737) 1 ? from ruffed grouse, Catskill
Mts., VIII. 1909, presented by Dr L. O. Howard.

As previously stated, Koch's type $ came from Brazil.

Remarks regarding H. cinnabarina, " H. punctata"
and " H. chordeilis."

It was only after having described and drawn the forms known
commonly as H. punctata in the Old World and H. chordeilis in the
New that we became convinced that their differences were merely
varietal. This conclusion, together with the undoubted priority of the

378 Genus Haemiaphy sails

name cinnabarina, has forced upon us a change of nomenclature which
we only adopt with reluctance in view of the large amount of literature
connected with the name H. punctata. The specimens of "H. chordeilis "
to which we first had access did not, in their general facies, strongly
recall "H. punctata," though when examined point by point their
differences were unimportant.

Koch's type of //. cinnabar^ina is a dried specimen not well preserved,
and it was with some hesitation that we admitted its identity with
"H. punctata." This hesitation is explained as soon as it is recognised
that there are two varieties of the species, and that the type specimen,
which came from Brazil, belongs to the American variety. The
American form thus necessarily becomes the type variety. Under the
circumstances we have thought it better to depart from our usual
procedure in dealing with a variety, and instead of merely indicating
the points which distinguish it from the type form, we have allowed to
stand the full description we had prepared of H. punctata — now
H. cinnabarina var. punctata — when we believed "H. chordeilis" to he-
generically distinct.

Varietal Differences : The main differences between H. cinnabarina
and H. cinnabarina var. punctata will be seen at a glance by reference
to the figures. In the type the body of the (/ is broader, the dorsal
furrows more distinct, palpal article 2 more protruding, the spur on
coxa IV straighter. In the $ palpal article 2 has a more sharply
protruding external angle. In the O and L the differences are slight.

Haemaphysalis cinnabarina var. punctata
(Canestrini & Fanzago, 1877).

Plates VIII-XI and Text-figs. 321-324.

Lit,, Icon, and Synon. :

Hae7naphysalis punctata Canestrini and Fanzago, 1877, p. 121 (reprint) and
1877-1878, p. 189. The original description refers only to the ^ and 9 and

is so brief that it would be impossible to recognise the species by it. 1890.

Canestrini, pp. 523, 525, PI. XLI, Figs. 6 and Qa., gives a fairly accurate
description, especially of the $ ; the figures of the ^ venter and capitulum
are inaccurate but show the main characters. All the points in his
description, except a few measurements, are included in Neumann, 1907,

pp. 327-330. 1891, Berlese, fasc. 58, PI. X, gives an inaccurate coloured

figvire of the ^ , also poor figures of the ^ venter, capitulum, palp, spiracle
and of the $ dorsum venter and spiracle. 1895. Pocock, p. 326, records

H. cinnaharina var. punctata 379

it in England. Raillict, p. 714 ; brief mention.— —1897. Neinnann, p. 237,
Figs. 1, 2 gives the first good description, the ilUistrations, however, only

relate to the hypostome ( ^ ) and digit ( ^ $ ). 1905. Nuttall, Cooper and

Smedley, pp. 439-441, give a desci'iption f)f the buccal apparatus. Cooper,
pp. 3, 4, describes 19 accompanying photomicrographs of all stages showing
details of the mouthparts, microscopic sections, etc., which served to illustrate
the foregoing paper by Nuttall, Cooper and Smedley. Lahille, p. 44 ; merely

lists the species. 1906. Wheler, p. 421, PI. X, Figs. 31, 32 (photographs);

describes all stages briefly and records it on sheep and hedgehog in England.
The figures of the ^ and 9 show the general character. He gives some

accurate measurements. 1907. Coward, p. 323 ; states that Oldham

found it at Dungeness in England. Donitz, p. 71 ; quotes Neumann.

1908. Bonnet, p. 261 ; poor description, condensed from Neumann, 1897 ;
his Fig. 29 of $ capitulum is original but bad. Nuttall (VII), pp. 16-18
and (VIII), pp. 398-399, describes and figures the process of oviposition.
Nuttall, Cooper and Robinson, pp. 152-181, Pis. XII-XVI, Text-Figs. 1-9 ;
give the biology, synonymy, iconography, distribution and a full illustrated
account of the external structures of all stages together with a bibliography
to 1908 inclusive. Nuttall (IX), pp. 514, 522 ; biology and part jjlayed in
disease transmission. Nuttall, Cooper and Robinson (X), pp. 238-242,
PI. XVIII, Text-Fig. 1 ; describe and illustrate the structure of Haller's
organ. Nuttall, Cooper, and Robinson (XII), pp. 347-351, Pis. XXII,
XXIII ; describe and figure the structure of the spiracle. Stockman
(VIII), reprint, p. 8, records the species on cattle and sheep and demon-
strates by experiment that it may convey piroplasmosis (redwater) to

cattle. 1909. Blanchard, pp. 154-157, Figs. 192-194, gives some more

recent data regarding distribution ; his figures ai"e taken from Bonnet

and Neumann, q.v. Rohr, p. 142 ; quotes Neumann. 1911. Knuth,

reprint 12 pp., illustrated by figures taken from Nuttall, Cooper and
Robinson, q.v. ; records its presence on cattle in Northern Germany.
Nuttall (X), p. 180, illustrates its type of parasitism graphically. Stockman,
pp. 23-32 ; account of the biology. Yakimoff" and Kohl-Yakimoft', p. 418 ;

record occurrence in Gouv. Cherson, Russia, and in Caucasia. 1912.

Eysell, Figs. 5, 6 ; buccal apj^aratus, copied from Nuttall, Cooper and

Robinson, q.v. 1913. Nuttall (IV), pp. 99-105; full account of the

biology (incorporated in our text). Patton and Cragg, p. 630 ; description
translated from Neumann 1911, q.v. ; p. 648 brief reference to Nuttall and
Stockman's experiments.

Haemaphysalis sulcata Koch, in Canestrini and Fanzago, 1877, p. 120, may well
be included here, although doubtfully, the description being somewhat vague.
Berlese, 1889, fasc, lv, N. 1 describes it. Berlese, 1891, fasc. lv, PI. I,
figures a replete o inaccurately, the capitulum shows the chelicerae situated
ventrally.

Rhipicephalus expositicius L. Koch, 1877 ft, pp. 196-198, certainly agrees very
closely in its description with that of H. cinnaharina; the author gives
no figure. We agree with Neumann, 1897, p. 327, in condemning the
species.

380 Genus Haemaphysalis

Haemaphysalis rhinolophi Canestrini and Fanzago, 1877, p. 189 ; Canestrini,
1890, p. 526 ; Neumann, 1897, p. 332. According to Neumann this is a
good species distinguishable from H. punctata through having a non-
punctate scutum. One of us (G. H. F. N.), in 1910, examined the only
existing type specimen through the courtesy of Prof. Carazzi of the Zoo-
logical Institute, Pixdua, a $ without capitulum and anterior third of the
body and with only three legs remaining. The specimen proved to be
only a poorly chitinised H, cinnaharina var. punctata.

Haamaphysalis punctata punctata Canestrini and Fanzago in Neumann, 1911a,
pp. 107-108, Figs. 50, 52. The figures show the ^ in dorsal and ventral
aspects, the palp seen ventrally.

Haemaphi/salis o-assaWsivhuvton, 1908, pp. 516-517, Fig, 8 (reproduced); see
discussion in our text.

Not Pseudixodes holsatus (Fabricius) in Haller, 1882, p. 311, PI. V, Fig. 5
{$ capitulum) as stated by Canestrini, 1890, pp. 485, 526, who includes it
in his synonymy oi H. punctata ; Neumann, 1897, p. 360, appears justified
in referring the tick to the synonymy of Dermacentor reticulatus (Fabricius
1794).

Male (Text fig. 321, Plates IX, X^, Figs. 3, 4 ; XI, Figs. 2, 5, 6) :
Scutum : elongate, narrowed in front ; size variable, averaging about
3x2 mm. but ranging from 3*3 x 2*0 down to 2*4 x 1"5 mm.'-; colour
usually dark brown ; cervical grooves rather short, concave externally ;
lateral grooves very long, beginning at the level of legs II, and
including from one to three festoons ; punctations very numerous and
small over the whole surface, some of them confluent ; two lateral
furrows (in ungorged specimens), sometimes resolved into a series
of shallow pits, and often connected posteriorly by three shallow pits
in front of the festoons ; festoons short and ill-defined ; emargination
deep and abrupt. Capitulum small ; base rectangular, about twice
as broad as long, punctate ; cornua short and blunt ; palps short,
the lateral salience slight, not angular, but rounded ; article 1 hardly
visible ; article 2 massive, and longer than article 3 ; the inner contour
of articles 2 and 3 rectilinear, and their dorsal surface somewhat
corrugated ; no dorsal spurs, but a strong ventral retrograde spur under
article 3 ; hypostome 5 j 5 (with sometimes an extra pair of files for
part of the length). Venter : spiracles large, elongate, with blunt dorsal
process ; anal grooves very slightly ogival. Legs strong, coxa I short,
with blunt internal spur ; coxae II and III with a blunt spur in

1 For description of Plates see p. ix.

2 Variation determined on over 60 <? collected in England and Schleswig-Holstein,
or raised in Cambridge. The proportion of length to width varies, tlius the scutum may
measure 3-0 X 1-8, 2-95 x 1-7, 2-9 x 1-8, 2-9 x 1-6, 2-6 x 1-4, 2-4 x 1-5 mm., etc.

H. cinnabarina vjir. punctata

381

the middle of the posterior border ; a long, strong, sharp, inwardly
curved spur on coxa IV, as long as the article itself (see note on

Fig. 321. H. cinnabarina var. punctata <? . Dorsum, part of venter, spiracle and tarsus
IV. Specimen (N. 2708) from Germany. Original, G. H. F. N. del.

Transcaucasian forms, p. 384) ; trochantal spurs faintly indicated ;
tarsus IV, small and tapering ; pads short.

Female (Text figs. 322, 323 ; Plates IX ; X, Figs. 1, 2 ; XI, Figs. 1, 4) :
Body, when unfed, measures 2'7 to 3'2 x 1"8 to 2 mm., attaining, when
fully gorged, 13 x 8 mm. Scutum about 1'3 x 1*2 mm., ranging from
1'4 X 1*2 down to 1*25 x 1 mm.\ sub-cordate, but the evenness of the
contour usually broken by slight postero- lateral angles; cervical
grooves well-marked for about two-thirds of the scutal length ;
punctations medium, irregular, not very numerous. Capitulum : base
rectangular, nearly thrice as broad as long, without cornua ; porose
areas large, rather indefinite, far apart, with a slight depression in
the interval ; palps with lateral salience more angular and more
posterior than in the (/ , and with shorter spur under article 3 ;
hypostome more spatulate than in the </, with similar dentition,
ordinarily 5 [ 5, but frequently 6 ; 6. Venter : spiracle rounded, with

' As in the d , the proportions vary. The range, determined on about 50 ? , varied
being 1-4 x 1-2, 1-3 x 1*2, 1-3 x 1, 1-25 x 1-1 mm., etc. Nuttall, Cooper and Kobinson (1908,
p. 1.57) record the length as 1-08-1 -37, the breadth as 1-05-1 '31 mm.

382

Genus Haemaphysalis

short dorsal process and large macula ; anal grooves slightly ogival.
Legs : coxae I-III armed as in the ^ , but the spur on coxa IV is only
represented by a spur somewhat stronger than on coxa III.

Fig. 322. Fig. 323.

Fig. 322. H. cinnabarina var. punctata ? . Dorsum, coxae, spiracle and tarsus IV.

Specimen {N. 2706) from Germany. Original, G. H. F. N. del.
Fig. 323. H. cinnabarina var. punctata ? . Capitiilum and scutum, spiracle and tarsus

IV, Warburton 1908, Fig. 8. Sketch of fZ". crassaWarb. Specimen from Transcaucasia

with unusually rugose scutum.

Nymph (Text fig. 324 ; Plates IX ; X, Figs. 5, 6) : Body : 1-32 x 0-85
to 1-26 X 08 mm. Scutum : 0*37 x 0*4 up to 0"o5 x 057 mm.\ as broad
as long or broader than long ; like that of the $ but less angular and
appearing broader in proportion to its length, with very faint punctations ;
cervical grooves long, broad, sub-parallel. Capitulum : base hexagonal
dorsally, the sides being produced to sharp lateral points; ventral cornua
but no dorsal cornua ; hypostome 2 | 2 ; palps like those of the $ but
with article 2 more sharply recurved at its lateral angle and less clearly

^ The scutums of 5 O raised in Cambridge (N. 2780) measured in mm. :
0-57 X 0-57 0-5 X 0-53

0-53 X 0-53 0-5 X 0-51

0-5 xO-54

H. cinnabarma var. punctata

383

marked off from article 3. Ventei': spiracle sub-circular. Legs as in
the % , except for the tarsi (see Figs.). When gorged, the nymph may
attain a length of 3 mm.

Larva (Plates VIII ; X, Figs. 7, 8 ; XI, Fig. 3) : Body, when unfed,
measures about 0-50 x 0-45 mm., when fully gorged the larva may
measure 1'6 x 1 mm. Hciituin broad in front, 025 x 0"32 to 0"23
X 0'3 nnn., narrowing and rounded posteriorly ; cervical grooves

Fig. 32i. H. cinnabarina var. punctata Q. Dorsum, ventral aspect of capitulum, coxae,
anus and anal grooves, spiracle and tarsus IV. Specimen (N. 814) raised in Cambridge.
Original, G. H. F. N. del.

deep and parallel. Capituliau : base wiuh slight lateral points ;
hypostome 2 | 2 ; palps like those of the O.

Described from numerous specimens collected and raised experi-
mentally by us in England.

See further under Notes on Biology, p. 518.

25-2

384 Genus HaentapJiysalis

NOTE ON TRANSCAUCASIAN FORMS OF
H. GINNABARINA.

In the collection of ticks left with us for determination by Dr
Dschunkowsky of Surnabad, we have found a number of specimens
of typical H. cinnaharina var. punctata intermixed with forms in
which the characteristic long spine on the fourth coxa of the </• is
either reduced in size or practically absent ; these specimens conform
to the type in other respects. Details regarding these specimens
will be found in the section relating to Asia under Geographical
Distribution (see p. 887). One lot especially (N. 780) contained
transition forms : a typical long-spined (/, 1 cT with a moderate
spine and 5 (/s with very short spines. We have received precisely
similar specimens from Smyrna (N. 2554) 1 (/• 2 $ taken from sheep,
but the cf had the usual long spine.

Haeinaphy salts crassa Warburton, 1908. While examining the
ticks in the British Museum in 1907, we found, in a tube labelled
" No. 254. H. papuana," two gorged $ s which certainly did not
belong to this species and which we were then unable to identify.
One of us described them under the name of H. crassa (see Fig. 323),
in allusion to their unusually rugged appearance. They were taken
from cattle at Surnabad and presented to the Museum by Dr
Dschunkowsky to whom Ave have since been indebted for the oppor-
tunity of studying all the material he collected in Transcaucasia.
After due consideration we believe that H. crassa represent only
aberrant forms of H. cinnaharina var. punctata.

Geographical Distribution and Hosts.

Although widely distributed, H. cinnaharina var. punctata is a
relatively uncommon tick. It occurs in Europe, Asia, and Africa as
the following records show. [Records b}- other authors are enclosed
in square brackets.]

EUROPE : England : We have received all our adult specimens
from sheep in Kent, as follows: (N. 2151) Lydd, 13. in. 1905, S. T.
Sellens coll. ; (N. 899 and 1564), ditto, iii. and iv. 1905 ; (N. 2780)
ditto, 25. V. 1914 ; (N. 2152) Littlebourne, 5. iv. 1905 ; (N. 1565)
Heme, lY. 1905 ; (N. 1207) Ashford, v. 1900, Mr Spanton coll. ; all
of these specimens were collected at the instance of Mr W. F. Coojjer.
Prof. R. T. Hewlett has presented (N. 304) specimens from sheep.

H. cinnaharina var. punctata 385

found at New Roimiey, iv. 1907, and Mr J. Davidson found (Liverpool
No. 5) speciiiiciis on sheep at Lydd, ill. 1910. [Pocock, 1900, p. 326,
reports that F. Pickard-Cambridge collected specimens from shingle
on the beach and from hedgehog at Dungeness. Coward, 1907, p. 323,
records the finding of a gorged $ on Dungeness beach, Kent, in
the nest of a plover, Oedienus scolopax, the tick having been found
lying on the ground beneath two young birds by C. Oldham, vi. 1903.
Apparently the first record of the tick in England is that of Neumann \
1897, p. 330, who refers to a specimen found on sheep (coll. Bur.
Animal Industry, Washington D.C.)] Wales: (Liverpool No. 11) from
grass on clifts, Gower, Glamorganshire, S. Wales, v. 1909, Dr J. W. W.
Stephens coll. France: We have seen (Brumpt No. 9) a (/* found
on deer at Fontainebleau, 13. ii. 1912, Baron Lestrange coll. [Neumann^
records specimens from a hidl, found at Alfort, near Paris, Railliet coll. ;
specimens from Digne and Beaune, E. Simon coll. ; from Villefranche in
Aveyron ; from sheep and cattle at St-Jean-de-Luz ; nymphs and larvae
were found in Aveyron on horse, hare, red and grey partridge (Neumann
coll.)] Holland: (Neumann^ records a $ from Numenius arquatus
(Curlew) ; a o from Plecotus auritus, found at Utrecht^ ; a ? from
Erinaceus europaeus; all in Oudemans' coll."-). Denmark: We have
determined (D.E.M. No. 22) a ? collected at Fano, vii. 1913 by
W. Horn. Germany: [L. Koch, 1877 h, p. 196, states that his type
of Rhipicephalus eocpositicias { = ^. H. c. var. punctata) was found near
Niirnberg] ; the tick was first recorded by Knuth in 1911-1913, who
found it in North Schleswig-Holstein. Through the courtesy of
Prof Knuth we have been able to examine his numerous specimens,
some of which he has allowed us to retain. The specimens were
collected in four districts as follows : {a) in Kreis Apenrade :
(Kn. Nos. 11, 12) cT from cattle and slteep, Schmedaggar, xi. 1911 ;
(Kn. No. 15) 15 j/" from cow dying of " Milzuptur," an obscure disease,
Schmedaggar, 1. xi. 1911; (Kn. No. 14 = N. 2710) 3 c/s from cattle
Kasso-Hof, XI. 1911 ; (Kn. No. 16) 1 c/" 3 $ from cattle, Reppel,
Gemeinde Schmedaggar, no date; (Kn. No. 7=N. 2708) 16 cT from
cattle, Krasso-Feld, ix. 1912. (6) In Kreis Apenrade and Tondern :
(Kn. No. 5) 2 c/ 40 $ from cattle, ix. 1912. (c) In Kreis Tondern:
(Kn. No. 10 = N. 2709) ^ ^ \ %, from cattle, Bau, ix. 1912;
(Kn. No. 13) 1 ^ horn cattle, Fanderup, ix. 1912 ; (Kn. No. 9) 1 cf 6 ?,
from cattle, Braderup, 22. ix. 1913. (d) In Kreis Husum : (Kn. No. 8)
1 $ with /. ricinns %, from cattle, 2\. IX. 1913. (e) In East Friesland :
' Neumann, 1897, p. 330. - Neumann, 1901, p. 2G0.

880 Genus HaemapJij/salls

(Kn. No. 1 = N. 2705) c/'s $ s from dairy cattle, Island of Norderney,
30. X. 1913; (Kn. No. 6 = N. 2707) ^s $ from cattle, Sportplatz,
Island of Norderney, 3. xi. 1913; (Kn. No. 3 = N. 2706) c/s ?s from
cattle, Island of Juist, 22. xi. 1913. Prof. Knuth dwells on the notable
absence of $s in some cases ; presumably the (/s remained upon the
hosts after the $ s had dropped off. Russia (South) : (N. 3008 and
Berlin Mus. No. 353) 2 os found on Turdus viscivorus L.,in Gouv. Poltawa,
20. IV. 1907, have been recently determined by us. [Yakimoff and
Kohl-Yakimoff, 1911, p. 418, record the species from Ananiewsk
District, Gouv. Cherson, and from the Caucasus. Their specimens
were doubtless determined by Neumann.] Italy: (N. 2544 h) gorged
$ s from sheep, Tuscany, XI. 1913, Prof. A. Berlese coll. [According to
Canestrini, 1890, p. 528, and Berlese, 1891, the tick is fairly common
on sheep, goat, and fallow-deer. " Herpetobia sidcata " Can. and Fan. also
favours these hosts. The tick attaches itself to sheep mainly behind
the eai*s. " Haemapfhysalis rhinolophi' Can. and Fan. was found on
Rhinolophus ferrum-equinum. The Italian authors state that the
immature stages of "H. sulcata" occur on Lacerta viridis, and
Neumann, 1901, p. 260 records os from Vipera aspis (C. Parona
coll., Genoa),] Spain: Rev. L. Navas of the College de Salvador,
Saragossa, sent us for determination a $ captured on the ground
in the environs of Saragossa, v. 1897. Hungary: (N. 1695) c/s ?s
from Equus cahallus, Budapest, vi. 1912, Prof, von Ratz coll. Croatia
and Dalmatia: [Neumann, 1897, p. 330, records a c/ from Fiume,
E. Simon coll. Canestrini and also Berlese state that nymphs and
larvae occur on Lacei'ta muralis var. pelagosae in Dalmatia.] Roumania
and Greece: [Neumann records specimens from Jassy, Leon coll.^ and
from Athens'-.] Islands in the Mediterranean: [Neumann records
specimens from Corsica^ E. Simon coll. ; nymphs and larvae found
on lizards on Cyprus, now in his collection ^ ; adults from the Cyclades
and Crete, without mention of hosts^].

AFRICA: Canary Islands: (N. 2080, 2081) d's ?s from goats,
Puerto Orotava, Teneriffe, ii. 1913, G. V. Perez coll. [Neumann states
that there are specimens in the Paris Museum from the islands and
(/s $s collected at Orotava and at Funchal, Madeira by Kraepelin
(Paris Museum 1).] Algeria: [Neumann records the species as having
been found on stones and on a lizard, Acanthodactylus vidgaris, at
Oran, Dumergue coll.; on cattle and goat at Blida and Medea; a

1 Neumaun, 1897, p. 330.

2 Neumanu, 1901, p. 260.

H. lei^oris-pahistris 387

$ from Mamia'; nymphs and larvae from Lacerta ocellata var.
tingitana, at Djebel Ksel, collected by Diimergue'-]. Egypt: We
have received specimens through the courtesy of l)r M. A. Ruffer
(N. 1926, 1928) ?sfrom sheep sind goat, Alexandria, xi. 1912 ; (N. 1929)
t/'s ?s from sheep, Mersyne, 1912. [Neumann states that the Smith-
sonian Institution coll., Washington, D.C., contains specimens taken
from sheep in Egypt \]

ASIA: Asia Minor: We have received two lots from Smyrna
(N. 2552a) $ from horses and cattle; (N. 2553a) cfs $s from goats;
both collected, xi. 1913 by Mr W. H. J, van Heemstra. Trans-
caucasia: Dr E, Dschunkowsky has sent us numerous specimens for
determination that were collected by him about Surnabad from sheep,
X. 1903 and lii. 1904; (N. 782) from hare, x. 1903 and ill. 1904;
(N. 783) from hare including typical forms and 4 J" without spined
coxa IV; (N. 784) 4 £/• 5 ? from fox, 1. x. 1903, J's without spined
coxae; (N. 787) 6 </ from bear, viii. 1903, cfs without spined coxae ;
(N. 780 a) (/"s with spine on coxa IV of normal size, medium size and
practically absent ; (N. 789) c/s ?s ivom fox, 1903, normal cfs and 5 c/s
without spines. We refer to these forms in the text, p. 384. Japan :
[Neumann^ records a $ from a horse, taken at Aoraori. Determination
correct ?] '

4. HAEMAPHYSALIS LEPORIS-PALUSTRIS
(Packard 1869).

Figs. 325-329.
Lit., Synon. and Icon. :

Ixodes leporis-palustris Packard, 1869 a, p. 67.

Not Ixodes chordeilis Packard, 1869 «, p. 67 ; as stated by Neumann, 1897, p. 34.3.

Gonixodes rostralis Dug^s, 1888, p. 129, Fig. 2. (The author describes under

this name an unfed $ which he regarded as a $, whilst the ^ and L

he describes and figures appear to be either Hyalomma or Amblyomma,

but the poor description and figures leave the matter in doubt.) It appears

clear that his G. rostralis ^ =-H. leporis-pahistris $ ).
Rhipistoma leporis Osborn, 1896, p. 261, with poor text-figures of $, capitulum

and tarsus of ^ ; wrongly states it is called the " Lone Star Tick " { = Am-

blyomina americanum !).
Haemaphysalis leporis (Packard) in Neumann, 1897, p. 343, Fig. 9 ; 1901 a, p. Ill,

Fig. 54 (condensed from previous description, the figure also reproduced.

We reproduce Neumann's figure). Lahille, 1905, p. 45 (quotes Neumann's

1 Neumann, 1897, p. 330.

2 Neumann, 1901, p. 260.

388 Genus Raemapliysalis

description and states the species (?) occurs in Argentina). Blanchard,
1909, p. 154 (sccircely more than hsted ; follows Neumann).

Haemaphysalis leporis Packard in Rohr, 1909, pp. 144-146 (merely quotes
Neumann's description).

Haemaphysalis hporis-palustris Packard in Hunter and Hooker, 1907, pp. 53, 54.
Text-Figs. 7 and 8, PI. Ill, Fig. 2 ; (species not described, brief mention of
distribution, biology, hosts ; Fig. 7 of 9 capitulum and scutum is in-
accurate ; Fig. 8 shows ^ and $ coxae ; the plate figure is a poor photo-
micrograph of a mounted $ ). Hooker, 1908 a, pp. 47, 48 (referred to as
" the rabbit tick of this country " (U.S.A.) and treats of its biology). Banks,
1908, pp. 33, 54, PI. IV, Figs. 8, 10, and PI. X, Figs. 2, 6 ((^ and ? described,
notes geographical distribution ; the o described in 14 words ; Fig. 8 of (^
capitulum ; Fig. 10 of $ capitulum, scutum, spiracle, coxa I and tarsus I ;
Fig. 2 of (iJ dorsum, $ capitulum in ventral aspect ; Fig. 6 of $ dorsum,
unfed and replete). Hooker, 1909, p. 423 (geographical distribution),
Hunter and Bishopp, 1911, pp. 228, 229 (brief notes on distribution, hosts
and feeding habits). Hooker, Bishopp, and Wood, 1912, pp. 89-96, PI. VII,
Figs. 1-6 ; (important in that the authors raised the tick experimentally ;
the figures are photomicrogi'aphs of unfed o and X, replete o, replete and
partly fed 5 , venter of ^ ; the photomicrographs are from balsam-mounted
specimens excepting those of the replete o and $ ; Fig. 5, a map, giving
distribution in the United States and Mexico ; they confer the [superfluous]
vulgar name of " rabbit tick " upon the species).

Haemaphysalis proxima Aragao, in Rohr, 1909, pp. 100-110, 146, 201, PI. II,
Figs. 12, 13, 16. (Rohr states that Ai'agao had thus named, but not as
yet described, the supposed new species. Rohr gives details regarding its
biology and distribution ; his figures, microphotographs, illustrate the
spiracle, etc. Not to be confused with Haemaphysalis proxima Warburton
and Nuttall, which = ^. cornigera Neumann, 1897 ; see further under follow-
ing paragraph.)

Haemaphysalis leporis var. proxima Aragao, 1911, p. 167, PI. XI, Figs. 4, 5 (we
have seen the type and do not consider the variety should be recognised ;
the figm-e is of the ^).

Haemaphysalis leporis-pabistris (Packard) in Hadwen, 1912, pp. 97, 98 (records
the tick in Canada ; raised the species experimentally and has kindly sent
us specimens).

Male (Fig, 325)-: Scutum about 2 x 1*3 mm.^ widest at the posterior

third, punctations coarse and confluent but not deep ; a pseudo-scutum
^ The scutums of 8 (? measured as follows in mm. :

N.

N.

1196 from Canada

1-95 X 1-34

384 from Texas

1-6 xl-2

1196 „

1-9 xl-30

1144 „ Canada

1-6 xl-17

719 „ Texas

1-9 xl-3

1144 „

1-56 X 1-12

384 „

1-75 X 1-25

1954 ,, California

1-14 X 0-8

Banks, 1908, p. 33, states that the <? is 1'6 mm. long. Hooker, Bishopp and Wood,
1912, p. 89, give the size of the <J (capitulum included?) as ranging from 2'25 x 1-25 down
to 1"6 X 1*0 mm.

H. leporis-palnstris

389

generally indicated ; cervical grooves rather long, convergent and deep
throughout their length ; lateral grooves commencing behind the pseudo-
scutum and therefore short, well-marked to the spiracle and faintly con-
tinued to include two or three festoons ; festoons rather broad. Capituhtm :
base broadest in front, the sides nearly straight and converging pos-
teriorly; cornua slight, but there are also ventral cornna ; palps longer

Fig. 325. H. leporis-palustris i . Dorsum, capitulum in ventral aspect, coxae with tro-
chanters, spiracle and tarsus IV. Drawn from (N. 719) a specimen from Texas.
Original, N. C. del.

than broad, sub-cylindrical, article 2 very salient beyond the base, but
the lateral contours of articles 2 and 3 form normally nearly a straight
line, only slightly recurved at the base of article 2^; no dorsal spurs,
but a slight point under article 3 ; hypostome 3 | 3. Venter : anal
grooves slightly ogival ; spiracle large, with slight dorsal process.
Legs : two short spurs, internal and external, on coxa I ; a slight spur
on coxae II-IV ; very slight trochantal spurs ; tarsus IV long, stout,
tapering rapidly.

1 We have, however, received specimens (N. 1144) from Canada which are undoubtedly
H. leporiti-iHiluitris, but in which article 2 of the palp is much more recurved at the base.

390 Genus Haemaphysalis

Female (Figs. 826, 327): Scutum about 0'9 x OvS mm.\ appearing

Fig. 326. H. leporis-palustns ? . Scutum, capitulum in dorsal and ventral aspects,
coxae with trochanters, spiracle and tarsus IV. Drawn from (N. 719) a specimen
found with the g in the foregoing Fig. Original, N. C. and E. W. del.

Fig. 327. H. leporis-palustris ? . (P) Palp of left side in ventral aspect, (Ti and 1*)
tarsi I and IV. x 80. Neumann, 1897, Fig. 9. (Eeprinted from original block.)

1 The scutums of 9 ? measured as follows in mm. :
N.

1-1 xl-0
0-9 xO-82
0-9 xO-8
0-9 xO-8

0-85 x 0'8 — the body of this specimen (unfed) measured
1-7 X 1-2 mm.

1196 from Canada
1144 .,

719

384

1144

Texas

Canada

1954

California 0-72 x 0-65
0-72 X 0-6

„ „ „ 0-7 xO-64

,, „ ,, 0-66 X 0-56

Hooker, Bishopp and Wood, 1912, p. 89, give the size of the ? (capitulum included ?)
as ranging from 2-5 x 1-5 down to 2-25 x 1-25 mm., when unfed, and attaining 6x3*5 to

H. leporis-palustris

noi

decidedly longer than broad, oval, narrowing posteriorly, with coarse
confluent punctations ; cervical grooves long and deei). Cajntulum :
base broader than in </ ; porose areas oval, converging anteriorly, far
apart; cornua slight, ventral cornua well-marked; palps with all the
characters of the ^ but relatively longer. Venter : spiracle sub-circular,
with slight dorsal process. Legs as in the cT- When replete, the ? may
attain 11-8 x 7-5 mui.

Nymph (Fig. 328) : Scutum relatively broader than in the $ .
Hypostonie 2 , 2. Other characters as in the $ ; the dorsal and ventral
cornua being even emphasized. When replete the O may attain
2-5 X 1-75^

Fig. 328. H. leporis-palustris O. Scutum, capitulum in dorsal and ventral aspects, coxae
with trochanters, spiracle and tarsus IV. Drawn from (N. 2324) Texan specimen.
Original, N. C. del.

Larva (Fig. 329): resembles the o, with still shorter palps
and broader, almost diamond-shaped scutum : the latter measuring

11-3 X 7-5 X 5-3 mm. when replete. Banks, 1908, p. 33, states that the scutum is 0-9 mm.
long.

1 The following measurements, in mm., were made from (N. 2324) nymphs received

from Mr Bishopp :

Scutums Body (unfed)

0-44 X 0-44 1-0 X 0-32

0-44 X 0-43

0-44 X 0-42

0-43 X 0-44

0-43 X 0-42
Hooker, Bishopp and Wood, 1912, p. 89, give the size of the o (including capitulum?)
as 1-33 X 0-8 when unfed, 2oxl-75 when replete; scutum 0-42 x 0-43 ; capitulum 0 22
long (between tips of cornua and palp).

392

Genus HaemaphysalU

0*26 X 0*3 mm/ Capitulum with ventral coiiiua. Legs: coxae feebly
armed. When replete the L may attain 1"33 x 0"9 mm.

Our descriptions of the ^f and $ refer more particularly to Texan
specimens (N. 719) named by Mr N. Banks, whilst the o and L are
described from specimens (N. 2322-2324) raised in Texas by Mr F. C.
Bishopp, who kindly presented us with them.

Fig. 329. H. leporis-palustris, larva. Scutum, capitulum in dorsal and ventral aspects,
coxae and tarsus III. Drawn from (N. 2321) Texan specimen. Original, N. C. del.

This is the only known species of which the ^ and $ have ventral
cornua or points projecting postero-laterally from the basis capituli
as well as dorsal cornua. Such ventral cornua occur, however, in the
immature stages of H. cinnaharina and H. cinnaharina var. punctata,
which are nearly allied forms.

See further under Notes on Biology, p. 530.

^ The following measurements, in mm., relate to larvae :

Bodies (replete)
N. 2322 0-26 x 0-31 0-55 x 0-45 N. 2323 1-2 x 0-38

1-15 X 0-37
1-09 X 0-37
1-05 X 0-35
1-05 X 0-35
Hooker, Bishopp and Wood, 1912, p. 89, give the size of the scutum at 0-24 x 0-3 mm.,
that of the unfed larvae (capitulum included?) at 0*53 x 0*38 mm., the replete larvae
attaining 1-33 x 0-9 mm. Capitulum 0-16 mm. long. The larvae we received from
Mr Bishopp were somewhat shrivelled by preservation in alcohol, consequently our
measurements of the replete Os are no doubt too low.

Scutums

Bodies (unfed)

0-26 X 0-31

0-55 X 0-45

0-26 X 0-3

0-53 X 0-45

0-26 X 0-3

0-25 X 0-3

//. leporis-palusfris 893

Geographical Distribution.

According to Aniorican authors, //. leporis-palastris is widely dis-
tributed in the United States, occurring in the following States :
Alabama**"; Arizona^'", uncommon'*; Arkansas'*; California^'", in Kern
County^ it is uncommon on rabbits^ though Marx states it is
common^'-"; Colorado «• '^^ « ; Florida'*-"; Idaho**; Illinois'*; Kansas'*", being
common^ and having been found on hare by Marx'^ ; Louisiana'**", at
Shreveport"^ ; Massachusetts"'"; Minnesota'*'"; Montana'*-'' where it
occurs in large numbers on rabbits in and about the State. Mr W. V.
King is stated to have killed 2 Lepus bairdi at Florence, 3. iv. 1911,
which were infested with 10o3 ticks, many of which were replete $s;
New Mexico'*, uncommon; New York**", in Keene Valley and Dunnc-
mora^; North Carolina'*"'^'", Packard^ describes the species from a $
found on Lepus palustris at Fort Macon; Nevada^-"; Oklahama**-";
Oregon*; Tennessee'*; Texas", where it was first collected by Marx^,
uncommon in the western portion of the Stated it occurs at Columbus,
Victoria and Maverick ^ stated to be common by Marx", and found on a
horse by Curtice^'^ found chiefly on the ears of hare and rabbit in 1906-
1907**; Virginia''"'*; Washington**; Wyoming*.

American authors^ record the following hosts of the tick in the
United States : Besides the type-host, Lepus palustris, the tick occurs
on 6 other species of hare and rabbit, these appearing to be the chief
hosts. Adults also occur on Felis domestica, robin, quail and nieadoiu-
lark. Immature stages are abundant on quail and meadow-lark, scarcer
on chaparral cock and Breiuers blackbird (assuming that they were
rightly determined, the immature stages are also found on thrush, field-
lark, jackdaw, blue jay, magpie and pine squirrel). Rabbits are nearly
always infested about the head, on the crest and occasionally about the
ears and eyes; when heavily infested^" the rabbits are often much
weakened and can be easily captured.

1 Packard, 18G9«, p. G7.

- Neumann, 1897, p. 343, who examined Marx's specimens in the Bureau of Animal
Industry collection and in the Paris Museum.

^ Marx, cited by (4) who give no reference ; his specimens examined by (2).

■* Hunter and Hooker, 1907, pp. 53-54.

5 Banks, 1908, p. 33.

'- Hooker, 1909, p. 423.

' Hunter and Bishopp, 1911, p. 2'28.

s Hooker, Bishopp and Wood, 1912, pp. 90-96.

» Hooker, Bishopp and Wood, 1912, p. 90.

'" Hunter and Bishopp, 1911, p. 228, give fewer hosts than the authors previously cited.

394 Grenus Haemaphysalis

Mexico : [Duges, 1888, j). 129, found his Gonixodes rostralis (see
our synonymy) at Guanajuato, Curtice is stated to have also found the
tick in Mexico \ Assuming that the determinations were correct, im-
mature stages have been found on birds in Tamaulipas and Monterey^.]

We have received or determined specimens from the following
countries :

UNITED STATES: from Texas: (N. 719) ^ $ from Victoria,
presented by the U.S. Dept. of Agriculture ; (N. 384) (/ $ found on
rabbit, Maverick County, v. 1906, presented by Mr W. D. Hunter ;
(N. 2321-2324) fed and unfed os and L raised at Kefugio and Utopia
in 1912-13 and presented by Mr F. C. Bishopp. California : (N. 1954)
small $s from Claremont (N. 2772) 12 $ from Lepus auduboni, San
Francisco, 30. iv. 1909, coll. M. B. Mitzmain.

CANADA: (N. 1144) (/" % from Xepiis a ?nmca7M(s, Aweme, Mani-
toba, 13. V. 1910, coll. N. Criddle and (N. 1196) adults, from Lepus dalli,
Peardonville, British Colombia, 17. vi. 1910, collected by Dr S. Hadwen,
who (1912, p. 98) also found the tick at Mt Lehman and at Nelson, B.C.

CENTRAL AND SOUTH AMERICA: Panama: (N. 1907) % s,
with their progeny of eggs and larvae, the $ s having been taken from
Dasyprocta sp., at Mataichan, Canal Zone, ix. 1911, by Dr S. T. Darling.
Brazil : (N. 1894) </ % from Lepus braziliensis, Manguinhos, vii. 1907,
presented by Dr H. de B. Aragao {H. leporis var. py^oxima Aragao, of which
we have also seen the type): [Neumann, 1901, p. 262, records a $ from
Brazil, collected by Delalande (Paris Mus.), and Rohr, 1909, pp. 100-110
states it occurs on wild rabbits and Dasyprocta agouti]. Paraguay:
(Berlin Mus. 113, 132) $ o collected at San Bernardino by K. Fiebrig.
Argentine: [Lahille, 1905, p. 45, states that the species attacks man,
and that S. Venturi has found it on Penelope obscura (a bird) in the
Province of Santa Fe, Colonia Mocovi, in September ; his determination
is uncertain, as he merely quotes Neumann's description].

1 Huuter and Hooker, 1907, p. 53 ; Hooker, 1909, p. 423.
- Hunter and Bishopp, 1911, p. 228.

H. montfjomcryi 395

5, HAEMAPHYSALIS MONTGOMERYI Nuttall, 1912.

Figs. 330, 331.
Lit. and Icon. : Nuttall, 1912, pp. 57-59, Fig.s. 7, 8 (reproduced).

Male (Fig. 330): Scutum Id x 13 to 25 x 1-6 mm.>, long-oval ;
cervical grooves normal ; lateral grooves including the first festoon

Fig. 330. Haemaphy salts moiitgomeriji 6 ■ Dorsum, right palp (P) in profile, spiracle,
anal grooves, coxae with trochanters, tarsus IV. (G. H. F. N. and F.M.H. del.)
Nuttall, 1912, Fig. 7.

1 Five (? gave the following measurements

in mm. :

Scutums

Capitulums

2-5 X 1-6

0-4

2-3 X 1-4

0-5

2-3 X 1-4

0-5

2-0 X 1-4

0-5

1-9 X 1-3

0-4

396

Genus Haemai)liy sails

and attaining half the body-length ; festoons longer than broad. Gapi-
tulura : 0'4 to 0'5 mm. long, base with lateral borders almost straight,
converging behind ; corniia pointed, continuous with the crescentic
dorsal ridge ; ventral ridge sharp, with trenchant lateral angles ; palps
with article 2 protruding slightly, about a third longer than article 3 ;
articles 2 and 3, viewed laterally (P in Fig. 330), bear sharp protruding
recurved spines ventrally ; hypostome 5 j 5 or 6 | 6, armed nearly to
the base with 12 distinct teeth per external file, besides finer denticles
and a large corona. Venter : hairy, genital orifice between coxae II ;
spiracle large with well-marked postero-dorsal elongation. Legs rela-
tively strong ; coxae I-IV with a long, pointed, retrograde spur, longest
on coxa I ; trochanters with pointed spurs ; tarsi short, tapering
from near the pseudoarticulation, bearing a small distal spur, and, in
some specimens, a slight median protuberance ventrally; pads almost
as long as the claws.

Female (Fig. 331): Scutum 0*9 x 11 mm.^, cordiform, with slight
lateral angles, broadly rounded behind ; cervical grooves well-marked,
extending slightly beyond half the length ; punctations poorly marked.

Fig. 331. Haemaphijtialis montgomeryi ? . Capitulum and scutum, capitulum in ventral
aspect, coxae with trochanters, spiracle and tarsus IV. (F. M. H. del.) Nuttall,
1912, Fig, 8.

1 All of the ? were but partly gorged ; they measured roundly 4 mm. in length.

H. klnneari 397

uniformly distributed. Capituluin 0 5 iiini. long, resembling that of (/*,
but broader, shorter, with cornua less pron(nmced ; porose areas small, oval,
far apart, converging in front, placed anteiiorly. Venter : vulva between
coxae II; spiracle with dorsal and posterior margins flattened. Legs
resembling those of tlie (/, but the spurs on coxae and trochanters less
pronounced; tarsi tapering gradually, unarmed.

Described from 9 </ and 2 ? taken from the ears of ponies, at
Muktesar, United Provinces, India, 30. v. and 1-7. vi. 1905, also on
a bull's ear at Bhulumaya ; 3 ^Z* found on a dog, Muktesar, 8. Vili.
1905 ; 2 $ found on the ear of a bull, at Berinag, U.P., 9. ix. 1905 ; all
of these specimens (N. 760, 761, 762) were collected by Dr R. E.
Montgomery, after whom the species is named. We have since
received: (N. 1407) cT from a dog, Balaghat, Central Provinces, India,
1908, coll. S. H. Gaiger; (N. 2251) $ from sheej), Gilgit, Kashmir, v.
1913, coll. Dr M. Abdullah ; (N. 2923) cf from dog, Kashmir, 1912, coll.
J. E. M. Mellor ; and specimens have been sent to us for determination
by the Indian Museum : cT and $ , collected at AlmoraS Kumaon
(5500 ft. elevation), vi. 1911, by C. Paiva.

Types in Cambridge ; we have presented co-types to the Berlin
Museum and to the Neumann collection, Toulouse.

6. HAEMAPHYSALIS KINNEARI Warburton, 1913.

Fig. 332.
Lit. and Icon. : Warburton, vii. 1913, pp. 127, 128, Fig. 6 (reproduced).

Male : Unknown.

Female (Fig. 332) : Scutum sub-circular, slightly broader than long,
0-8 X 09 mm., with numerous medium or rather large punctations,
most numerous on the lateral fields ; cervical grooves, beginning as
shallow depressions, deepening into oval pits at some distance from the
anterior border, then broad and shallow to within a short distance of the
postero-lateral borders. Capituluni base twice as broad as long, rect-
angular, with blunt cornua ; porose areas apparently nearly circular, but
ill-defined and not easily distinguished ; palps long and, in general
facies, much like those of H. montgonievyi, having the lateral salience
slight and very obtuse ; instead, however, of being smoothly rounded

' Almora is distant about 14 miles from Muktesar where the types were found.
N. I. 26

398

Genus HaemapJiTj salts

as in that species, their dorsal surface presents corrugations, article
3 has an irregular posterior border, and the internal contour of article 2
presents a strong rounded protuberance anteriorly ; moreover, article 2
bears no ventral retrograde spur, though one is present under article 3 ;
h3-postome 4 | 4. Venter : spiracle rather large, oval, with blunt dorsal
process ; anal grooves ogival. Legs : coxae with very slight armature of
the normal type ; tarsus IV rather long (especially the proximal pseudo-
segment), somewhat tapering ; no trochantal spurs.

Fig. 332. H. kinneari ? . Capitulum and scutum, ventral aspect of capitulum, anus
with anal grooves, tarsus IV, spiracle and coxae. (N. C. del.) Warburton, 1913^
Fig. 6.

Described from (N. 1997)1 $ taken from a tiger at Kadra, Kanara,
India (Bombay Nat. Hist. Mammal Survey), xi. 1911, by Mr N. B.
Kinnear, after whom the species is named. The specimen was found
in company with H. hispinosa and Anihlyomma prolongatum.

Type in Cambridge.

A- large species, recalling H. montgomeryi in general appearance but
easily distinguishable from that species by the rounded scutum, the
absence of a ventral spur on the second palpal article and the absence
of trochantal spurs.

7. HAEMAPHYSALIS ABORENSIS Warburton, 1913.

Fig. 333.

Lit. and Icon. : Warburton, vir. 1913, pp. 122, 123, Fig. 1 (reproduced).

Male : Unknown.

Female (Fig. 333) : Scutum broader than long, 1*5 x 1'8 mm., broad-
oval, pale yellow ; punctations numerous and fairly large, distributed

H. ahorensis

899

laterally and posteriorly, inconspicuous in the median field ; cervical
grooves far apart, extending to tht' posterior border. Gapituhim: base
much broader than long, cornua shoi-t and blunt ; porose areas ill-defined,
ovate, the interval greater than their diameter ; palps with very slight
and obtuse lateral salience on article 2, of which the internal border is
nearly straight and much longer than the external border; the dorsal
surface of articles 2 and 3 smooth, rounded and destitute of spines ;
a short, blunt spur under article 3 ; hypostorae spatulate, well-covered

Fig. 333. H. aborensiti ? . Capitulum and scutum, anus with anal grooves, spiracle,
tarsus IV and coxae. (N. C. del.) Warburton, 1913, Fig. 1.

on its anterior f by 4 j 4 sub-equal teeth. Venter : spiracle large,
trans\ersely oval, with slight blunt dorsal projection ; anal grooves nearly
semi-circular, with hardly any visible median post-anal groove. Legs :
very long and strong ; coxa I with a short, blunt internal spur ;
coxae II-IV with slight spur in the middle of the posterior border;
tarsus IV long and tapering, pad short.

Described from 1 $ found on grass at Yambung, India, 13-17. I.
1912, by S. W. Kemp (Abor Expedition); the species is remarkable
because of its size.

Type in the Indian Museum, Calcutta (No. 1251/17).

26—2

400

Genus Haemaphysalis

8. HAEMAPHYSALIS FORMOSENSIS Neumann, 1913.

Figs. 334, 335.

Lit. and Icon. : Neumann, 1913, pp. 135-137, Figs. 1, 2 ( (^ capitulum in ventral
aspect with coxa I, $ capitulum in dorsal aspect with scutum and trochanter
I, not reproduced).

Male (Fig. 334): Body broadly oval, livid brownish-yellow, legs
lighter. Scutum about 2*4 x 1-9 min.^ ; punctations numerous, but

Fig. 334. H. formosensis i . Dorsum, capitulum in ventral aspect, coxae with
trochanters, spiracle and tarsus IV. (N. 2730 Co-type.) Original, N. C. del.

very shallow and inconspicuous ; cervical grooves slight oval pits ;
lateral grooves long and rather noticeably straight, including one
festoon ; festoons long. Capitulum : base rectangular (in some speci-
mens rather broader posteriorly), cornua strong and blunt; palps

^ The scutums of the 5 cT co-types in our collectiou (N. 2715, 2730) measured
respectively :

2o X 1'9 mm.
2-4 xl-85 „
2-35 X 1-85 „
2-3 xl-9 ,,
2-1 xl-75 „

H. formosensis

401

short, very slightly salient laterally, with rounded contours and with-
out dorsal spines ; article 2 rather longer than article 3 ; a small but
distinct retrograde spur under article 3 ; article 2 ending in a point
which, however, does not project; hypostome short, broad in front,
corona rather small, dentition 6 | 6, the median teeth very small, the
three external files much larger, about 8 teeth per file. Venter :
spiracle pear-shaped, the pointed end dorsal. Legs : coxae I-IV with
rather strongly-developed spurs ; coxa I has its posterior end trun-
cated, and the spur proceeds from its postero-external angle ; small
trochantal spurs, progressively diminishing, and almost absent on tro-
chanter IV^ ; tarsus IV stout, with pseudo-segments of almost equal
length, tapering moderately, with a small terminal ventral spur ; pad
moderate.

Female (Fig. 335): Scutum about 1"1 x 1'3 mm., sub-circular, but
rather broader than long, with numerous fine shallow punctations ;

Fig. 335. H. formosensis ? . Capitulum in dorsal and ventral aspects, scutum, coxae
with trochanters, spiracle and tarsus IV. (Deutsches Entomol. Mus., Type.)
Original, N. C. del.

cervical grooves beginning as small pits behind the anterior border and
visible as shallow grooves almost to the posterior border. Capitulum :
base rectangular, twice as broad as long, with slight blunt cornua ;
porose areas large, far apart, oval, converging anteriorly; palps with
hardly any lateral salience, longer than in the (/, but with the same
characteristics ; hypostome 4 1 4, with a trace of 5 | 5 at its posterior
end. Venter : spiracle transversely oval, with faint dorsal protuberance.
Legs : coxae as in the cT ; trochantal spurs absent ; tarsus IV more
slender than in the </, unarmed.

' The slight spur on trochanter I, is omitted in the figure.

402 Genus Haemaphysalis

Neumann founded this species on 20 </ and 2 $ found on the dog
at Kosempo, Formosa, iv. 1912. We here redescribe the species from
5 </ and 1 % belonging to the type material : 1 ? (type) lent and 4 cT
(co-types) presented by the Deutsches Entomologisches Museum, and
1 </ (co-type) presented by Professor L. G. Neumann. Our description
and figures differ somewhat from the author's. We have found (N. 2958)
a % from Burma (in a tube together with a $ i/". hystricis probably
from Hystrix hengalensis or Ursus torquatus) in the L. Fea collection at
the Museo Civico di Storia Naturale, Genoa.

Types (D.E.M. No. 14 c/* ?) in the Deutsches Entomologisches
Museum, Berlin-Dahlem ; co-types ((/ $ ) in the Toulouse and (N. 2715,
2730 ; 5 (/"s) Cambridge collections.

9. HAEMAPHYSALIS JAPONICA Warburton, 1908.

Figs. 336, 337.

Lit., Icon, and Synon. :

Haemaphysalis flava Neumann, 1897, p. Z'i^, pro parte ; Neumann, 1905, p. 237,
pro parte (see discussion under Zr._^ai-a Neumann, p. 408).

Haemaphysalis japonnica Warburton, 1908, pp. 512, 513, Figs. 3, 4 (reproduced).
Blanchard, 1909, p. 152, wrongly gives H. japonica as a synonym of
H. flava Nn.

Haemaphysalis flava flava Neumann, 1911 a, p. 12.

Male (Figs. 336, 337) : Scutum 2*5 x 1-8 mm., oval, rather broad, not
much narrowed in front, glossy yellow, with numerous rather shallow
punctations, absent in places ; cervical grooves, oval pits ; lateral grooves
fairly long, markedly concave internally, including distinctly one and
faintly two festoons ; festoons rather short, almost square, punctate.
Venter glossy, punctate, genital orifice large, between coxae II ; anal
grooves ogival ; spiracle short comma-shaped. Capitulum base broad,
punctate, with short blunt cornua ; palps with article 2 moderately
salient, the lateral contours of articles 2 and 3 unbroken ; no dorsal
spines ; ventrally article 2 is much corrugated and forms a retrograde
angle ; a short spine under article 3 ; hypostome spatulate with very
large corona, dentition 5 j 5, small pointed sub-equal teeth. Legs :
strong ; coxae large, especially coxa IV, all with a short spur, sharp
on coxa I, conical on coxae II-IV ; tarsus IV fairly long, considerably
more slender than the preceding article.

Female : unknown.

H. japonica

403

Our description is based on numerous specimens found in a tube
labelled "No. 173, H. flava" in the British Museum, the specimens
having been wrongly determined by Neumann. The ticks were found
on Neniorhaedus crispus at Hondo, Japan, by the Duke of Bedford's

•Fig. 336. Fig. 337.

Fig. 336. H. japonica <? , dorsum. Warburton, 1908, Fig. 3.

Fig. 337. H. japonica d , venter and spiracle. Warburton, 1908, Fig. 4.

(Sketches, C. W. del.)

collector. We have since received (N. 1374) 24 </ taken from a roe-
buck at Kansu, China, the specimens were purchased by us from
Mr T. V. Sherrin, taxidermist.

Types in the British Museum and co-types (N. 1247, 2 j/*) in
Cambridge.

Haemaphysalis japonica var. douglasi Nuttall
and Warburton, 1915, n. var.

This variety differs from the type as follows :

Male : Distinctly smaller. Scutum 2*1 x r5 mm., narrower in front,
and with straighter sides and lateral grooves. Palps with ventral spine
on article 3 distinctly longer and sharper.

Female : unknown.

Described from 4 (^ taken from roe-deer, at Ten-an-fu, Shiensi,
Northern China, 19. v. 1909, by Captain H. E. M. Douglas.

Types in the British Museum and co-types (N. 1248 2 j/") in
Cambridge.

404

Genus Haemaphy sails

10. HAEMAPHYSALIS PAPUANA Thorell, 1882.

Figs. 338-341.

Lit. and Icon. : Thorell, 1882, pp. 62-66, PL VI, Figs. 40-45 ( S dorsum and
venter, ends of digits, foot ; $ dorsum ; good figures). Canestrini, G., 1884,
p. 705 (repr. p. 13) PI. VI, Fig. 4 (figures the capitnlum of a specimen he
determined as H. papuana from Queensland ; his determination is probably
wrong, if not the figure is inaccurate). Neumann, 1897, pp. 336, 337. Rainbow,
1906, p. 165 (lists the tick as occurring in Australia on Canestrini's authority
cited above q.v.). Warburton, 1908, pp. 514-516, Fig. 6 (reproduced). Blan-
chard, 1909, p. 154 (species listed only). Neumann, 1911 a, p. 108.

Male (Fig. 338) : Scutum about 22 x 1*6 mm.S yellow, flat, broadest
towards the posterior end, coarsely pitted except along certain ridge-

Fig. 338. H. j)apuana <? , dorsum and venter ; ? capitulum and scutum ; 3 and ?
spiracles. Specimen from Borneo. Warburton, 1908, Fig. 4. E. Wilson del.

like tracts where punctations are absent ; festoons about as broad
as long, the intervals often dark and broadening distally; cervical

The following measurements, in mm., relate to 20 <? from various sources :

N.

Scutums

N.

Scutums

309 from

Borneo

2-5 xl-7

295 from Borneo

2-2 xl-6

1563 ,,

,,

2-45 X 1-8

1563 ,

,,

2-15 X 1-55

496 „

Java

2-4 xl-65

1563 ,

) ))

2-15x1-52

1563 ,,

Borneo

2-4 xl-76

220 „

,,

2-14 X 1-5

496 ,,

Java

2-34 X 1-6

2116 ,

Malaya

2-1 xl-6

220 „

Borneo

2-25 X 1-6

1563 ,

Borneo

2-1 xl-6

1563 „

,,

2-3 xl-7

1563 ,

,,

2-1 xl-57

1563 „

,,

2-27 X 1-6

309 ,

, ,

2-05 x 1-65

220 ,,

,,

2-24 X 1-7

1563 ,

,,

2-04x1-54

309 ,,

,,

2-2 xl-65

1563 ,

,,

1-97 x 1-43

The size of the <? given by Thorell (about 3x2 mm.) is apparently only roughly
approximate and doubtless includes the capitulum.

H. papnana

405

grooves consist of oval pits with slight shallow continuation ; lateral
grooves very short and ill-defined, beginning about midway along
the body-length and ending behind the spiracles. Capitulum : base
rectangular, with slightly convex sides, broad blunt cornua, coarsely
punctate dorsally ; palps only slightly salient, the external angle of
article 2 fairly sharp, its inner dorsal border has an inward projection
distally ; no distinct dorsal spines, but the posterior border of article 3
is sharply angular in the middle ; article 2 is sharply angular ventrally ;
a short, retrograde ventral spine on article 3 ; hypostome 4 \ 4.
Venter : yellow-brown, darker than the scutum, glabrous ; finely
punctate ; sexual orifice broad, between coxae II ; sexual grooves far
apart, nearly parallel ; festoons well-marked ventrally, with dark
intervals ; spiracles yellow, ovoid, with fairly pronounced dorsal process.
Legs : a short sharp spur on coxa I, a very slight spur near the middle
of the posterior border of coxae II and III, a very short spur at the
inner angle of coxa IV ; trochanters slightly spurred ; tarsi relatively
short, somewhat humped ; pad medium.

Atypical cT from Malaya (Fig. 339). We include under H. papuana

3:1:5^

Fig. 339. H. papuana i (N. 2116, atypical form from Malaya). Dorsum, ventral aspect
of capitulum with coxae and trochanters, spiracle and tarsus IV. Original, N. C. del.

a specimen (N. 2116) from Malaya which only differs from the typical
form in that the characteristic coarse punctations are absent ; the scutum
is smooth, irregularly undulating, with very faint shallow punctations.

406

Genus HaemaphysaUs

Female (Fig. 338): Scutum nearly circular, about 1x1 mm.^
bright yellow, coarsely pitted; cervical grooves shallow, concave
externally. Capitulum : base broader and shorter than in the cT, with
very slight cornua ; sides somewhat convex ; porose areas large, oval,
far apart, converging anteriorly; palps longer and narrower than in
the (/ ; article 2 with sharper external angle, its inner dorsal contour
like that of the </. Venter: spiracle ovoid, with blunt dorsal pro-
tuberance, yellowish. Legs : coxae as in the ,/ ', tarsus IV tapering
gradually, the distal pseudo-segment rather long.

Nymph (Fig. 340) : Scutum deeply emarginate, oval, about as long
as broad, but appearing longer (045 x 0"45 mm.), of uneven surface

Fig. 340. H. papuana. Nymph. Capitulum in dorsal and ventral aspects, scutum, coxae
with trochanters, spiracle and tarsus IV. Specimen (N. 220) from Borneo. Original,
N. C. del.

and more punctate than is usual with nymphs of this genus. Cervical
grooves well-marked, nearly parallel for half the scutal length. Capitulum :
base rectangular with slight cornua ; palps slightly salient laterally,
fairly long ; article 2 about equal in length to article 3 ; no dorsal
spines, but the inner angle of article 3 slightly recurved ; a slight
retrograde spur under article 3 ; hypostome rather narrow, 2 j 2.
Venter: spiracle circular. Legs: coxae normal; tarsus IV short and
tapering rapidly ; pad fairly long.

1 The following measurements, in mm., relate to 8 ? from various places in Borneo :

N.

Scutums

■ N.

Scutums

309

115xl-2

1563

0-95 X 1-07

220

1-05 X 1-15

1563

0-95 X 1-05

295

1-0 xl-0

220

0-9 xl-05

220

0-97 X 1-0

1563

0-9 xl-03

H. papimna

407

Larva (Fig. 341): Scutum cordate, broader than long (about
2-3 X 3-3 mm.). Capitiduiii : like that of the O, with obscure division
between palpal articles 2 and 3. Lefjs : as in the O, but the spurs on
coxae II and III practically absent. When replete, the larva attains
1*35 X 11 mm.

Thorell's excellent figures of \\\e ^ and % leave no doubt as to the
identity of this species, and show clearly various characteristics not
mentioned in his description, such as the ridge-like tracts, referred
to in our description, the short indefinite lateral grooves, and the
inward dorsal projection of article 2 of the palps.

Our description is based on specimens collected in Borneo by
Dr A. R. Wellington : (N. 220, 295) 2 c/ 2 $ 3 o from dog, Sarawak,

Fig. 341. H. pajmana. Larva. Capitulum in dorsal and ventral aspects, scutum, coxae,
tarsus III. Specimen (N. 1563) from Borneo. Original, N. C. del.

I. 1907, and (N. 309) 3 j" 1 ? from pig, Sarawak, vir. 1907. We have
also received (N. 1563) 30 cT 1 ? 1 o and 1 L from Ursus malayanus,
collected ii. 1912 at Kuching, Sarawak, by J. C. Moulton. We possess
specimens from Java: (N. 496 h J") fi'om Felis pardus, collected by
Dr J. C. Koningsberger in 1908 and (N. 3117 a (/) from wild pig,
Tjibaroesa near Buitenzorg, 1914, Dr de Blieck coll. ; we have deter-
mined adults found in Sumatra on a tiger (Schliffner 3 a) 1912.
The atypical form (N. 2116 cT) we have figured, was found on a pariah
dog, near Kota Baru, Kelantan, Federated Malay States, 17. iii. 1913,
by Dr J. D. Gimlette.

Thorell's types (1 </ 1 ? ) were found on a mammal at Ramoi, New
Guinea; they were collected by L. M. D. Albertis. We do not know
where these types are deposited. Our types of the o and L are in
Cambridge.

408 Genus Haemaphysalis

11. HAEMAPHYSALIS FLAVA Neumann, 1897.

Figs 342, 343.

Lit., Ssmon. and Icon. :

Haemaphysalis flava Neumann, 1897, pp. 333-336, Fig. 3 (digit ; not reproduced) ;

Neumann, 1901, p. 260 {pro parte).
Haemaphysalis flava var. armata Neumann, 1905, pp. 237, 238.
Haemaphysalis flava Neumann in Donitz, 1905, pp. 129, 130.
Haemaphysalis flava Neumann 1897, in Warburton, 1908, pp. 510-512, Figs. 1

and 2 {$ and ? de.scribed ; figures reproduced); Blanchard, 1909, p. 151

(only a few lines).
Haemaphysalis flava armata Nn. in Neumann 1911a, p. 112 (raised to a

sub-species of his H. flava of which it constitutes the type !).

Nothing can exceed the confusion which has arisen with regard to
this species. Neumann's original description (1897) was fairly definite^
but he subsequently (1905) came to the conclusion that two varieties
were in existence, and these he interchanged, making the last found
form the type, and the original type the variety. Donitz (1905)
pointed out that this was inadmissible, and we entirely agree with
him. Moreover, Neumann, in identifying specimens, confounded yet
other species with H. flava, for in a tube (No. 137 in the Brit. Mus.)
labelled by him H. flava, we found three distinct species". Donitz
(1905), moreover, found two species^ among alleged H. flava which
Neumann sent him. After reviewing the situation, Warburton (1908)
reinstated what appeared to be the original H. flava and removed two
other forms from it under the names of H. japonica and H. campanulata.

Male (Fig. 342): Colour dull yellow. Scutum 2*1 x 1-4 to
2'8 X 1'9 mm.*, broadest at about the level of the spiracles; punctations
numerous, of medium size, regularly distributed ; cervical grooves short,
faint, converging, then separating ; lateral grooves short, beginning about
at half the body length and ending just behind the spiracles ; festoons
long, curved. Capitidum : base rectangular, twice as broad as long,
with strong sharp cornua;- palps with article 2 fairly salient, but not
sharply angular externally; lateral contour of articles 2 and 3
continuous; no dorsal spines; ventral border of article 2 having a

^ Neumann here specifically mentious the yellow colour and the long-spurred coxa IV
of the (J . Donitz (190-5, p. 130) lays stress on the characteristic colour. We would note
that the name " armata." chosen by Neumann, refers to the long sharp spur on coxa IV.

2 Including H. japonica Warburton 1908, H. campanulata Warburton 1908.

3 H. neumanni Donitz 1905 ( = H. bispinosa Nn. 1897) and H. flava Nn.
* The second measurement is from (N. 763) specimens from India.

H. flava

409

somewhat prominent retrograde angle; a distinct retrograde spine,
directed inwards at the ventro-lateral angle of article 3 ; hypostome 4 | 4
on its distal half, then 5 | 5, the additional teeth being median. Venter :
yellow, spiracles white, ovoid, with slight blunt dorsal protrusion.
Legs : a fairly long blunt spur on coxa I ; shorter blunt spurs on co.xae II
and III ; a long sharp spur (longer than the width of the article) at
the inner angle of coxa IV; tarsus IV rather short, about thrice as
long as broad, tapering gradually.

Fig. 342. Fig. 343.

Fig. ?4'2. H. flava i . Dorsum, spiracle and coxae. (Sketch by C. W.) Warburton,
1908, Fig. 1.

Fig. 343. H. flava ?. Capitulum and scutum, coxae, spiracle and tarsus IV. (Sketch
by C. W.) Warburton, 1908, Fig. 2.

Female (Fig. 343): All yellow when unfed, the scutum a bright
yellow when the body is dark and distended. Scutum as broad as long,
about 1x1 mm.^, broadest at the posterior third, bright yellow, with
numerous fairly large punctations ; cervical grooves long, concave
externally, beginning somewhat behind the anterior border as fairly
deep depressions and continuing faintly to the postero-lateral margin ;
no lateral grooves. Capituluin : base comparatively broader than in
the cT, and with short blunt cornua, porose areas large, oval, converging
anteriorly, their interval equal to their long diameter, a few irregular
punctations in the middle of the interval ; palps more salient and
more sharply angular outwards, otherwise like those of the (/ ;
hypostome 4 [ 4 or 5 I 5. Venter : spiracles white, rounded, with blunt

1 The scutums of two other females measured in mm. : (N. 893) from Japan, 0-85 x 1 ;
(N. 763) from India, 1 •2x1-2.

410 Genus Hae^naphy salts

dorsal protuberance. Legs : coxa I with weaker spur than in the cT ,
a slight sharp spur at the middle of the posterior border of coxae II and
III, a short blunt spur at the internal angle of coxa IV ; tarsus IV
slightly longer than in the (/, tapering gradually. When replete the
$ may attain a length of 8"5 mm.

Our description is based on (N. 893) 1 (/ 1 $ from Japan, kindly
presented by the late Prof W. Donitz. The specimens were found on a
dog at Ise, and on cattle or on a hoise at Hiroshima. Neumann (1905
p. 238; 1911 a, p. 112) states that his armata (i.e. his original Jiava
were found in Japan upon a hmse and on vegetation. We have received
and determined specimens from India: (N. 763) </ $ from dog, Madras,
collected v. 1906 by Mr V. I. Phadke ; (Montgomery coll.) adults from
Sus cristatus, Muktesar, viii. 1905 ; (Indian Mus. coll.) cT from sheep,
on hillside below Phagu (7000 ft.), Simla Hills, Western Himalayas.

T3rpes in Toulouse.

12. HAEMAPHYSALIS TURTURIS Nuttall and
Warburton, 1915, n. sp.

Fig. 344.

Male (Fig. 344): Scutum 2 x 1'5 mm., ovate, broader posteriorly,
covered by very numerous small shallow punctations also on the
festoons ; cervical grooves shallow, comma-shaped, beginning at some
distance behind the anterior border; lateral grooves almost absent,
represented only by short shallow depressions dorsal to the spiracle.
Capitulum : base rectangular, not greatly broader than long, cornua
strong; palps short, only slightly salient laterally, with broken lateral
contour ; articles 2 and 3 of about equal length ; a very slight
retrograde point at the middle of the posterior dorsal border of article
3 ; the posterior ventral border of article 2 ends in a sharp point, and
there is a strong retrograde spine under article 3 ; hypostome 4 j 4,
small equal teeth, about 7 per file, corona large. Venter : spiracle
ovoid, with distinct dorsal process. Legs fairly long and strong ; coxa I
with spur well-developed, a slight spur, progressively diminishing,
on coxae II-IV ; trochanters with distinct retrograde ventral spurs,
largest on trochanter I ; tarsus IV fairly long and strong, tapering
rather rapidly.

Female : unknown.

H. turturis and H. acicnlifer

411

Described from a single (/ found on a dove (Turtur suratensisy,
Ceylon, 2. Iii. 1906. H. Schoede coll.

The specimen is of a uniform pale yellow colour. In size and
general facies it greatly resembles H. Jiystricis Supino, but appears to
be separated from this species by the presence of trochantal spurs,

Fig. 344. H. turturis cf . Dorsum, capitulum in ventral aspect, coxae with trochanters,
spiracle and tarsus IV. Drawn from the type. Original, N. C. del.

which are perfectly distinct, though not very strongly developed. The
dorsal spine on the third article of the palp of H. hi/stricis is represented
in the present species by a very small retrograde cusp.
Type (No. 167) in the Zoological Museum, Berlin.

13. HAEMAPHYSALISACICULIFERWarburton, 1913.

Figs. 345, 346.
Lit. and Icon. : Warburton, vii. 1913, pp. 125, 126, Figs. 4, 5 (reproduced).

Male (Fig. 345) : Scutum about Vd x 1*2 mm., oval, widest in the
middle, glossy, with very fine, shallow, unequal punctations ; cervical
grooves short, fairly distinct ; lateral grooves distinct, rather short,
concave internally, including one festoon ; festoons fairly long and rather
broad. Capitulum : base rectangular, with strong cornua ; palps
short ; article 2 short, but slightly salient laterally, prominent, but
without ventral spine ; article 3 slightly salient laterally, breaking the

1 The original label refers to the host as " Kowadall's Taube " which no doubt stands
for " Kawada," the Mahrati name for this species.

412

Genus Haemajjhy salts

external contour, without dorsal spine but slightly protuberant where
a spine might be expected, recurved at its postero-internal margin ;
the whole posterior border of the palp is characteristic ; hypostome
short, dentition 4 | 4, teeth small. Venter : spiracle rather broad, with
very slight, blunt, dorsal process ; anal grooves slightly ogival. Legs :

Fig. 345. H. aciculifer d . Capitulum in dorsal and ventral aspects, scutum, coxae and
trochanters, spiracle, tarsus IV and anus with anal grooves. (N. C. del.) Warburton,
1913, Fig. 4.

coxa I with strong internal spur and a slight hint of an external spur ;
coxae II and III with slight protuberance in the middle of the posterior
border; coxa IV with long, needle-like, internal spine (hence
aciculifer) longer than the article itself, and narrow from its origin ;
slight, progressively diminishing trochantal spurs ; tarsus IV tapering
rather abruptly ; pads rather long.

Female (Fig. 346) : Scutum cordiform, as long or slightly longer
than broad (0'9 x 0"9 or 0'8 x 0'7 mm.), widest at the anterior third ;
punctations small and inconspicuous ; cervical gi'ooves well marked,
slightly converging, visible for about two-thirds the scutal length ;
emargination deep. Capitulum : base much broader than long, cornua
strong; porose areas oval and far apart; palps as in the </, especially as
regards their internal contour, but the dorsal protuberance on article 3

H. kochi

413

is more marked and pointed, almost amounting to a spine ; hypostorae
4 I 4. Venter: spiracle sub-circular ; anal grooves slightly ogival. Legs:
coxa I Avith fairly sharp internal spur ; coxae II-IV with slight pro-
tuberance near the middle of the posterior border, strongest on
coxa IV: tarsi and trochanters as in the </•

Fig. 346. H. aciculifer ? . Capitulum in dorsal and ventral aspects, scutum, anus
■with anal grooves, spiracle, coxae and trochanters. (N. C. del.) Warburton, 1913,
Fig. 5.

Described from (E. 463 a) 1 ^ and 1 ? found on Cohus thomasi
(antelope) on the N.E. shore of Lake Edward, Uganda, x. 1911, by
S. A. Neave. We have since received (N. 2625) 1 $ from a reedbuck,
Wandara, Gold Coast, West Africa, 16. v. 1913, coll. J. J. Simpson.

Types in London (Imperial Bureau of Entomology).

14. HAEMAPHYSALIS KOCHI Aragao, 1908.

Fig. 347.

Lit. and Icon. : Aragao, 1908, reprint, pp. 3-6 ; Kohr, 1909, pp. 142-144, 200
(merely quotes Aragao) ; Blanchard, 1909, p. 152 (merely listed) ; Aragao,
1911, pp. 178-181, PI. XII, Figs. 16, 17, 17a (S venter, capitulum in dorsal
and ventral aspects)

Male (Fig. 347): Scutum 1-77 x 1*12 mm., regularly oval, broadest
at the posterior third, earthy yellow; cervical grooves short comma-
shaped, the concavity outward, deep anteriorly ; lateral grooves very
short, beginning with a shallow depression opposite leg IV and including
one festoon ; festoons distinct, longer than broad ; punctations fine
and discrete on scutum and festoons. Capitulum short (0*46 mm.) ;
base broader than long (012 x 0'28 mm.) with very strong cornua ; palps

N. I. 27

414

Genus Haemaphysalis

without dorsal spines, articles 2 and 3 nearly equal in length, lateral
salience of article 2 slight, a strong hook-like spine under article 3 ;
hypostome spatulate, dentition 4 | 4, 7 to 8 teeth per file. Venter pale
yellow, finely punctate, bearing a few short hairs ; anal grooves V-shaped ;
spiracle ovate, narrowing dorsally. Legs long, hairy, yellow ; coxa I with
rather strong internal spine, a blunt tuberosity, decreasing progressively
in size, near the middle of the posterior border of coxae II-IV ; the
trochanters bear slight, progressively decreasing spurs ; tarsi tapering ;
pads as long as the claws.

Fig. 347. H. kochi s . Dorsum, coxae and ventral aspect of capitulum ; lateral aspect
of palp, spiracle. ? capitulum and scutum. Sketched from the types. Original,
C. W. del.

Female (Fig. 347) : Body, when unfed, regularly oval, 279 x 1*66 mm.
Scutum nearly circular, 1"02 x 1'16 mm. ; cervical grooves attaining
the posterior border, but deepest at the anterior half; punctations fine
and evenly distributed. Capitulum : base rectangular, 0"1 x 0"56 mm.,
cornua much less marked than in the (/ ; porose areas small, oval, far
apart, converging anteriorly, their posterior ends encroaching on the
lateral ridge ; palps with article 2 much longer than article 3, and
more salient laterally than in the cf ; hypostome spatulate, dentition
5 I 5, 9 to 10 small teeth per file. Legs as in the ^ , but the coxal spines
are shorter and the trochantal spurs are reduced to minute tubercles.

Description based on 1 </ and 2 % taken from Cervus campestris,
at Bauru, East of S. Paulo, Brazil, by Dr Castro Goyanna, 15. xi. 1907,
and 1 cT taken from Cervus rufus at Jacutinga, S. Paulo, in the collection
of Dr A. Lutz, iv. 1907 (dry specimens).

H. hirinaniae 415

The species is apparently allied to H.flava. Through the courtesy
of Dr H. de Beaurepaire Aragao of Rio de Janeiro, we have been able
to examine and redescribe the typos and to figure their salient features.

Types in Rio de Janeiro.

15. HAEMAPHYSALIS BIRMANIAE Supino, 1897.

Fig. 348.

Lit. and Icon. :

H. Urmaniae Supino, 1897, iii, fasc. 1 ; iv, p. 251, PI. XIII, Figs. 17, 18
Supino only figures the $ and $ tarsi which do not serve to distinguish the
species ; his description of both sexes is so meagre as to be useless.

H. hirmaniae Supino, in Neumann, 1897, p. 3.36.

Not identical with H. semermis Neumann, 1901, p. 263, as stated by Neumann,
1902, p. 128, when he condemned his own species after examining Supino's
types (the latter were found by Neumann to include specimens of Am-
bli/omyna besides H. hirmaniae). H. semermis Nn. = ^. hystricis Supino q.v.
Neumann, 1911 a, p. 109, gives the following confusing synonymy :
H. birmaniae Supino + H. hyiitricis Supino in Supino, 1897, ser. 2, v. 3,
p. 236 ; H. punctata (part.) Can. and Fanz. + H. hispinosa Nn. in NeTxmann,
1897, pp. 327, 341 ; H. semermis Neumann, 1901.

It will be seen that we regard the species H. hirmaniae Supino, //. hystricis
Supino, and H. hispitiosa Nn. as valid. H. punctata Can. and Fanz. = H.
cinnaharina var. punctata (C. & F.), and cannot be confused with any of the
foregoing; whilst^, semermis Nn. =ir. hystricis Supino. Because of this
confusion we do not record data from other authors who may have supposed
they were dealing with H. hirmaniae Supino.

Male (Fig. 348): Scutum rather broadly oval, about 2x1-4 mm.^ ;
punctations very numerous and fine, but so shallow that the general
effect is a smooth surface ; cervical grooves only represented by small
oval pits behind the anterior border ; lateral grooves obsolete. Capi-
tulum : base with strong blunt cornua and with slightly rounded lateral
margins ; palps only very slightly salient laterally and without dorsal
spur ; article 2 rather longer than article 3 ; a moderate retrograde spur
under article '4 ; hypostome well covered with 4 ■ 4 to 5 | 5 small, sub-
equal teeth, about 8 per file. Spiracle pear-shaped, or sub-circular,

1 Supino's measurement. The scutums of the 4 <? types examined by us measure
respectively :

1-9 X 1 35 mm.
1-9 X 1-3 „
1-8 X 1-3 „
1-7 X 1-2 „

27—2

416

Genus Haemajyhysalis

with a slight dorsal process. Legs : coxae, especially coxae III and lY,
short and broad (antero-posteriorly) ; coxal armature normal ; well-
marked, but rather blunt trochantal spurs ; tarsus I V tapering rather
rapidly ; pad long.

Female : unknown.

Fig. 348. H. hirvianiae <? . Dorsum, capitulum in ventral aspect, coxae with trochanters,
spiracle and tarsus IV. Drawn from the type. Origiual, N. C. del.

Closely allied to H. hystricis, but without the dorsal spur on
article 3 of the palps, and to H. for^mosensis, from which it is dis-
tinguished by the absence of the lateral groove and its different
dentition.

Our description is based on 4 </ (labelled types) found on Cervulus
muntjac and Atherura macrwa, at Carin-Gheen and Yado-Carin-
Asciuii-Cheba, Burma, by L. Fea in 1885-89, types (Nos. 27,44) in the
Museo Civico di Storia Naturale, Genoa; co-types (N. 2962, 2970) in
Cambridge.

16. HAEMAPHYSALIS SILACEA Robinson, 1911.

Fig. 849.

Lit. and Icon. : Robinson, 1911, pp. 478-480, Fig. 1 «-/(? rejarodviced).

Male : unknown.

Femiale (Fig. 349): Body 24 x 1'7 mm. (unfed), ovate, narrowing
anteriorly ; colour earthy yellowisEi-brown ; dorsum punctate, almost

H. silacea

417

glabrous; marginal groove commencing some distance behind the
scutum and terminating at the antepenultimate festoon; postero-
median and accessory grooves represented by broad shallow depressions ;

Fig. 349. Haemaphysalis silacea ? : (a) dorsum (capitulum foreshortened) ; (/;) capitulum,
dorsal aspect; (c) capitulum, ventral aspect; {d) spiracle (the arrows indicate the
anterior and lateral margins, respectively) ; [e) coxae I-IV ; (/) tarsi I and IV.
/ — same magnification as b and c, d — same magnification as e. (L. E.E. del.)
Bobinson, 1911, Fig. 1.

festoons well-defined. Scutum 1'04 x 1"17 mm., broadly ovate, almost
circular ; colour somewhat darker than the general body colour ; deeply
emarginate ; scapulae rounded ; punctations fine, fairly numerous and
regularly distributed ; cervical grooves of moderate depth, slightly

418 Geims Haemapliy salts

convergent posteriorly. Capitulimi (0"44 mm. long) very short and broad ;
base rectangular, about three times as broad as long ; cornua short, with
rounded extremities ; porose areas well separated, the interval as great
as or exceeding the double diameter ; palps hemicylindro-conical, with
moderate lateral protrusion ; articles 2 and 3 with postero-ventral,
-lateral and -dorsal margins slightly salient ; article 3 with a retrograde
process on the ventral surface only ; hypostome dentition 4 | 4. Veyiter
scattered with pale hairs ; genital orifice opposite interspace between
coxae II and III ; genital grooves widely separated and divergent ;
spiracles sub-circular. Legs : coxa I with a small wide triangular spur;
coxae II and III with an oblique salient ridge ; coxa IV with a wide
triangular spur, larger than that on coxa I ; tarsi tapering gradually,
unarmed ; tarsus IV more than three times as long as broad.

Described by Robinson from 4 $ found by Mr R. J. Davys on oxen
allowed to run on " starvation camp " from which stock had been
excluded for two years at Gonubie Park, East London, Cape Colony,
South Africa.

Types in Cambridge 4 $ (N. 1629, 2944).

17. HAEMAPHYSALIS PARMATA Neumann, 1905.

Figs. 850-353.

Lit. : Neumann, 1905, pp. 228-230 ; described but not figured. Donitz, 1907,
p. 71 ; quotes Neumann. Blanchard, 1909, p. 154 ; description abstracted
from Neumann. Galli-Valerio, 1909, p. 539 ; listed as occurring (?) in Sumatra.
Ziemann, 1905, pp. 116-117 ; 1912, p. 58 ; listed as occurring in W. Africa.
Neumann, 1911 a, pp. 109, 110 ; abstract of author's earlier description. None
of the foregoing authors figure the species.

Male (Fig. 350) : Scutum rather broadly oval, 1*8 x 1*1 mm., strongly
and uniformly punctate with moderate-sized deep punctations ; cervical
grooves short ; lateral grooves very short, including one festoon ; festoons
short and broad, punctate like the rest of the scutum. Capitulum :
base much broader than long, with strong sharp cornua; palps short
and broad, a strong retrograde spur dorsally on article 3, directed
somewhat outwardly ; ventral retrograde spurs under articles 2 and 3 ;
hypostome broad, dentition 4 | 4. Venter : spiracle rather large, sub-
oval, with slight dorsal process. Legs: coxae rather short and broad, with
normal armature very feeble, but with a slight external spur on coxa I

H. parmata

419

and indications of trochantal spurs (not shown in figure) ; tarsus IV
fairly long and tapering; pad medium.

Female (Fig. 351) : Scutum circular, 07 x 0-9 mm. in the specimen
we figure ; cervical grooves broad, parallel, far apart, visible for rather

Fig. 3-50. H. parmata d • Dorsum, capitulum in ventral aspect, coxae, spiracle and tarsus
IV. Specimen (N. 504) from West Africa. Original, N. C. del.

a

Fig. 351. H. parmata ? .
spiracle and tarsus IV.
N.C. del.

Scutum, capitulum in dorsal and ventral aspects, coxae,
Specimen found with the s depicted in Fig. 350. Original,

more than half the scutal length. Capitulum : base short and very
broad, with moderate cornua ; porose areas ill-defined, far apart ; palps
and hypostome as in </. Venter : spiracle sub-circular, sometimes with
a slight dorsal protuberance. Legs as in the </.

420

Genus Hae^naphysalis

Nymph (Fig. 352) : Scutum broad-oval, with deep eraargination in
which the short, broad, basis capituli is deeply set ; cervical grooves
forming conspicuous broad depressions, widely separated, extending
to more than half the scutal length. Capitalam : palps as in the % ,
except that the dorsal spur on article 3 is absent ; hypostome 2 | 2.
Venter : spiracle very small, sub-circular. Legs as in the % , but
relatively thicker.

Fig. 352. H. parmata nymph. Scutum, capitulum in dorsal and ventral aspects, coxae,
spiracle and tarsus IV. Specimen (N. 2354) from Congo Free State. Original,
N.C. del.

Fig. 353. H. parmata larva. Scutum, capitulum in dorsal and ventral aspect, coxae and
tarsus III. Specimen (N. 2354) from Congo Free State. Original, N. C. del.

Larva (Fig. 353) : Scutum essentially like that of the o. Capitulum :
base with cornua obsolete ; palps with rounded lateral contour, most
salient laterally in the middle of their length, otherwise approximating
to those of the o. Legs : coxae almost destitute of armature except for
a fairly marked internal spur on coxa I ; tarsus IV tapering to a point.

H. parmata 421

This species is closely allied to H. Itystricis Supino.

Our description is based especially upon the examination of (N. 1899)
adults from Sierra Leone and of co-types (N. 2887, 2888 (/ % ) from
Cameroon ; the immature stages (N. 2354), now first described, came
from the Congo Free State.

Geographical Distribution and Hosts.

We have received or examined and determined the following
specimens from Africa :

Uganda : (N. 884 c) 2 $ from bushbiick, iii. 1909, and (N. 877) 1 ?
from cattle which came from Bukedi to Mpumu, Chagwe, ix. 1909, both
lots having been sent us by Sir David Bruce. J. H. Ash worth's
collection contains 1 $ from a bashbuck shot at Toro. British East
Africa: (E. 307 6) 1 ? from Jackson's hartebeeste, V -piper Nzoia River,
North Kavirondo, collected vi. 1911, by S. A. Neave. Congo Free
State : (Liverpool N. 45 a) 1 ? from antelope, Kasongo, collected i. 1905,
by Drs Button and Todd ; (N. 2354) (/ O s and larvae from antelope,
Kimaka, 15. vi. 1918, and (N. 2906) a large ^ from buffalo, Beni, 31. vi.
1913, collected by F. Harker. Gold Coast : (N. 504) 2 c^ 1 ? 2 o,
collected in 1908 at Prahsu by Dr W. Graham. Sierra Leone :
(E. 584 = N. 1899) cfs $s os from harnessed antelope, Port Lokko, v.
1912, and (E. 106 a) a $ from Cross River, Obubura, vi. 1912, both
collected by Dr J. J. Simpson. J, H. Ashworth's collections contain 1 %
from cattle, Securella, 19. ii. 1913, collected by Dr J. Y. Wood.
Cameroon : Prof Neumann has presented us with co-types (N. 2887
(/) from Bos taurus, Njanga, 1904, and (N. 2888 $) from Capra hircus,
Mbula, 1904, collected by Dr Ziemann, who also found the adults on
sheep and pig in this region. [Ziemann (1905, pp. 116-117), whose
specimens were determined by Neumann, records that the tick occurs
on cattle at Lagos (Coastal Region), on pigs in Cameroon, on cattle and
goats in Duala (Coastal Region), on cattle, goats and pigs in and about
the Bakossi Mountains and on cattle in the Cameroon Mountains.
Ziemann, i. 1912, p. 58, states that he found specimens on cattle, goats
and dogs in the Cameroon Coastal Region. We learn by correspondence
that he also found the tick on "pigs" i.e. Fotamoclioerus porcus L., at
Akonolinga, 6. vii. 1910; he informs us that the latter were determined
either by Neumann or Donitz.]

[Galli-Valerio, 1909, p. 539, states that DrNarbel found H. parmata
on a monkey in Sumatra, 1909, but the determination appears to us
doubtful, as the species has not hitherto been found outside Africa.]

The types are in Toulouse, the co-types in Cambridge.

422 Genus Haemaphysalis

18. HAEMAPHYSALIS HYSTRICIS Supino, 1897.

Figs. 354-357.

Lit., Synon, and Icon. :

Haemaphysalis hystricis Supino, 1897, III, fasc. 1, pp. 251, 252 ; fasc. 2,
PL XIII, Figs. 19, 20. The author only figures the tarsi, these being of no
use for classification ; his description is valueless.

Haemaphysalis hystricis Supino in Neumann, 1897, pp. 342, 343.

'■'■Haemaphysalis bispiriosa Neumann, 1897" in Neumann, 1901, pp. 261-263.
Description of ^ with notes on $ .

Haemaphysalis hystricis Supino, 1897, in Warburton, 1908, pp. 518, 519, Fig. 11
( $ reproduced). Describes the $ .

Not Haemaphysalis bispinosa Neumann, 1897, pp. 341, 342, Figs. 7, 8, as stated
by Neumann, 1902 a, p. 128, when he wrongly degraded his species, regard-
ing it as identical with H. hystricis Supino. See our description of H.
bispinosa.

Not Haemaphysalis birraaniae Supino as stated in Neumann, 1911 a. Neumann
here describes H. bispinosa.

Male (Figs. 354, 855) : Scutum ovate, narrow in front, broad pos-
teriorly, 3 X 2-2 to 2-54 x 2 mm.^ broadest just in front of the spiracles.
Numerous minute very inconspicuous punctations; cervical grooves
small deep pits, behind the anterior border, followed by shallow divergent
depressions ; no lateral grooves ; festoons only slightly longer than
broad. Capitulum : base rectangular, twice as broad as long ; cornua
strong and rather sharp ; palps very slightly salient laterally ; articles
2 and 3 nearly equal in length ; the lateral salience of article 2 about its
middle and forming an obtuse angle ; a ftiirly strong dorsal retrograde
spine on the middle of the posterior border of article 3 ; strong ventral
retrogi'ade spine on article 3 ; h3rpostome rather broad anteriorly, well
covered with 4 | 4 to 5 | 5 strong teeth of uniform size, about 8 per file.
Venter : spiracle short comma-shaped, with bluntly rounded dorsal
process. Legs : coxae normal ; trochanter I with the dorsal retrograde
spur rather sharp, and with a small sharp ventral retrograde spur ;
tarsus IV sloping rather rapidly, with small spur.

' The scutums of 9 <? measured as follows in mm. :

N. 2956 3-0 x2-2 N. 1060 27 x2-0

N. 1060 2-9 x2-l N. 576 2-6 x2-0

N. 679 2-8 x2-0 N. 1060 2-6 x 1-9

„ „ 2-76 X 2-0 „ „ 2-54 X 2-0

„ 1060 2-75 X 2-05
Lots N. 679 and 1060 are from Malaya, N. 576 from Assam, N. 2956 (co-type) from
Burma.

H. hystricis

423

Fig. 354. K. hystricis i . Dorsum, part of venter, hypostome (highly magnified), spiracle
and tarsus IV. Specimen (N. 679) from Federated Malay States. Original,
G. H.F.N, del.

Fig. 355. H. hystricis i . Dorsum, venter and spiracle. Drawn from the type of
H. scmermis Nn. in the Paris Museum. Original, G. H. F. N. del.

i

424

Genus Haemaphysalis

Female (Figs. 356, 357) : Scutum. 1-4 x I'l to 08 x 0-8 mm.\ usually
sub-circular, with fairly numerous moderate punctations, more conspicu-
ous in the lateral fields ; cervical grooves forming pits at some distance
from the anterior border, followed by fairly well-marked sub-parallel

Fig. 356. H. hystricis ? . Scutum, capitulum in dorsal and ventral aspects, coxae
with trochanters, si^iracle and tarsus IV, Specimen (N. 576) from Assam. Original,
G. H. F. N. del.

Fig. 357. H. hystricis ? . Dorsum, etc. (not drawn to scale). Sketched by C. W. from
a specimen lent by Prof. Neumann. Warburton, 1908, Fig. 11.

depressions not reaching the posterior border. Capitulum with the
characteristics of that of the </ ; porose areas oval, far apart, a depres-
sion in the middle of the interval ; palps with article 2 longer than 3,

1 The scutums of 6 $ measured as follows in mm. :

N. 2956 1-4 X 1-1 N. 572 1-1 x 1-3

N. 572 1-2 X 1-35 N. 1070 0-95 x 1-1

N. 576 1-2 X 1-2 N, 576 0-8 xO-8

Lots N. 2956 (co-types) from Burma, N. 572 and N. 576 from Assam, N. 1070 from
Malaya.

H. hystricis 4t2f)

the lateral salience more pronounced and more acute than in the (/ ;
palpal spurs as in the </ ; hypostome 4 1 4, about 10 teeth per file.
Venter: spiracle pear-shaped, the narrow end dorsal. Legs as in the cf .
Fully gorged specimens may attain 11 x 8"5 mm.

Our description is based on the examination of the types and
co-types, some of the latter (N. 2956, 2957 </ $) having been pre§ented
to us by the Genoa Museum.

Geographical Distribution.

We have received or examined and determined specimens from the
following countries^ :

Burma: Supine's specimens, (/ $ (Genoa Mus. Nos. 20, 21, 51),
of which we possess co-types (N. 2950, 2957), were found on Hystrix
hengalerisis and Ursus torquatus at Yado-Carin-Asciuii-Cheba and at
Carin-Cheba (900 m. elevation), by L. Fea in 1887-1889; (Indian Mus.
1798 b/17) 1 (/ found at Arakan Yomas, Chauung, Upper Myinudaung
Reserve, Hedzada District, Lower Burma, 2. I. 1912, by C. G. Rogers;
(Indian Mus.) 1 $ found on Geoemyda spinosa. Assam: (N. 572) 3 $
from Canis familiaris, Lhassia, and (N. 576) 1 cT 3 $ from the same host,
Lushai Mountains, both lots having been collected in 1907 by R. A.
Lorrgin (presented by the Hon. N. C. Rothschild). We have determined
specimens from the Indian Museum (N. 1253/17) collected at Sadiya in
N.E. Assam, Xli. 1911, and also at Sibsagar, hosts not recorded. China:
(D.E.M. No. 13 = N. 2905, N. 2977) ?s from hedgehog, Tsingtau, Prof.
Hoffmann coll. Federated Malay States : (N. 1060 6) J^s from tiger,
Pahang, 1910, collected by Dr A. T. Stanton who also found this tick on
a tuild boar; (N. 2106) (^s $s from tiger, Kuantan, Pahang, I. 1913,
collected by Dr Oscar Pou ; Messrs J. H. Ash worth and H. C. Pratt
have also sent us (N. 1070 a) 1 $ from Kelantan, and (N. 679) 2 c/
from an unrecorded place. Mr C. Strickland has sent us (N. 2953) 2 $
from Ursus malayawis, iv. 1913, and (N. 2952) ^f $ from dog, xi. 1913,
both collected at Ulu Gombak, Selangor ; (N. 2951) 1 % from dog, Port
Dickinson, North Sembilan, ii. 1912. Sumatra: (N. 1274) 1 (/ from
dog, V. 1911, presented by the late Prof W. Donitz. Borneo : (N. 954)
2 % from a dog. Upper Sarawak, xi. 1909, collected by Mr J. C. Moulton.
Ceylon : (Indian Mus. No. 847/17) 1 ? fi-om Kandy. Celebes : (N. 72)

1 Owing to the confusion in the synonymy, we confine ourselves to mentioning
only such specimens as we have ourselves examined.

426 Genus Haemaphysalis

1 $ from Sas celebensis, presented by Hon. N. C. Rothschild. Formosa :
(Berlin Mus. No. 174) ^ $ from wild hoar, Taihorin.vi. 1911, collected
by H. Sauter.

19. HAEMAPHYSALIS BISPINOSA Neumann, 1897.

Figs. 358-362.

Lit., Syn. and Icon. :

Haemaphysalis bispinosa Neumann, 1897, pp. 341-342, Figs. 7, 8 ( $ palp in

dorsal aspect, tarsus IV, reproduced). Blauchard, 1909, p. 148 ; merely

listed.
Not Haemaphysalis bispinosa Nn. in Neumann, 1901, pp. 261-262.
Not Haemaphysalis hystricis Supino, as stated in Neumann, 1902 a, p. 128.
Haemaphysalis bispinosa in Warburton, VI. 1907, p. 11, Fig. 9 (poor) incomplete

description ; 1908, pp. 517-518, Figs. 9-10 {^ dorsum, part of venter,

spiracle ; 9 scutum and capitulum, spiracle and tarsus IV ; sketchy, not

reproduced).
Haemaphysalis neuvianni Donitz, 1905, pp. 127-129, 134, Figs. 4-6 (reproduced).

We have co-types.
Not Haemaphysalis birmaniae Supino, as stated in Neumann, 1911 a, p. 109.
Haemaphysalis neumanni Donitz, in Neumann, 1911 a, p. 109.
1 Haemaphysalis neumanni Donitz, in Galli-Valerio, 1909, p. 539, found on

Canis aureus, Ceylon, 1907, Dr Narbel coll. ; Yakimoif and Kohl-Yakimoff,

1911, p. 418. (They record 6 $ from cattle coming from China, Primorsk

Government, E. Siberia.)
Haemaphysalis bispinosa in Patton and Cragg, 1913, p. 648 (raised on onongoose),

PI. LXXIII, Fig. 11 ; 9 capitulum in ventral aspect ; PI. LXXXI, Figs.

5, 6, ^ dorsum and (?) $ venter (legend confused).

There has been a great deal of confusion about this species: Neumann
(1897) founded the species bispinosa on a single % from Ramnad, India,
the description being fortunately accompanied by figures (Fig. 361 a
and h here reproduced). In 1901 he described the </ and $ of what
was evidently another species under the name of " bispinosa" ; these ticks
came from Japan and China. The second description agrees with that
of H. hystricis Supino, 1897. In 1902, Neumann, after examining
Supino's types, regarded bispinosa as identical with " hystricis" and
(wrongly) gave the latter name priority. He had evidently been led
astray by his second description of the purported "bispinosa" and 'the
fact, which he mentions, that Supino's types {^ $ ) are devoid of
capitulums.

Donitz (1905), like ourselves, perceived the confusion which had
arisen, and to clear up matters selected one of the forms described as

H. hispinosa

427

' hispinosa " by Neumann as the type of a separate species, which he
named neumanni. The form Donitz selected is, we find, identical with
H. hispinosa Neumann, 1897, and therefore falls into synonymy (see
Fig. 359 from Donitz). Warburton (1908) restored and redescribed
H. hispinosa.

Male (Figs. 358, 359) : Scntiwi 1-45 x 1 to 2-3 x 1-65 mm.', yellow
or brown (in well chitinised specimens), long oval, broadest in the middle,

Fig. 358. H. hispinosa s . Dorsum, venter, tarsus IV, spiracle. Specimen (N. 1826
from Madras). Original, N. C. del.

glabrous, uniformly punctate with medium inconspicuous punctations ;
cervical grooves very faint, lateral grooves long,well- marked, ending behind
spiracles ; festoons very long. Venter : spiracles white, sub-oval, broadest
posteriorly, almost without dorsal protuberance. Capitulum : base fairly
long, broadly rectangular, with straight sides and stout cornua;
palps only slightly salient, articles 2 and 3 about equal in size and

1 The following measurements show the variation in the size of the c? scutum in mm. :

N. 1826, from India

N. 1373, from ,China

1-8 xl-1
1-8 X 1-0.5
1-7 xl-1
1-65x1-15
1-7 xll
1-45 X 10

chosen for range of size from amongst 83 <f
contained in the lot.

N. 129, from Japan 2-3 x 1-65
205 X 1-0

428

Genus Haeinaphysalis

equally angular externally; no spines on article 2, a short, sharp
retrograde spine at the middle of the dorsal border and at the middle of

Fig. 359. H. bispinosa <? : (A) dorsum, x 12 (the original figure showed eyes which are
absent, they have now been erased; vide Donitz, 1906, p. 143) ; (B) part of venter
and — (C) capitulum more highly magnified. Donitz, 1905, PI. Figs. 4-6, illustrating
H. neumanni Do., 1905 (from horse, Japan), which we find = H. bispinosa Nn.

the ventral border of article 3 ; hypostome 4 | 4 to 6 | 6, with equal
teeth. Ler/s : a moderate sharp spur on coxa I ; a very slight pro-
tuberance on coxae II-IV.

Female (Figs. 360, 361): Scutum yellow or brown, nearly circular,

H. hispinosa

429

0"8 X 0*8 to r25xl*2mm.^ with unbroken contour; cervical grooves
beginning rather behind the anterior mai-gin as fairly deep furrows
which converge and then curve outward, almost reaching the postero-
lateral border; numerous regularly distributed punctations of medium
size. Venter: spiracles white, nearly circular, practically without dorsal
protuberance, Gapitulum : base broader than in the </, the sides

Fig. 360.

Fig. 361.

Fig. 360. H. bispinosa ? . Capitulum in dorsal and ventral aspects, scutum, coxae with

trochanters, spiracle and tarsus IV. (N. 1826, same origin as i in Fig. 358.) Original,

N. C. del.
Fig. 361. H. hispinosa ? : (a) right palp in dorsal aspect, x 65 ; [b) tarsus IV, x 75.

Neumann, 1897, Figs. 7, 8, illustrating his original description of the species (clearly

drawn from balsam-mounted specimen).

straight, the cornua short and blunt ; porose areas fairly large, far
apart, oval, not very clearly defined ; palps as in the ^ ; hypostome 4 | 4
to 6 I 6, with teeth of equal size. Legs : coxae as in (/ ; tarsus IV
long (at least four times as long as broad), tapering gradually.

^ The scutums of 12 ? measured as follows in mm.

N. 1826, from India 0-95 x 0-9
0-85 X 0-9
0-85 X 0-8
0-8 xO-8
0-8 xO-85
0-7 xO-7

N. 129, from Japan
N. 1852, from Australia

1-0 xl-1
1-25 X 1-2
1-15 X 1-15
1-1 xl-25
1-1 xl-2
1-1 xl-2

28

430

Genus Haemaphysalis

Nymph (Fig. 362) : Body elongate. Scutum sub-circular, about
0'2 X 0'2 mm.^, a few faint punctations generally visible in the posterior
region ; cervical grooves well-marked, not very concave externally, not
quite reaching the posterior border. Capitulum : like that of the ? ,
but without an erect dorsal spine on article 3 ; its inner edge is, how-
ever, recurved (as in the $ of II. hispinosa var. intermedia) ; hypo-
stome 3 | 3. Venter : spiracle nearly circular. Legs : coxal armature
normal ; tarsus IV fairly long and sharply pointed ; pad short.

Fig. 362. H. hispinosa nymph. Capitulum in dorsal and ventral aspects, scutum, coxae
with trochanters, spiracle, tarsus IV. (N. 1826, same origin as c? in Fig. 358.)
Original, N. C. del.

Fig. 362 a. H. Msjiinosa larva. Capitulum in dorsal and ventral aspects, scutum, coxae
and tarsus III. Drawn from (N. 2745) specimens raised in Cambridge, the progeny
of a ? from Madras. Original, G. H. F. N. del.

1 The scutums of 5 nymphs measured as follows in mm. :

N. 169, from India 0-21 x 0-23 N. 169, from India 0-18 x 0-24

0-21 X 0-23 0-18 X 0-2

0-2 xO-21

H. Mspinosa 431

Larva (Fig. 3G2«) unfed, measures about 0*53 x 0'44 mm. Scutum
broader than long, about 0'25 x 0'8B mm., with undulating postero-
lateral borders; cervical grooves distinct, parallel. Capitulum with
base rectangular, posterior border almost straight, no cornua; palps
with article 2 protruding more than in the o; hypostome 2 | 2. Legs:
coxa I with blunt tuberosity postero-internally, coxae II and III
unarmed ; tarsus IV short, tapering to a point.

Description based on numerous specimens from India and Ceylon,
where it appears to be common.

Geographical Distribution and Hosts.

India : The tick is widely distributed, occurring apparently
throughout India. We possess, or have examined, specimens col-
lected as follows : Punjab : at Kasauli (Liverpool 149 a). United
Provinces : Mr Abdal Ghani collected specimens from goats, at Hald-
wani, VI. 1905 (Liverpool coll.) ; Mr R. Hodgart collected adults from
Felis tigris, at Motisal, Garthwal District, III. 1910 (Indian Mus.).
Nepal: (Ind. Mus. 1082/17) </, from Felis tigris, Neapalganj, near the
Frontier, ii. 1911 ; (Ind. Mus.) $ , from Felis tigris, Burdwar, Nepal Terai.
Bengal : (Ind. Mus. cc/367 c.) cf , found at Singla, Darjiling District, iv.
1913, collected by Lord Carmichael ; (Ind. Mus.) cT $, from dog, at
Bhogaon, Purneah District ; (Ind. Mus. 917/17), found at Katihar,
Purneah District, by C. Paiva ; (Ind. Mus. 999-17) from buffalo,
Purneah District, L. R. Paiva, XI. 1910 ; (London Sch. Trop. Med.) from
dog, Calcutta, collected by C. R. M. Green; (Ind. Mus. 1079/17 =
N. 1804), cT from dog, Calcutta, ii. 1911 ; (N. B. Kinnear) $ from
Canis aureus, Andheri, near Calcutta, xii. 1908; (Ind. Mus. 2140/17 =
N. 2864) adults from Viverricula malaccensis (Gmel.), Satpara, Puri
District, Orissa, 17. viii. 1914, Dr N. Annandale coll. Eastern Bengal
and Assam: (N. 890), from the Chin Hills, just east of Chittagong,
1909, H. M. Lefroy coll. ; (N. 892), from Canis familiaris, Southaia Mts. ;
(N. 582 and 583), from Eqaus caballus and Talpa sp., Lushai Mts.,
collected vi. 1907 by R. A. Lorrain ; these three lots were presented
by the Hon. N. C. Rothschild ; (Ind. Mus. 1202/17, 1244/17) ^ ?, from
N.E. Assam, Xl. 1911, found on grass and on floor at Sadiya re-
spectively. Madras Presidency : Beginning in the north : (Ind. Mus.),
(/, from dog, Berhampur, i. 1910; (Ind. Mus.) $, from grass at Lake
Chilka, collected in. 1910 by Dr A. Annandale ; (N. 1826), c^ % o, from
a cow, Madras collected in 1912 by Mr J. F. Valladares; (N. 167 and

28—2

432 Genus Haemaphysalis

168) Avere collected by Colonel C. E. Nuthall, A.V.C., xi. 1906, from
cattle, at the slaughter-house and from a goat at Madras; (N. 1906), $,
from cattle, collected vii. 1906, at Bellary by Mr F. M. Hewlett ; (N.
343) cT ?, from Coimbatore, collected in 1906 by Colonel C. E. Nuthall,
A.V.C. ; (N. 1958) cT %, from sheep, at Agricultural Farm, Coimbatore,
collected in 1912 by Mr T. Bainbrigge Fletcher; (N. 2745) ?, from
calves, Guindy, in. 1914, Captain W. S. Fatten coll. Travancore :
(Ind. Mus.) %, from Macacus sinicus, at Maddathory, western base of
Western Ghats; (N. 1253 a) cT %> from dog, place not stated, collected
III. 1911, by Mr E. R. Hewlett; (Ind. Mus. TM/3, TM/6) $s, from
bullock, collected xii. 1911, at Nagercoil ; (N. 3072 = Ind. Mus. 2173/17)
^ $ from Hemitragus hylocrius, Parambikulam, Cochin State, ix.
1914, F. H. Gravely coll. Mysore: (N. 1071 h), ^ ?, from cow.
Bangalore, collected i. 1910, by Mr W. H. Marshall. Bombay
Presidency : (N. 863), from calves, and (N. 869) from foxhounds, at
Belgaum, collected in 1909, by Captain F. H. C. Hutchinson, I.M.S. ;
(Ind. Mus. 1117/17) d", found at Gaunter, Panwell, ii. 1911 ; (N. 2529 h),
(/ $, from bullocks, collected x. 1913, at Poena by Major Eassie and
presented by Colonel E. R. C. Butler ; (Mr Hewlett's coll.) adults from
lion, shot 11. HI. 1909, in Gaur Forest, Kathiawar ; (N. 2002) ?, from
Felis tigris, Kadra, Kanara, xi. 1910, N, B. Kinnear coll. Ceylon :
Dr Aide Castellani, in 1906, sent us four lots (N. 150), from dogs,
Nugegoda; (N. 148) from polecat, KoUupitiya ; (N. 151) from Mouse-
deer, Cinnamon Gardens ; (N. 152) from pig. Government Model Farm,
near Colombo. Colonel B. Skinner, R.A.M.C, in 1906, sent us three
lots; (N. 186 a) from country cat, Borlesgama; (N. 187) from dog;
(N. 188) from black monkey, Raygam Koili. Mr E. E. Green sent us
two lots (N. 510) $s, from dog ; (N. 515 6) $s, from cattle, Matara, vii.
1906. We have a specimen (N. 891) $ , from cattle, presented by
Mr E. G. Wheler in 1906. We have determined (E. 876 a = N. 2901,
E. 771) adults and nymphs found on the goat, at Hakgala, 23. vii. 1913
and 27. ill. 1914, and (E. 708) on cattle, at Peradeniya, 31. v. 1913,
by A. Rutherford.

Burma: (N. 1384 a) from goat, Rangoon, collected ix. 1911, by
Dr H. H. Marshall. Andaman Islands : (Ind. Mus. 1118/17) ^, from
dog, at Ross, collected iii. 1911, by O. Paiva. Federated Malay States :
Dr W. M. Scott has sent us (N. 2459) cfs $s, found on fowls, in. 1913,
at Singapore. China : We have received through Mr T. V. Sherrin
(N. 1372) 1 $, from Nemorhaedus cinereus, Jehang, and (N. 1373) 6 cT
5 $ , from Cervulus sp.. Fen Tuang Shan, An- Wei ; we have, moreover.

H. blsplnosa 433

determined numerous specimens (Berlin Mus. No. 179) from Chinese
roebuck {II ydropotes inermis), Herr Lemm coll., and (D.E.M. 3, 4, 10 ;
N. 2905, 2976) from dog and other (?) hosts, collected at Tsingtau,
7. VIII. 1914, by Prof. Hoffmann.

Japan : The late Professor W. Donitz presented us with co-types
(N. 129) of his H. neumanni, </ and ?, found on the horse. Dr M.
Miyajima has sent us (N. 2911 a) \ % from cattle, Tokio, 16. x. 1911,
found with Boophilus ; (N. 2907 a) $ s from horse, Aomori, North-Central
Japan, 15. vii. 1912, found with Dennacentor ; (N. 2908) adults and
larvae, from horse, Daisen, N.W. of Central Japan, 2. viii. 1912, found
with Ixodes ricinus; (N. 2909) $s from horse, Takanabe, Kiushiu, 5.
VI.-12. X. 1912. Borneo: (N. 307 6), from wild red-deer, Kuching,
Sarawak, collected vil. 1907, by Dr A. R. Wellington. Australia : We
are indebted to Dr J. Burton Cleland for (N. 1851) $ , from horse,
Ballina, N.S. Wales, x. 1911 ; (N. 1852) $ , from ccdf, Bonville. British
East Africa : Dr J. G. Parham, iv. 1912, collected specimens (N. 1787 b,
1781 d), c/s and $ s respectively, from bullocks at Bet-el-Ras and Chuck-
w^ani ; (N. 1783 a) $, from a goat at Mahonda ; (N. 1790 6) $, from a
bidl at Kikwaguni ; he also found a $ on a bullock at Bububu.
(N. 2846= E. 870) </, from imported Indian (Bombay) cattle, Zanzibar,
2. IX. 1913, Dr W. M. Aders coll. We have also determined specimens
collected by Mr Last in Zanzibar, x. 1905, which are in the collection
at Liverpool (Liverpool coll. Nos. 74 and 87) $s and nymphs from cattle,
and c/s, $s fi-om goat, and specimens (cf $ Berlin Mus. No. 71) labelled
merely " Zanzibar." From the foregoing records it would appear that
H. bispinosa has recently been imported into Africa as it has not pre-
viously been reported from this continent.

Haemaphysalis bispinosa var. intermedia
Warburton and Nuttall, 1909.

Figs. 363, 364,

Lit. and Icon. : Warburton and Nuttall, 1909, pp. 69-70, Fig. 16 (reproduced).

Differs from the type in little else than the structure of the palps \
In the $ (Fig. 364) the third article, instead of bearing a somewhat

1 The scutums of 4 c? and 4 ? measured as follows in mm. :

cT ?

1-8 X 1*15 0 "9 X 0-9 (3 specimens)

1-75 X 1-15 0-8 X 0-8

1-7 xl-2
1-6 xl-05

434

Genus Haemaphy sails

erect dorsal retrograde spine at the middle of its posterior border, has
its internal angle produced to a retrograde point, hardly raised above
the dorsal surface of the palp. In the cT (Fig. 363) the point may be
altogether absent, the postero-internal angle of article 3 being bluntly
rounded. The spiracle is, as a rule, somewhat narrower and more
transverse than in the type.

This variety was described from specimens marked * ( ? types)
in the subjoined list relating to the geographical distribution ; it
was named intermedia because it seemed to connect the group with
a dorsal spur on article 3 of the palp with another group, destitute of
such a spur, of which H. campanulata is the type.

Types in Cambridge (N. 1686 cT I N. 34 d and h; N. 327,
N. 515 a $). We presented co-type N. 34 /i $ to the collection of
the London School of Tropical Medicine.

ZI=^

Fig. 363. Fig. 364.

Fig. 363. H. bispinosa var. intermedia t? . (N. 1686) capitulum of type. Original,
N. C. del.

Fig. 364. H. bispinosa var. intermedia ? . Capitulum and scutum, tarsus IV, spiracle
and ventral aspect of right palp. Warburton and Nuttall, 1909, Fig. 16. Sketch by
C.W.

Although widely distributed in India, this variety has not reached
us from the same places as H. bispinosa, and it does not appear to
be as common. We have received the following specimens: (*N. 327),
from a cat, Agra, collected in 1907 by Dr E. H. Hankin ; (*N. 1686,
cT types), from Lepus ruficaudatus, collected, ill. 1911, at Shamghar
by Mr N. B. Kinnear.

The following were collected in Cutch by Mr N. B. Kinnear: —
(N. 1680), 3 c/ 1 $, from Ganis pallipes, at Charwa, 1911 ; (N. 1682),
(/s, from Felis caracal, at Bhuj, viii. 1911; (Kinnear No. 289 6), c/*,
from Felis caracal, at Dhonsa, 1911 ; (Kinnear, No. 348), from

II. blspuiosa var. intcniwdia 435

Erinaceus micropus, at Bhuj, vii. 1911; (Kinnoar No. 241a), ^ %,
from Felis ornata, at Bhuj, 1911; (Kinnear No. 264 a), ^/'s, from
Felis affi,nis, at Nohaman, 1911. Major O. A. Smith collected on
9. viii. 1912 numerous (/"?"! Chota Nagpur (Indian Mus. 1524/17 =
N. 2056, N. 2077), from a wolf, four miles west of Koderma Station.
Mr N. B. Kinnear collected three lots in Kandeish, at Ghodasgaon,
V. 1911, namely (N. 1665), (/, from Hyaena hyaena: (N. 1666) cTs,
from Felis affmis ; (Kinnear, No. 402), $, from Millardia meltada^ ;
also one lot at Pili, Sipna Valley, Berars ; (N. 1672) cT ?, from Cyan
dukhunensis. Colonel C. E. Nuthall, A.V.C., sent us several lots
(*N. 34 rf, /<) collected in 1906 in the Bangalore, Mysore, Secunderabad
and Hyderabad Districts, the hosts not having been recorded. One lot
(N. 3071 = Ind. Mus. 2172/17) (^ $ was found on Sus cnstatus, nesir
Barhi, Hazaribagh Distr., Chota Nagpur, 21. vii. 1914, by Major
O. A. Smith.

One lot has reached us from Ceylon (*N. 515 a), from cattle, at
Matara, collected, vii. 1906, by Mr E. E. Green.

20. HAEMAPHYSALIS PARVA Neiunann, 1908.

Figs. 365, 366.

Lit. and Icon. : Neumann, vii. 1908, pp. 89-91, Fig. 10 (reproduced). Blanchard,
1909, p. 154, and Galli-Valerio, 1909, p. 539 (listed only).

Male (Figs. 365, 366) : Very small, body measuring 11 x 08 to
1"4 X I'O mm. Scutum a rather broad oval, broadest posteriorly, the
posterior border being rather flat or truncate; strongly punctate, the
punctations deep and often confluent ; cervical grooves shallow ; lateral
grooves medium, including no festoons ; festoons fairly long, well-marked.
Capitulum : base rectangular, relatively rather long, cornua strong ;
hypostome short, with dentition 4 4; palps with articles 2 and 3
of about equal length, article 2 not very salient laterally, without
spines ; article 3 with strong ventral retrograde spine and a dorsal
tooth not very erect and somewhat internal. Venter : spiracle sub-
circular. Legs : a strong spine on coxa I, a slight internal spur on
coxae II-I V ; tarsi medium, not humped ; pad long ; the hairs along
the ventral surface of the legs are longer and more conspicuous than in
H. bispinosa.

1 Family Muridae.

436

Genus Haemaphy sails

I

Female : Body oval, red-brown or yellowish. Scutum oval
0'7 X 0*65 mm., with punctations as in the (/, a little lighter than
the rest of the body ; cervical grooves well marked, spiracle as in

Fig. 365. H. parva 3 . Dorsum. Drawn from co-type No. 1038 in Galli-Valerio coll.
Original, N. C. del.

Fig. 366. H. parva i . Capitulum in ventral aspect with coxae I. Neumann vii. 1908,
Fig. 10.

the cT- Dorsum conspicuously hairy. Capitulum 0"33 mm. long ; base
more than twice as broad as long ; cornua strong ; porose areas small,

H. parva 437

indistinct, far apart ; liypostouie 4 1 4 ; palps as in the (/ . Legs
medium, as in the cT-

Nymph and Larva: These only differ from the corresponding
stages of H. bispmosa (sec pp. 430, 431) in having stronger cornua
and more hairy dorsum and legs.

The species was originally described from 16 (/ 8 $ 3 o and 7 L,
from Ganis aureus, Ceylon, 1907, collected by Dr Marbel. The types
and co-types are in Toulouse, in Prof Galli-Valerio's collection at
Lausanne and in Cambridge (N. 2889 cT %), the last having been
presented to us by Prof Neumann.

Prof Galli-Valerio has kindly allowed us to examine all of his
specimens of H. parva (Nos. 1037, 1038), which are co-types if not
types. No. 1037 contained 2 (/ iT. parva and 2 $s which differed
much from each other and do not belong to the </s; one was
H. hystricis $ , and the other was indistinguishable from H. bispinosa $ ,
No. 1038 contained 10 cT, 1 $, 2 o and 4 L; all of which agree fairly
well together and are doubtless the various stages of the tick named
H. parva by Neumann.

When the dorsal spur on article 3 of the palps is added to
Neumann's description, the likeness to H. bispinosa is very close,
and it is not without hesitation that we regard it as specifically
distinct. Besides the difference in size, the (/s have a different
facies, H. parva (/ being short and broad, truncated posteriorly, and
more deeply punctate than H. bispinosa. The $s, os and L have
stronger cornua and are distinctly more hairy.

We have determined the following as H. parva : (Ind. Mus.
2141/17, 2143/17 = N. 2865 and 2931a) c/s $s from Felis viverrina
Bennett and Lepus rujicaudatus Geoff., Satpara, Puri District, Orissa,
India, 16. in. 1914, coll. Dr N. Annandale ; (Berlin Mus. = N. 2822)
(/ $ , origin unknown, collected by Schoede ; (N, 3073 = Ind. Mus.
2176/17) (/s $s from Lepus sp., Satpara, Lake Chilka, Orissa, ix. 1914,
Chilka Survey coll. ; (N. 2388 b) cTs $s from paiither, no locality stated
presented by R. Hancock ; this lot contained some large specimens.

438

Genus Haenia2)hysalis

21. HAEMAPHYSALIS CUSPIDATA Warburton, 1910.

Figs. 367-370.

Lit. and Icon. : Warburton, xii. 1910, pp. 401-402, Figs. 6, 7, ^ -^nd ? (repro-
duced).

Male (Fig. 367). Scutum: 1-8 x ll mm., fairly glossy, with
numerous very small, shallow, discrete punctations; cervical grooves
deep under the cornua, then faint and gently diverging ; lateral grooves
medium, faint, including one festoon ; festoons long, curved, with
distinct broad intervals widening distally. Capitulum : base rectangular,
broader than long, with straight sides and very long cornua, longer
than the base (hence cuspidata) ; palps of medium length ; article 2
fairly salient laterally, unarmed, article 3 with long dorsal retrograde
spine reaching to the posterior border of article 2, and almost equally
long ventral spine, article 4 unusually long ; hypostome 4 1 4. Ve)iter :

oa

cQo^

Fig. 367. Fig. 368.

Figs. 367 and 368. H. cuspidata. (Fig. 367) <? dorsum, capitulum in ventral aspect,
coxa and trochanter I, palp in profile, spiracle, anal grooves. (Fig. 368) ? capitulum
and scutum, palp in profile, spiracle. (Sketches by C. W.) Warburton, 1910, Figs. 6, 7.

genital oriiice between coxae II ; anal grooves ogival ; spiracle sub-
triangular, with narrow rounded anterior border and nearly straight
posterior border. Legs : coxa I with long narrow spur, coxae II-IV
unarmed, trochanter I with a strong rather sharp retrograde spur
both dorsally and ventrally ; otherwise as in H. aculeata (p. 440).

Female (Fig. 368) : Body dark-brown, very punctate. Scutum :
yellowish, nearly circular, 0*96 x 13 mm., numerous medium-sized
punctations ; cervical grooves rather long and slightly converging
till they reach the middle of the scutum. Capitulum : base about
twice as broad as long, with long cornua, but not so long as in the </ ;

H. cusjndata

439

porose areas rather large, oval, converging in frcjnt, separated by more
than their diameter; palps with the same characteristics as those
in the cT, but the ventral spur on article 3 longer than the dorsal;
hypostome 4 | 4. Venter: spiracle nearly circular, but rather pointed
dorso-laterally ; vulva between coxae II and III. Legs: as in the (/'.
Nymph : (Fig. 369) : Scutum : 0-42 x 0-48 mm. Capitulum :
without dorsal spur on article 3 of the palp ; otherwise like the $
in such structures as are present. The long comua give it a very
characteristic appearance.

Fig. 369. H. cuspidata. Nymph. Scutum, capitulum in dorsal and ventral aspects,
coxae with trochanters, spiracle and tarsus IV. (N. 2692 type.) Original, N. C. del.

Fig. 370. H. cuspidata Larva. Scutum, capitulum in dorsal and ventral aspects, coxae
and tarsus III. (N. 2692 type.) Original, N. C. del.

Larva (Fig. 370) : Scutum : 0*25 x 0'31 mm., cordate, emargination
deep ; cervical grooves well marked for half the scutal length, parallel.
Capitulum : base sub-rectangular, with cornua only slightly developed :
palps as in the nymph, but less salient laterally. Legs : coxae

440 Greniis Haemaphysalis

practically unarmed ; tarsus III tapering finely ; pad short. There
is little to indicate the connection of the larva with the adult.

Described from (N. 1108) 30 (/, found (in company with
U. aculeata) on Tragulus memmina (the " Mouse Deer "), at Colombo,
Ceylon, 3. viii. 1909, by Mr C. C. Dobell ; and (N. 2692) 2 ?, 7 os
and 4 larvae, found on Paradoxurus niger (Palm civet) at Colombo,
Ceylon, 16. ii. 1910, Dr A. Willey coll. We have since received
(N. 2766) 3 (/ 1 $ , found on Herpestes mungo, Peradeniya, Ceylon,
II. 1912, J. S. Fryer coll.

This species much resembles H. aculeata in most respects, but
it is easily distinguished by the capitulum.

Tj^es in Cambridge (N. 1108, cfs; N. 2792, 1 $ 7 o 4 L).
We have presented co-types (N. 1108 cf) to the Zoological Museum,
Berlin, and to the Neumann collection, Toulouse.

22. HAEMAPHYSALIS ACULEATA Lavarra, 1905.

Figs. 371, 372.

Lit., Icon, and Synon. :

H. aculeata Lavarra, 1905, pp. 255-258, no figure {^)-

H. longipalpis Warburton, 1910, pp. 399-400, Figs. 4, 5 ( (J and $ reproduced).

H. aculeata Lavarra, in Neumann, 1911a, p. 110 ; no figure.

Male (Fig. 371). Scutum 2 x 1-3 mm.\ glossy, with numerous
very small punctations, many of them confluent ; cervical grooves
deep to receive the cornua, barely visible behind them ; lateral grooves
rather short and faint, including one festoon ; festoons long, with well-
marked curved intervals. Capitulum : base sub-rectangular, with
rounded sides, broader than long, with cornua of almost equal length ;
palps very long, three times as long as broad : article 2 very slightly
protuberant laterally, unarmed ; article 3 with a long, strong retro-
grade spine doi'sally and ventrally ; hypostome small, with numerous
very small sharp teeth, apparently 6 | 6, considerably shorter than the
palps. Venter : genital aperture between coxae II ; spiracles sub-
rectangular, more rounded anteriorly ; anal grooves rather ogival.
Legs : coxae I with a rather long blunt spur ; coxae II-I V unarmed ;
trochanter I has a distal retrograde spur both dorsally and ventrally,

^ The scutum of (N. 2731 co-type) the s presented by Lavarra measures 1-7 x 1-2 mm.
but the specimen appears somewhat shrivelled.

H. aculeata

441

the former blunt, the latter shorter and more pointed ; tarsus IV
roundly tapering ; pads large.

Female (Fig. 372). Scutum nearly circular, 12 x 1-2 inm. ; cervical
grooves deep pits behind the cornna, followed, at a short interval, by
broad, shallow, nearly parallel depressions extending slightly beyond
the middle of the scutum ; numerous very small punctations. Capitu-
luni : base rectangular, twice as broad as long, porose areas much
longer than broad, very far apart ; cornna short and blunt ; palps very

'vxj:)

Fig. 371. Fig. 372,

Uaemaphysalis aculeata.
Fig. 371 <?. Dorsum, tarsus IV, spiracle, palp in profile.

Fig. 372 ? . Capitulum and scutum, palp in profile, spiracle. Warburton, 1910, Figs. 4, 5
('-ketches ; scutums to scale).

long, four times as long as broad, nearly cylindrical, armed as in the
(/, but with the dorsal retrograde spur of article 3 shorter; hypostome
5 I 5, well covered with small teeth ; spatulate. Vente7~ : spiracle smaller
than in the cf and more pointed dorso-laterally. Legs: as in the (/.

Described from 10 cf and 1 $ taken by Mr C. C. Dobell from
Tragulus memmina (the Mouse Deer), at Colombo, Ceylon, 3 viii. 1909,
and named H. longipalpis by Warburton (1910). The cT was described
before as H. aculeata by Lavarra (1905) ; the latter name has, therefore,
priority. Lavarra's specimens (16 cT) were found on Tragulus memmina
Bodd. in the East Indies, the host, which died in a menagerie, having
been purchased by the Zoological Museum, Rome, A specimen kindly
presented to us by the author (N. 2731, co-type) agreed fully with the
foregoing description of the (/,

This remarkable species, though undoubtedly a Haemaphy salts, has
the abnormal character of palps much longer than broad. It belongs
to the H. bispinosa group.

442

Genus Haemaphysalis

Types and co-types (cT) in Lavarra's collection, Rome, in Toulouse
and Cambridge. The types oiH. longipalpis Warb. (N. 1107, </ $ ) are in
Cambridge.

23. HAEMAPHYSALIS CALCARATA Neumann, 1902.

Figs. 373-377.

Lit. and Icon. : Neumann, 1902, pj). 113-115, Figs. 2, 3 {,$ reproduced).
Description of (^ and o, the latter not figured. Neumann, 1910, pp. 173-174,
Figs. 11, 12. Description of $ . The figures represent the $ capitulum and
scutum, ventral aspect of palp, digit ; not reproduced (see our text). Neumann,
1911a, pp. 113-114; account condensed from Neumann, 1902, $ not de-
scribed, species not figured. Blanchard, 1909, p. 148, Fig. 178 ; species
merely listed, the figure taken from Neumann.

Male (Figs. 373-375) : Body about 1-7 x 1-1 mm. in average speci-
mens\ Scutum : punctations very fine, discrete, inconspicuous ; cervical
grooves short, deep, converging ; lateral grooves medium, well-marked, in-
cluding one festoon ; festoons relatively broad, hardly longer than broad.

Fig. 373. Fig. 374.

Fig. 373. H. calcarata <? . Dorsum and venter, spiracle and tarsus IV. (Drawn from the

type in Neumann's collection, No. 1292.) Original, G. H. F. N. del.
Fig. 374. H. calcarata <? , left palp, ventral aspect. Neumann, 1902, Fig. 2.

Capitulum : base trapezoid, broadest in front, with straight or somewhat
rounded sides converging behind, cornua slight ; palps with article 2
very salient laterally, the salience hollowed to receive coxa I, and
produced on the ventral side to a sharp retrograde spur ; article 3
broader than long, with a long sharp ventral retrograde spur ; no

1 This is the size of the c? type we illustrate in Fig. 373 ; our 4 <? s from Somaliland
measured 1-95 x 1-1; l-8xl-l; 1*75 x 1-1 ; and 1-65x1 mm. respectively.

H. calcarata

443

dorsal spurs on the palps ; hypostome 4 | 4. Venter : spiracle sub-
triangular or somewhat comma-shaped, rather small. Legs : coxae
with strong internal spurs, strongest on coxa I; trochanter I with
long sharp ventral (as well as dorsal) retrograde spur ; a similar but
less-marked spur on trochanters II and III, nearly obsolete on
trochanter IV ; tarsi short and very stout almost to the tip, where
they taper very abruptly ; pad moderate.

Female (Fig. 376). Scutum : 0*95 x 0"85 mm.', appearing distinctly
longer than broad, narrowing posteriorly ; punctations rather few,
indistinct, of moderate size. Capitulum with base of similar shape to
that of cT though relatively broader ; porose areas oval, well separated,

Fig. 375.
Fig. 375. H. calcarata s

Fig. 370.
Coxae with trochanters. Neumann, 1902, Fig. 3.
Fig. 376. H. calcarata ? . Scutum, capitulum in dorsal and ventral aspects, coxae and
trochanters, spiracle and tarsus IV. (N. 1224.) Original, N. C. del.

converging anteriorly ; palj)S as in the ^^ but relatively longer and
with the ventral spur on article 2 less marked. Venter : spiracle sub-
circular, small. Legs as in the ^f , with the same coxal and trochantal
armature less marked, and with similar thick abruptly-tapering tarsi.

Nymph (Fig. 377) : Scutum sub-circular, shagi-eened but without
definite punctations ; cervical grooves well-marked and converging at
first, then broad and shallow. Capitulum with base broadening
anteriorly, cornua slight ; palps with article 2 sharply salient laterally,

Measured on our specimen from Somaliland.

444

Genus Haemaphysalis

with its posterior border almost straight dorsally, but with a ventral
retrograde spur directed outward ; lateral contour of articles 2 and 3
continuous and concave outwardly ; article 2 twice as long as article 3 ;
hypostome 2 j 2. Legs with a sharp internal spur on all the coxae.

Through the courtesy of Professor Neumann we have been able
to study and figure one of his type specimens {^ ), the species having
been founded by him on 5 </ and 1 o from Spermosciurus sp., Sichi Baz,
Abyssinia, collected by Hilgert (received in 1901 from Erlanger).

Fig. 377. H. calcarata Nymph. Capitulum and scutum, capitulum in ventral aspect,
with coxae and trochanters, spiracle and tarsus IV. (N. 1224.) Original, N. C. del.

Neumann, 1910, p. 173, records 2 </ and 8 $ from a squirrel,
Roseres, Upper Blue Nile, Egyptian Soudan, 1906, coll. C. Alluaud
(Paris Mus.). The female as described and figured by Neumann (1910)
does not agree so closely with the male as do our specimens ; we do not,
therefore, reproduce his figures.

Our description of the adults and nymph is based on specimens
(N. 1203, 1224), 4 cT 2 $ and 1 o, from Xerus rutilus, Biacobaba,
Somaliland, iv. 1901, presented by Dr E. Brumpt, and the study
of a type </.

Types in Neumann collection, Toulouse. Co-type (N. 2882) </
in Cambridge.

Haemaphysalis calcarata var. houyi
Nuttall and Warburton, 1915, n. var.
Male : Resembles the type but for its somewhat sturdier build and
the fact that the trochantal spurs are short and blunt, the palps being
somewhat shorter and more massive.

H. calcarata var. houyi

445

Female : Resembles the type except for the reduced trochantul
spurs.

Nymph : Possesses a scutum rather longer than in the type.
Trochantal spurs absent.

Described from 7 cT 1 ? '^^^^ 2 o from Bate, New Cameroon,
collected by Dr Houy. The variety is closely allied to a small form of
H. leachi found in Abyssinia by Dr Brumpt but it is distinguished
therefrom by its humped tarsi.

Types in the Berlin Museum (No. 279), co-types in Cambridge
(N. 2996).

24. HAEMAPHYSALIS CALVUS Nuttall and
Warburton, 1915 n. sp.

Figs. 378, 379.

Male (Fig. 378). Scutum about 2-4 x 1-7 mm.\ broadly oval,
widest at about the fourth coxae, glossy, with very numerous minute

Fig. 378. H. cahnts s . Dorsiim and venter, right i^alp in profile, hypostorae (highly
magnified), spiracle and tarsus IV. (N. 2120 type.) Original, N. C. del.

shallow punctations, often confluent and tending to a linear arrange-
ment ; cervical grooves short deep depressions close behind the anterior
border; lateral grooves short, beginning far back and ending very

1 The scutums of our 3 i specimens measure respectively :

2-6 X 1-8.5 mm. 2-3 x 1-7 mm.

2-45 X 1-8 „

N. I. 29

446

Genus Ilaemaphysalis

indefinitely ; festoons rather longer than broad. Capitidum : base
rectangular, twice as broad as long, finely punctate, with moderate
cornua ; palps short, article 2 only slightly salient laterally, the salience
about the middle of the article, considerably longer than article 3 in
its ventral aspect ; no dorsal spines, but long sharp ventral retrograde
spines on articles 2 and 3, those on article 2 so divergent as to be
visible dorsally ; hypostome short and broad, with 6 | 6 small sub-equal
teeth, about 8-10 per file, and large corona. Venter : anal grooves
ogival ; spiracle sub-rectangular, rather broader posteriorly. Legs :
strong sharp spines on all the coxae, longest on coxa I ; well-marked
sharp trochantal spurs, progressively diminishing, and shortest on
coxa IV ; tarsus IV long, tapering gradually ; pad long.

Female (Fig. 379). Scutum broader than long, 0*9 x I'l mm.,
broadest near the posterior end, glossy, with numerous moderate
punctations ; cervical grooves well-marked, sub-parallel. Capitulum :
base twice as broad as long, cornua slight, porose areas large, sub-
circular, far apart, but indefinite in contour except where defined

Fig. 379. H. calvus ? . Scutum, capitulum in dorsal and ventral aspects, right palp
in profile, coxae with trochanters, spiracle and tarsus IV. (N. 2120 type.) Original,
N. C. del.

by the lateral ridge ; palps of the same character as in the (/ , but
rather longer and entirely without the divergent ventral spurs on
article 2 ; hypostome short and broad, 4 [ 4. Venter : anal groove
semi-circular ; spiracle sub-circular. Legs as in the cT > with shorter
spurs.

H. calciiH

447

Described from 3 ^ and 1 % (N. 2120 a), taken from huffalo,
Sekong River, Sandakan, British North Borneo, 19. iv. 1918,
Dr H, F. Conyngham coll.

A large species allied to H. montgomeriji Nuttall, 1912, but
differing therefrom as follows : basis capituli and scutum broad, fourth
coxal spur less developed, venter not hairy, ventral spur of article 2
of (/ palp more divergent, etc.

Types in Cambridge.

25. HAEMAPHYSALIS SPINIGERA Neumann, 1897.

Figs. ;380-383.

Lit. and Icon. : Ncuuiaun, 1897, pp. 3.52-3.54, Fig. 18 (^J pii-lp, digit aud coxae,
herein reproduced) ; Neumann, 1901, p. 264 ; Neumann, 1911 a, p. 113, Figs. 59,
60 (condensed from his earlier description ; omits the figure of the digit) ;
Blanchard, 1909, p. 157, Fig. 195 (four-line description with figure ; both taken
from Neumann).

Male (Figs. 380, 381). Scutum oval, about 2*5 x 1-5 mm., broader
posteriorly, finely punctate all over ; cervical grooves shallow, parallel ;

Fig. 380. H. spinigera i . Capitulnm in dorsal and ventral aspects. Drawn from a
balsam-mounted type specimen taken from a bear. Only the surface structures are
shown. Original, G. H. F. N. del.

lateral grooves short, very concave internally, including no festoon ;
festoons long. Capitulum. : base rectangular, finely punctate dorsally,
cornua strong ; palps short, article 2 fairly salient laterally, with a
retrograde spine at the ventro-lateral angle ; article 3 much broader
than long, with a short stout ventral tooth; hypostome 5 | 5, 9 or 10
teeth per file. Venter: spiracle oval, with slight dorsal process. Legs :
relatively stout ; coxa I with a long mternal spur; short spurs at the
middle of the posterior border of coxae II and III, a very long internal
spur on coxa IV ; tarsi tapering, pad long.

29—2

448

Genus Haemaphysalis

Fig. 381. H. spinigera g : (1) left digit, x 320 ; (2) left palp in ventral aspect, x 65 ; (3)
coxae, x45. Drawings from the type. Neumann, 1897, Fig. 18.

Female (Figs. 382, 383). Scutum sub-circular, 1-2 x 1-1 mm., with
very fine punctations irregularly distributed ; cervical grooves faint.

o

Fig. 382. Fig. 383.

H. spinigera, ? .
Fig. 382. Capitulum and scutum, capitulum in ventral aspect, tarsus IV. Drawn from

one of our specimens (N. 1998). Original, N. C. del.
Fig. 383. Coxae of right side and spiracle. Drawn from a type specimen mounted in
balsam. The tick was found on an Indian bullock. Original, G. H.F. N. del.

H. spiiiigera 449

sub-parallel. Capitulum : base rectangular, much broader than long ;
porose areas oval, for apart, a small median pit in the interval ; cornua
fairly strong; palps relatively longer than in the c/, but with the same
characteristics; hypostome 4 1 4. Venter : spiracle sub-rectangular, dorsal
process almost absent. Legs : coxa I with a fairly strong internal spine ;
a very small spur on coxae II-IV ; tarsi tapering, pad long.

Our description is based on specimens (N. 1998, cf ? ) found on
Felis tigris, at Kadra, Kanara, India, xi. 1910, by N. B. Kinnear. and
on others (N. 2388 5 </ 2 ? 1 o) found on a panther and received in
1913 from R. Hancock. Neumann's types were (a) specimens from
S. Mahratta, India, mounted as microscopic preparations', namely, 2 cT
found on Felis tigris and a hear respectively, and 1 $ (now in
Toulouse) found on an Indian bullock, (/>) 2 </ 4 $ from Ceylon,
collected by Deschamps (Paris Mus.). Neumann has since (1901, p. 264)
recorded 3 $ from Judea, collected by Roux (Paris Mus.).

Haemaphysalis spinigera var. novae-guineae
(S. Hirst 1914) Nuttall and Warburton, 1915.

Figs. 384-386.

Lit., Synon. and Icon. :

Haemaphysalis novae-gxdneae Hirst, vi. 1914, pp. .328-330, Fig. 16 ; $ paljj in
ventral aspect and coxae with trochanters ; ^ capituhim and scutum (not
reproduced).

Both sexes are more glossy and less punctate than the type species.

Male (Fig. 384) differs from the type chiefly in the palps and
the coxal armature. Scutum : ranges in size from 2"45 x 1*6 down
to 1*66 X 1'05 mm. I Capitulum : the external contour of articles 2 and

1 These specimens were kindly lent us by Prof. Nenmann for purposes of study and
they were used in connection with our figs. 380 and 383. The specimens bear the labels
of the firm of E. Wheeler, London and formerly belonged to the Blanchard collection.

- The scutums of 12 g and 14 ? varied as follows in their measurements in mm. :

12 J

14?

2-45

xl-6

Type

from New Guinea.

11

xO-9

Type from New Guinea.

2-25 X 1-53

,,

It

))

0-95

xl-1

Specimens

from Queensland

2-2

xl-5

,,

,,

, ,

0-93

xl-0

1-93

xl-22

Specimens from

Queensland.

0-9

xl-0

1-9

xl-2

,,

,,

,,

0-9

xO-96

1-85

xl-2

,,

»>

,,

0-9

xO-9

1-81

xl-24

,,

>■>

,,

0-87

xO-9

1-8

xl-18

,,

,,

,,

0-87

xO-9

{Footnote continued overleaf)

450

Genus Haema/physalis

Fig. 384. H. spinigera var. novae-guineae i . Dorsum and venter, spiracle and tarsus IV.
• (N. 2673.) Original, N. C. del.

3 of the palps is almost unbroken ; the lateral salience of article 2 is
longer and more retrograde in direction ; there is a slight but distinct
dorsal spine in the middle of the posterior border of article 2. Legs :
the spurs on coxae I-III are very short and blunt ; whereas in the
type there is a long spur on coxa I and well-marked spurs, longer than
broad, on coxae II and III. The long needle-like spur on coxa IV is
present in both forms.

Female (Fig. 385). Scutum: ranges in size from 1"1 x 0*9 to
0'78 X 0'91 mm.^ Capitulum like that of the ^T, and presenting
similar differences from that of the type. Legs : spur on coxa I
slight.

Nymph (Fig. 386). Scutum about 0-52 x 0*54 mm.-, sub-circular,
with a few faint punctations and well-marked sub-parallel cervical
grooves. Capitulum with the characteristics of the % less pronounced ;

1-7 xl-14
1-7 xl-08
1-68 X 1-15
1-66 X 1-05

Specimens from Queensland.

0-85 X 0-95
0-85 X 0-92
0-84 X 0-95
0-8 xO-95
0-8 xO-9
0-78 x 0-91

Specimens from Queensland.

1 See note 2, p. 449.

'^ The scutums of 5 nymphs measured as follows in mm. :
0-54 X 0-54 0-48 x 0-53

0-52 X 0-54 0-44 x 0-53

0-51 X 0-55

H. spinigera var. novae-guinecie

451

Fig. 385. H. spinigera var. novae-guineae ? . Scutum, capitulum in dorsal and ventral
aspects, coxae with trochanters, spiracle and tarsus IV. (N. 2673,) Original,
N. C. del.

Fig. 386. H. spinigera var. novae-guineae, nymph. Dorsum and venter, spiracle and
tarsus IV. (N. 2673.) Original, N. C. del.

comua distinct ; hypostome 4 4. Spiracle circular. Legs as in
the $.

Described from ^ ^,1 % and os (N. 2673), from bandicoot, Manus,
Admiralty Islands, German New Guinea, 20. ix. 1913, A. F. Eichhom
coll. We have since determined (Sweet No. 463 = N. 2832) 9 ^ and
13 ? from kangaroo, Northern Territory, Queensland, Australia : and

452 Genus Haemaphy salts

(N. 3011 = Berlin Mus. No. 185) many cT ? o and L from Perameles
dorevanus, Astrolahe Bay, German New Guinea. After this description
was written and the figures completed, we were able to examine speci-
mens of H. novae-guineae Hirst (</ %) through the courtesy of the
author. We regard Hirst's species as but a variety of H. spinigera;
his description was based on numerous specimens taken from Kangaroo,
Perameles, and Hydromys sp., at Sattelberg, Huon Gulf, German New
Guinea, 1911, by Prof F. Forster.

Types (cT ? ) in the British Museum ; co-types (N. 2903, ^ % ,
from Hydromis), and our types (N. 2673) of the O are in Cambridge.

26. HAEMAPHYSALIS CONCINNA Koch, 1844.

Figs. 387-393.

Lit., Synon. and Icon. :

Haemaphy salis concinna C. L. Koch, 1844, pp. 237-238 ; 1847, p. 125, PI. XXVII,
Figs. 99, 100, (^ and $ , coloured, fairly good but for the capitulum of the ^.
Origin undetermined.

Haemaphy salis concinna C. L. Koch, in Neumann, 1897, pp. 338-341, Figs. 4-6
(reproduced). Neumann is wrong in giving Ixodes nigrolineatus Packard
and 7. cookei Packard in his synonymy of H. concinna (see Ticks, Part II,
pp. 280 et seq.) ; Salmon and Stiles, 1901, Figs. 219-221 (reproduced from
Neumann). Blanchard, 1909, p. 148, quotes Neumann.

Ixodes chelifer Megnin, 1880, p. 132 ; all the author says of this tick is that
it is distinct from Ixodes ricinus "the palp, of which the third article is
extended inward in a curved point, simulating, with its mate, a pair of
pincers." He collected the $ only. Canestrini, 1890, p. 526, and Berlese,
1891, fasc. LVIII, No. 10, wrongly give /. chelifer Megnin as a synonym of
H. punctata ( = cinnabarina var. punctata).

'■'■ Haemaphy salis hirudo L. Koch," in Neumann, 1897, p. 341 ; see Note, p. 456,
and our list of Condemned Species.

? Haemaphy salis concinna var. kochi Neumann, 1905, p. 239 ; see Note, p. 456.

Haemaphysalis concinna Koch, from ? Ovis aries, Brunswick, chosen as the type
of the genus Haemaphysalis by Neumann, 1901, p. 340.

Haemaphysalis concinna concinna Koch, a sub-species in Neumann, 1911 a,
p. 110, Fig. 53 (reproduced from Neumann, 1897, Fig. 4).

Haemaphysalis co;iem/ia^oc/ii Neumann, a sub-species in Neumann, 1911a, p. 111.
(See Note, p. 456.)

Haemaphysalis concinna Koch, in Bonnet, 1908, pp. 259-260, poor description,
condensed from Neumann, 1897 ; Fig. 28, of ? capitulum, is original, but
bad. Banks, 1908, p. 32, says the record of the presence of coiicinna in the
United States is " due to wrong synonymy " (refers to Neumann, 1897,
q.v. supra). Blanchard, 1909, pp. 148-150, Figs. 180-184, taken from
Neumann, 1897, 1901, 1905 ; Fig. 183, taken from Bonnet, 1908 ; descrip-

H. concinna

453

tion contains errors. Patton and Cragg, 191.3, p. G30, description taken
from Neumann, 1911a, p. 110; PI. LXXXI, Figs. 1, 2, original, of $ in
ventral and dorsal aspects. (For Haemaphysalis longicornis Neumann,
1901, Haemaphysalis concinna var. longicornis (Neumann) Neumann, 1905,
otherwi.se H. concinna concinna (Neumann) Neumann, 1911, refer to list
of doubtful and condemned species.)

Male (Figs. 387-390): Scutum oval, 2-3 x 1-9 to 3x1-84 mm. ^
glossy, with many fine but inconspicuous punctations ; lateral grooves

Fig. 387. Fig. 388.

Fig. 387. H. concinna s . Dorsum ; part of venter, tarsus IV and spiracle (more highly

magnified than dorsum). Specimen (N. 1075) from France. Original, G. H.F.N.

('el.
Fig. 388. H. concinna <? . Capitulum (A) ventral and (B) dorsal aspect, x 65. Neumann,

1897, Fig. 4. (Drawn evidently from balsam-mounted specimen.)

well-marked, including one festoon ; cervical gi-ooves short and faint ;
festoons long, the intervals well-marked. Capitulum : base with strong
sharp cornua ; palps longer than broad, article 2 moderately salient
laterally, article 3 elongate and curved inwards at the tip, giving the
palps when closed a chelate appearance ; a fairly strong retrogi-ade spine

1 The scutums of 20 cf captured on deer in France by Brumpt measured as follows
in mm. (One of the types in Berlin measures 2'7 x 1*8 mm.) :

3 X 1-84
2-95 X 1-85
2-9 xl-85
2-9 xl-8
2-85 X 1-84
2-82 X 1-77
2-8 xl-84
2-75 X 1-75
2-72 X 1-8
2-7 xl-8

2-7 xl-73
2-66 X 1-7
2-65 X 1-6
2-6 xl-8
2-6 xl-78
2-58x1-7
2-53x1-72
2-46x1-57
2-4 xl-54
2-3 X 1-9

454

Genus Haemaphysalis

i^ism

mi^mh

Fig. 389. Fig. 390.

Fig. 389. H. concinna <? , hypostome, x 210. Neumann, 1897, Fig. 6.

Fig. 390. H. concinna <? , left digit in dorsal aspect, x 230. Neumann, 1897, Fig. 5.

ventrally on article 3 ; hypostome markedly shorter than the palps,
corona large, dentition 6 | 6, teeth strong and uniform. Ventei" : spiracle
large, white, sub-oval, with slight dorsal process. Legs : coxa I with fairly
long pointed spur ; coxae II-IV with slight spurs ; tarsus IV tapering
rather rapidly ; pad medium.

Female (Fig. 391) : Scutum sub-circular, about 1'2 x 1-4 mm. (varies)^
glossy ; cervical grooves distinct for about two-thirds the length of the
scutum, concave externally, the interval rather broad ; punctations very
fine and inconspicuous. Capitulum : base broader than in the </ and
with less pointed cornua ; porose areas ill-defined, sub-circular, far
apart ; palps with article 2 rather prominent at postero-internal dorsal
margin ; article 3 much broader than in </ , the palps not being
chelate when closed ; no dorsal spines but a moderate blunt spur
ventrally on article 3 ; hypostome with dentition 5 1 5 or 6 j 6.
Venter: spiracle large, sub-circular. Legs: coxae as in (/ ; tarsus IV
longer, tapering gradually.

^ The type ? , seen and drawn by G. H. F. N., has the scutum distinctly longer than
broad ; but in Neumann's description it is given as circular, and it is very nearly so in a
specimen kindly given us by Neumann, which we figure ; the scutum may, however,
be slightly broader than long.

The scutums of 5 ? , captured on deer in France by Brumpt, measured as follows in mm. :
1-3 xl-33 1-2 xl-18

1-24 X 1-45 1-13 x 1-4

1-2 xl-45

H. concinna

455

Fig. 391. H. concinna ? , capitulum and scutum, ventral aspect of capitulum with
coxae, spiracle and tarsus IV. Same origin as the s in Fig. 387. G. H. F. N. del.

Nymph (Fig. 392) : Body (when gorged) may attain 3'4 x 23 mm.
ScutuDi nearly circular, about 0'5xO"6min.\ shagi'eened, with a few

Fig. 392. H. concinna nymph, capitulum in dorsal and ventral aspects, scutum, coxae
with trochanters, spiracle and tarsus IV. (N. 2763, specimen raised by E. Brumpt.)
Original, N. C. del.

scattered punctations ; cervical grooves well marked for two-thirds of
the scutal length. Capitulum : base with slight blunt cornua ; palps
with article 2 more salient laterally than in the $ ; hypostome 2 j 2.

' The scutums of 5 o, captured on deer in France by Brumpt, measured as follows in mm. :
0-54 X 0-6 0-5 X 0-6

0-5 xO-68 0-5 X 0-54

0-5 yO-64
The body length of 7 fully gorged nymphs, found on deer and hedgehog in France by
Brumpt, measured as follows in mm. :

3-4 x2-3 315 X 2-1

3-3 x21 30 x21

3-25 X 2-1 30 X 205

3-2 x2-0

456

Genus Haefmapliysalis

Venter: spiracle sub-circular. Legs: coxal' spurs distinct; tarsus IV
tapering gradually.

Larva (Fig. 393) : Body (unfed) 0*67 x 0"56 mm. Scutum broader
than long, about 0"28 x 0*34 mm.^ shagreened, but without puncta-
tions ; cervical grooves generally fairly distinct for half the scutal
length. Capitulmn : like that of the nymph, except that the comua
are almost obsolete and the distinction between articles 2 and 3 of the
palps obscure. Legs : a slight indication of a spur on coxa I ; coxae II
and III unarmed ; tarsus III tapering to rather sharp point.

Fig. 393. H. concinna larva, capitulum in dorsal and ventral aspects, scutum, coxae
with trochanters, spiracle and tarsus III. (N. 2763, specimen raised by E. Brumpt.)
Original, N. C. del.

Our description is based on numerous specimens from France and
an examination of the types.

Types (cT ? ) ii^ the Zoological Museum, Berlin ; origin unknown.
The types of the immature stages (N. 2763, o, L, figured) are in
Cambridge.

NOTE

Haemaphysalis concinna var. kochi Neumann, 1905.

We fail to recognize this variety as valid for the follovping reasons :
Neumann, 1897, p. 341, described the specimens concerned under the name
of " B. hirudo Koch," a species which he subsequently recognized as purely nominal
in view of the types not being available. (We include H. hirudo in our list of

1 The scutums of five larvae (N. 2778) measured as follows in mm. :
0-28 X 0-36 0-26 X 0-37

0-28 X 0-35 0-25 X 0-34

0-28 X 0-34

n. concinna 457

condemned species q.v.) Neumann, 1905, p. 239, renamed the specimens H.
concinna var. kochi. The specimens were (a) 6 $ from the Amur River, Eastern
Siberia, coll. Dickman, 28, vi. 1899 (Hamburg ]\Ius.) ; (i) 3 9 from Japan, coll.
Hilgendorf (Berlin ]\Ius.); (c) 2 ? from dog, Saga, Japan, coll. Yamaguchi; {d)-l $
from Saigon, Cochin China, coll. Harmand (Paris Mus.).

Through the courtesy of the Museum authorities in Hamburg, Berlin and Paris,
we have been able to study the types of (a), {b) and (c^), ten specimens having been
received from Hamburg', and one each from Berlin and Paris. The specimens
conform to H. concinna $ , but we do not find the suggested varietal characters at all
constant in these Hi)ecimens — and in themselves they are exceedingly slight. More-
over, it appears to us very unsafe to attribute the females to concinna at all in the
absence of males, more especially when the specimens are extra-Euroi)ean. The
$ of H. concinna has no very salient characteristics, and though the ticks in
question bear a strong general resemblance to our European specimens they are also
like other species, e.g. H. papuana. If the corresponding (^s should tvu-n out to
possess cheliform palps the attribution to H. concinna would be justified, but if
this be not the case we do not think that the similarity of the ? s would be
sufficient ground to maintain this form as a variety of that species.

Geographical Distribution.

There is no trustworthy evidence of H. concinna having been found
outside Europe, for no ^ has been recorded as accompanying the
supposed $s from other parts of the world. The tick occurs in
Germany ; there is a $ in the Berlin Museum, collected in Brunswick,
and a (/ of German origin. Neumann records 4 (/s from deer in
Poland. The bulk of the specimens collected comes from Prance ;
thus, in the north, Megnin (1880, p. 132) collected only (/s in the
Forest of Fontainebleau. We are indebted to Dr E. Brumpt, of Paris,
for the opportunity of examining many specimens of fed and unfed
adults, nymphs and larvae found by him on sixteen occasions on deer
at Fontainebleau (S. et M.), Chantilly and Compiegne (Dept. Oise),
and the Dept. Indre ; and twice (o and L) on hedgehogs at Chantilly ;
on nine occasions H. concinna was found with H. inermis on deer, and
once with H. cinnabarina var. punctata^. Dr Brumpt raised H. concinna
successfully upon the dog and hedgehog in the laboratory, thus giving
us an opportunity of publishing the first description of the immature
stages. He has kindly allowed us to retain some of his specimens
(N. 1529, 2754, 2755, 2760, 2761, 2778). Neumann (1897) who has
presented us with a (/ and ? (N. 1075), also records concinna from

1 We are much indebted to the Hamburg Museum for the gift of two of these specimens
(N. 1987).

2 See Table of Brumpt's observations under Biology, p. 544.

458 Genus Haemaphysalis

Southern France ; a </ from St Jean-de-Luz, near the Spanish border, a
% from Belesta, Ariege ; 33 c/, 22 % (Simon coll.) and 1 % from
sheep (R. Blanchard coll.) being from undetermined places.
Biology : see p. 542.

27. HAEMAPHYSALIS SIMPLEX Neumann, 1897.

Figs. 394-397.

Lit. and Icon. : Neumann, 1897, pp. 345, 346, Fig.s. 10-12 (drawn from balsam-
mounted specimens ; reproduced by us) ; Donitz, 1907, p. 70 (quotes Neu-
mann) ; Blanchard, 1909, p. 157 (species listed) ; Neumann, 1911 «, p. Ill,
Figs. 55, 56 (taken from author's earlier paper).

Male (Figs. 394-397) : Scutum cinnamon yellow, elongate,
1"8 X I'O mm., ovate, very convex, glossy, with very minute shallow
punctations, distributed all over, also on the festoons ; cervical grooves
shallow, sub-parallel ; lateral grooves practically absent ; festoons,
especially the anterior festoon, broad, involving an unusual proportion
of the body periphery. Capitulum small ; base with very slight
bluntly rounded cornua ; palps with article 2 strongly salient laterally ;
articles 2 and 3 of equal length, without dorsal spurs ; a small sharp
ventral spur under article 3 ; hypostome spatulate, dentition 4 | 4,
about 8 files of uniform fairly strong teeth. Venter : spiracle large,
w^ith a slight dorsal process ; a large proportion (nearly half of the
body) lies posterior to coxa IV. Legs short and thick; coxae I very
small, with slight blunt spur; coxae III and IV very broad (antero-
posteriorly), with short blunt spurs ^; the dorsal spur on trochanter I
almost obsolete ; the other trochanters unarmed ; tarsi short, thick,
humped, without ventral spur; pad almost as long as the claws.

Female : unknown.

Nymph : body 05 mm. long, corresponding in form and colour
to the (/. Scutum rounded, extending almost to half the body-length.
Capitulum : hypostome 3 j 3, with 6 or 7 stout teeth per file. Palps
and Legs as in the (/ .

Our description of the cT is based on a co-type kindly presented
by Prof. Neumann from his type material which originally consisted

1 Neumann's figure, here reproduced as Fig. 396, does not agree with his text, which
reads "Coxae contiguous, without spurs; a simple, almost obsolete tuberosity at the
postero-internal angle."

H. simplex

459

Fig. 394. H. simplex <? , dorsum, venter, left palp in profile (;; = ventral spur), spiracle
and tarsus IV. Drawn from British Museum specimen. Original, G. H. F. N. del.

Fig. 395. Fig. 396. Fig. 397.

Fig. 895. //. simplex s, (A) left digit, x 425 ; {B) left palp in ventral aspect, x 110.

(The spur on palpal article 2 is misplaced.) Neumann, 1897, Fig. 10.
Fig. 396. H. simplex i , coxae and trochanters, x 45. Neumann, 1897, Fig. 11.
Fig. 397. H. simplex <? , tarsus IV, x 60. Neumann, 1897, Fig. 12.

460 Genus Haemaphysalis

of 13 (/ and 1 o found on Erinaceus sp. in Madagascar, 1894, by Sikora,
together with H. elongata ? . The description of the o is taken from
Neumann. We reproduce the author's figures of parts of the (/.
Our figure is dra\^^l from a cT found on Ericidus setosus in Madagascar
(Cohen coll., 1862, Brit. Mus. ; determined by Neumann in 1905).
Types in Toulouse, co-type J' in Cambridge (N. 2891).

28. HAEMAPHYSALIS LEACHI (Audouin 1827)
Neumann, 1897.

Plates XII and XIII. Text-Figs. 398-410.

Lit., Syn, and Icon. : Ixodes leachii Aiidouiu in Savigny, 1826, PI. IX, Fig. 9 ;
an excellent figure of ^ dorsum, that is considering the date when it was
drawn. Audouin, 1827, p. 428.

Rhipistoma leachi Koch, 1844, p. 239.

Rhipistoma elUpticum Koch, 1844, p. 239.

Rhipicephalus ellipticus Koch, 1847, p. 135, PI. LXX, Fig. 111.

Rhipidostoma leachii (Audouin) in Karsch, 1878, p. 337.

Opisthodon canestrinii Supino, 1897 a, 1897 6, p. 252, PL XIII, Fig. 21. The
author only describes and figures the ^ tarsus. Neumann, 1902, p. 128,
states he examined the types and concluded they were $ leachi. Neumann,
1911a, p. 115, nevertheless lists canestrinii as a doubtful species but
probably leachi. We have examined the types and can state positively
that canestrinii=leachi. The types of H. canestrinii are in the Genoa
Museum, co-types in Cambridge.

Opisthodon gestroi Supino, 1897 6, p. 252, PI. XIII, Fig. 23. The author only
describes and figures the $ tarsus. Neumann, 1902, p. 128, examined the
types in Genoa and stated they were allied to H. leachi. Neumann, 1911 a,
p. 116, gives it as a doubtful species of Haemaphysalis. We have examined
the types ; there is no doubt about their being = ^. leachi.

Opisthodon asiaticus Supino, 1897 a ; 1897 h, p. 252, PI. XIII, Fig. 22 ; the
author only figures the (t, tarsus, his description is useless. We have
examined the types at Genoa and regard them as = 5". leachi. Neumann,
1897, p. 357 ; renamed Haemaphysalis asiatica (Supino). Neumann, 1902 a,
p. 123 ; types stated to be lost.

Haemaphysalis leachi (Audouin) Neumann, 1897, pp. 347-350, Figs. 13-15
(reproduced by us) ; 1901, p. 263, refers to distribution and hosts. Froggatt,
1900, p. I 542 (cited by Rainbow, 1906, p. 165). Lounsbury, xi. 1901,
pp. 1-12, first experiments on the transmission of canine piroplasmosis
by this tick.— 1902. Lounsbury, pp. 4-9, PI. I, Figs. 1-11 ; second
series of experiments on tick-transmission of canine piroplasmosis ; im-
portant ; the figs, illustrate all stages. Lounsbury, pp. 5-7 ; biology. —
1904. Lounsbury, pp. 27-29 ; biology. Lounsbury, pp. 22-43 ; experi-

H. leachi 461

mental tick-transmission of canine piroplasmosis ; important. Nuttall,
pp. 226 et seq., Pis. XII, XIII (photomicrographs of all .stages) ; treats
of biology of tick and relation to canine piroplasmosis. — 1906. Buy,
pp. 140-142 ; specific description taken from Neumann. Rainbow, p. 165 ;
species listed for Australia on Neumann's authority. — 1907. Dcinitz,
pp. 68, 69, PI. Ill, Fig. 20 a, PI. V, Fig. 32 ((^ in dorsal aspect, ? capitulum
and scutum) ; we cite this author in our text. Newstead, Button, and
Todd, p. 99 ; listed from Congo Free State ; cited in our text. Neumann
(a) p. 24 ; species listed from E. Africa (see our text).— 1908. C. W. Howard,
pp. 160-162, PI. XVI, Figs, a-m ; gives a good description of all the stages
together with figures of the capitulums (in dorsal and ventral aspects) of the
S ? and o, digits of the (J $ and o, (J ? scutums, ^ coxae (the latter
and the ^ capitulum being taken from Neumann, 1897, Figs. 14, 15, whose
synonymy is followed) ; refer to our text regarding the author's records
as to hosts and distribution. Meuleman, repr. pp. 12 et seq. ; nothing
original. Nuttall (VIII), p. 394 ; unfed adults survive 7 months. Nuttall
(IX) pp. 514, 522, 523 ; biology of tick, a carrier of canine piroplasmosis. —
1909, Blanchard, pp. 152, 153, Figs. 188, 191 (taken from Neumann) ;
contains two erroneous statements : that this tick conveys canine piro-
plasmosis in Italy (the tick does not occur in Italy) and that it is the
suspected carrier of " Babesia parva " in Japan (there is no evidence what-
ever for this statement, and East Coast Fever, caused by the parasite
named, does not occur in Japan). Manteufel, p. 16 ; nothing original ;
C. W. Howard, p. 162 ; short statement regarding hosts and distribution. —
1911, Lounsbury, pp. 5, 6 ; biology. Nuttall (X), p. 180 ; type of para-
sitism shown graphically. Ziemann, p. 58 ; occurrence in Cameroon. —
1913. Nuttall (IV), pp. 93-99 ; biology. Nuttall (X), pp. 309, 310,
Figs. 7-10, also pp. 308, 309, Figs. 5-8 (published in two journals, figures
herein reproduced). Patton and Cragg, PI. LXXIII, Fig. 12, ? capitulum
in ventral aspect ; PI. LXXXI, Figs. 3, 4, ^ venter, $ dorsum, sketchy ;
p. 647, cite Nuttall and Lounsbury.— 1915. Nuttall, p. 252, Fig. 3, mal-
formation in a (J.

Haemaphysalis canestrinii (Supino) Neumann, 1897, ^
p. 357

Haemaphysalis asiatica (Supino) Neumann, 1897, i See further under Opis-
p. 357 ... ' thodon fihovQ.

Haemaphysalis gestroi (Supino) Neumann, 1897,
pp. 357, 358

Haemaphysalis leachi var. australis Neumann, 1905, p. 238 ; 1911, p. 115.

Haemaphysalis koningshergeri Warburton and Nuttall, 1909, p. 65, Figs. 11, 12
(reproduced).

Haemaphysalis leachi var. indica Warbiu'ton, 1910, p. 402 (not figured).

Haemaphysalis leachi australis Neumann in Neumann, 1911 a, p. 115 ; the author
here raises his variety to a sub-species.

Haemaphysalis leachi leachi (Audouin) in Neumann, 1911a, p. 114, Figs. 62, 63 ;
condensed description with figures as in Neumann, 1897.

30

462

Genus Haemaphj/salis

Male (Text-figs. 398-402 ; PI. XII, Figs. 4, 5 ; PI. XIII, Fig. 2).
Very variable in size (see Fig. 399). Scutum, in average specimens

<5*^

Fig. 398.

Fis. 399.

Fig. 398. IL Icachi i , dorsum, capituhim in ventral aspect, spiracle and tarsus IV.

(N. 532 c, G. H. F. N. del.) Nuttall x. 1913 a, Fig. 7.
Fig. 391). //. leacld i s, contours of two e s collected from one host and showing the

variation in size. (N. 310, G. H. F. N.) Original.

about 26 X 1'3 \r\\\\}, long and narrow, widest at the level of the
spiracles, convex, with very many minute punctations ; cervical grooves
fairly well marked, the interval between them narrow ; lateral grooves

^ The scutums of 25 i from various sources measured in mm. :

N. 2264 from Sierra Leone

N. 310 from Congo Free State

3-4 xl-8]
305 X 1-7 V
2-9 xI-gJ

Three c? picked out of a lot of 17 typical
specimens, the largest, smallest, and an
average specimen being selected.

2-45 xl •2") >

All comprised in one lot of
typical specimens.

(Berlin 22 h) from G. E. Africa

N. 496 c from Java

N. 954 (I from Borneo

N. 1060 c from Paining

Ind. Mus. 5994/10 from Bengal

N. 1225 Abyssinia

\

Typical Specimens.

2-4 xl-3"!
2-4 xl-5h

2-45 X 1-5 j
1-5 xO-8
1-4 x0-8i
l-35xO-9J

Three specimens of 11. koningsbergeri type.

//. leachi var. indica type.
A small form.

H. leachi

463

Fig. 400. Fig. 401.

Fig. 400. H. leachi <? , capitulum {A) in ventral, (U) in dorsal aspect, x 55. Neumann,

1897, Fig. 14.
Fig. 401. H. leachi i , coxae I-IV, x 30. Neumann, 1897, Fig. 15.

Fig. 402. Fig. 403.

Fig. 402. H. leachi, ? and i digits of right side in ventral aspect, x 2fi0. Neumann,

1897, Fig. 13.
Fig. 403. //. leachi ? , dorsum, ventral aspect of capitulum with coxae, spiracle and

tarsus IV. (N. 532 c G. H. F. N. del.) Nuttall, x. 1913 a, Fig. 8.

long, near the lateral border, including one or two festoons ; the latter
as broad as long. Capitulum remarkable for the very obtuse angle
at which the palps meet anteriorly ; base broadest in front, the lateral
borders nearly straight and converging posteriorly; cornua strong;

30—2

464 Genus Haemaphysalis

palps broadly conical, article 2 very salient laterally, with a dorsal and
a ventral triangular retrograde spur from each lateral salience ; article 3
very small, with no dorsal spine but a strong ventral retrograde spur ;
the lateral contours of articles 2 and 3 continuous and typically
rectilinear, forming a very obtuse angle in front ; hypostome 5 | 5 or
4 I 4, stout teeth, 10-12 per file. Venter: anal grooves ogival ; spiracle
oval with slight, blunt, dorsal process. Legs : coxa I bluntly pointed
posteriorly ; dorsal spur on trochanter I strong and pointed ; coxae
II-IV with a slight spur at the internal angle ; all the coxae broad
antero-posteriorly ; tarsi rather stout with small, terminal spur ; pad
fairly long.

Female (Text-figs. 403, 402 ; PI. XII. Figs. 1, 4 ; PL XIII, Fig. 1).
Scutum long-oval, somewhat narrowing behind, about 1"2 x 0'8 mm. in
average specimens \ minutely punctate all over ; cervical grooves rather
long and near together. Capitulum : base rectangular, broader than
in the cT , cornua strong ; porose areas oval, far apart ; palps resembling
those of cf but relatively longer and meeting at a less obtuse angle ;
hypostome 5 | 5, at times 6 [ 6, rarely 4 1 4. Venter : spiracle sub-
circular, somewhat narrowing dorsally. Legs : coxae very slightly
armed, bearing only minute spurs situate as in the (/ ; tarsi rather
stout, tapering gradually; pad short. When gorged may attain
12 X 9 mm.'-

Nymph (Text-fig. 404 ; PI. XII, Figs. 3, 4) : Bodi/, unfed, about

1 The following measurements, in mm., relate to ? s from various sources :

Scutums

Bodies (unfed)

N.

2264 from Sierra Leone (10 S )

1-5

xl-2

N.

2264

(69)

3-35 x 2-1

1-5

xl-15

3-3 x20

1-5

xl-05

3-2 x2-05

1-5

xl-05

3-2 x2-0

1-45

xl-1

3-2 xl-95

1-45

xl-1

3-0 x2-l

1-4

xl-2

1-4

xl-0

1-35

xl-1

1-25

xl-1

N.

310 from Congo Free State

1-35

1-3

1-15

xO-9

xO-95

xO-75

N.

1223 from Abyssinia

0-8

xO-7

N.

1223

1-8 xl-2

N.

, 496 c from Java

1-2

xO-9

(H

. ki

yni

ngshergeri type).

(Ind. Mus. 5994/10) from Bengal

0-8

xO-6

[H.

leaclh

i var. indica type).

- The largest of about 80 ? raised in Cambridge (N. 1737).

H. leachi

465

l''ig. 401. //. Ic.acld nymph, dorsum, venter, tarsus IV, hypostome [highly magnified].
(N. 1048. G. H. F. N. del.) Nnttall x. 1913 a, Fig. 9.

1"5 X 1 mm. ' ; may attain 8'4 x 2-0 mm. when replete^ Scutum penta-
gonal, with rounded angles, about as wide as long (0-5 x 0-5 mm.) ;
cervical grooves short and straight. Capituluru like that of the $
with the characters less pronounced, the cornua being very small and
the palpal processes slight ; hypostome 2 1 2, 6-7 teeth per file.
Venter : spiracle small, transv(;r.so, bluntly comma-shaped. Legs as
in the ?.

Larva (Text-figs. 405-407; PI. XII, Fig. 2): Body, unfed,

Fig. 405. FiR- 400.

Figs. 40.J, 406. H. leachi larva. Capitulum in dorsal and ventral aspects, highly

magnified. Scutum, drawn to the same scale as Fig. 407. All figures of the larva
drawn from one specimen. (G. H. F. N. del.) Original.

' Tlie largest of a lot experimentally raised.

- Tlic following measurements in mm., relate to os raised in Cambridge :

Scutums Bodies (unfed)

N. 1737 from ii. E. Africa (.5 o) 0-5 x 0-51 14 x 0-82

0-5 xO'5 1-4 X 0-8.5

0-46x0-4(; 1-2 xO-8

0-40 X 0-5 1-2 xO-77

0-4:{x0-44 118 X 0-7

466

Genus Haemaphysalis

Fig. 407. H. leachi larva. Ventral aspect. Specimen raised in Cambridge. Nuttall,
X. 1913 a, Fig. 10.

05 X 0-4 mm.S attaining, when gorged, about 1'5 x 0*9 mm. Scutum
broader than long, with rounded posterior angle and rather sinuous
postero-lateral borders ; cervical grooves faint, converging behind.
Capitulum like that of the o. Legs : coxae unarmed ; tarsi tapering
to a point.

It is unusual to find so close a resemblance between the capitula
of the immature and adult stages as is presented by H. leachi, and
the nymphs and larvae can generally be identified by this character
alone. On the other hand the adult tick is subject to considerable
variation both in size and facies.

Our description is mainly based on numerous specimens from
Africa some of which have been raised in the laboratory in Cambridge.

See further in the Section on Biology, p. 536.

1 Measurements of larvae in mm. :

Scutums

Bodies (unfed)

N. 1049 0-28 X 0-32

0-54 X 0-48

0-24 X 0-32

0-49 X 0-39

0-24 X 0-32

0-23 X 0-31

0-23 X 0-29

H. leachi 467

Varieties of Haemaphysalis leachi.

Two varieties of H. leachi have been established : var. cmstralis
Neumann, 1905, and var. indica Warburton, 1910.

Neumann regarded H. leachi as an essentially African species, the
dentition of the hypostome in the cf being 5 | 5 and in the $ either
4 j 4 or 0 I 5. When, therefore, he received </* examples from
Australia and Sumatra with dentition 4 1 4 he considered this
difference, in conjunction with th(^ habitat, as of varietal importance,
hence his var. avstralis. Donitz (1907, p. 68) has however pointed
out that African leachi, both </ and $ , vary in respect to their
dentition which in one and the same lot may range from 4 | 4 to 5 | 5.
We have observed the dentition in African leachi to vary between 4 | 4
and 6 j 6 as the result of examining a large material, and it is clear
from the many specimens that have reached us from abroad that the
species is much more widely distributed than was at first believed. It
seems, therefore, impossible to maintain Neumann's variety australis.

Warburton (1910) also underrated the range of variation to which
leachi is subject, not only in dentition, but also in other characteristics.
In a typical leachi the palps meet at a very obtuse angle, their lateral
contour is nearly a straight line, and article 2 has dorsal and ventral
retrograde processes on the lateral salience ; moreover, the scutum in
both sexes is long and narrow. Warburton described a variety indica
(Fig. 408 ; from India) with dentition 4 j 4, other characteristics much
softened and the dorsal retrograde process absent ; it appeared necessary
to establish the variety in view of the difference in facies. True
examples of leachi (African type) have more recently been received
from India, and though there are indications that the species is
undergoing some change in that country, tending to have shorter
and broader scuta and less obtuse and spinous palps, intermediate
forms occur and a clearly marked variety is doubtfully present.

Similar considerations have led us to degrade H. koningsbergeri
(Figs. 409, 410) to a s}Tionym of H. leachi.

There remain three established species, H. spinnlosa, H. numidiana,
and H. obtusa, which we think it best to retain for the present, but
which may eventually have to be merged in H. leachi. They are very
closely allied to H. leachi and would appear to be recently derived
therefrom, but a wider knowledge of them is necessary to determine
how far their peculiarities are constant. It is interesting to note that
they correspond in size to the two extremes of the range of authentic

468

Genus Haema2)hysalis

Fig. 408 H. leachi ? (var. indica, Warburton). Capitulum (0-8 mm. long) in dorsal
and ventral aspects, scutum, palp in profile, coxae, spiracle and tarsus IV. Specimen
(N. 942 b) from Madras. Original, G. H. F. N. del.

Fig. 409. Fig. 410.

Fig. 409. H. leachi (koningshergeri) d , dorsum, palp and coxae, tarsus IV. Sketch

C. W. del. Warburton and Nuttall, 1909, Fig. 11. Drawn from the type.
Fig. 410. H. leachi {koninqshergeri) ? , capitulum and scutum, tarsus IV, spiracle.

Sketch C. W. del. Warburton and Nuttall, 1909, Fig. 12. Drawn from the type.

H. leachi, H. nuniidiana being a large species, while H. ubtusa is very
small ; see their descriptions on pp. 478, 477.

Geographical Distribution and Hosts.

The species is widely distributed, occurring on a variety of hosts,
mainly in Africa,, but it is also encountered in Asia and Australia.
The types, now lost, came from Egypt (Savigny, 1820). The following

H. leachi 469

records relate to specimens (usually </ $ ) we have examined or
determined [records by other authors are enclosed in square
brackets] :

AFRICA: Egypt and Soudan: (N. 539) (^ from Erinaceus albi-
ventris, Gebel Auli, White Nile, Egyptian Soudan, collected v. 1900 by
Messrs S. and T. Witherby ; (from information received from Mr H. H.
. King, the species occurs on hedgehog and dog at Taufikia, Upper White
Nile Province, at Khartoum, and at Roseires, Blue Nile Province) ; the
species was first described from Egypt by Audouin (1827, p. 428).
Abyssinia : (N. 2318 a) from dog, Dir6 Daoua, Ethiopia, collected
28. V. 1913 by M. Pellerin ; (N. 2819, received from Berlin Mus.)
cT $ from Herpestes sp., 5. iii. 1913, and (Berlin Mus. 676/1913) from
hwre, 7. ill. 1913, both collected in Erithrea by Dr Klatt ; (Brumpt coll.)
cfs $ s from Canis mesomelas, Hyaena crocuta, Pvocavia brucei, (N. 1223)
P. abyssinica, all collected v. 1901 at Harrar by Dr E. Brumpt (the last
two lots were small specimens). [Pavesi, 1884, p. 92, states that
Antinori found the tick on Felis pardus in the Mahal-Uonz Mts.,
Eastern Ethiopia.] British East Africa: (E. 599) $ from dog,
Gwasinyras Post, Northern Frontier, collected in 1912 by J. O. W. Hope ;
(E. 258) (/ ? from Cams adastus. Northern slopes of Mt Kenia
(7700 feet elevation); (E. 266, 267 = N. 3065) ? os from Tachyoryctes
audax Thos. (mole rat), near Embu, East of Mt Kenia, 2. ii. 1911,
collected by S. A. Neave. Several lots were received from Nairobi :
(N. 102 c) cf from Dr P. H. Ross; (E. 291 = N. 1422) ^ $ from dog.
collected iv. 1911 by T. J. Anderson; (N. 1737) ? and L from jackal,
collected iii. 1912 by R. E. Montgomery. (N. 48 h) from dog, collected
XI. 1904 by Dr P. H. Ross at Makindu ; (E. 711) from dogs, Massai
Preserve, 1913, and (E. 712) from cheetah, Mara River, Boma, 30. vi.
1913, collected by Captain A. O. Luckman ; (E. 184) ^ $ from terrier
dog, Usoga, viii. 1910, collected by Dr Hailstone ; (N. 2403 c) cT from
lion, near Kadjiado River, Magady Railway, viii. 1913, collected by
Dr L. Nicholls ; (E.) rf from dog, Meko, and (E.) $ from sheep, Ibesha,
collected, ii. 1910 by J. 3. Simpson; (E. 294 h) % from cattle, Kokolo,
IV. 1911, collected by C. (1 Gowdey ; (E. 587) cT collected at Mombassa,
I. 1912, by R. P. Thomas ; (E. 839, 844 = N. 2836, E. 847, 849 a) cTs $ s
from dogs, Mau, Narok, Massai Reserve, 15-20, i. 1914, and (E. 781 /;)
from Olorogoti, xii. 1913, collected by Captain A. O. Luckman.
(Probably from British East Africa are: (N. 2377 a) os from
rhinoceros, 1912, and (N. 2388 c) c/s $s from panther, no date,
received 1913 from R. Hancock; also (Armstrong coll.) from cat,

470 Genus Haemaphysalis

I. 1911, at Aburi.) Uganda: We have determined two lots for the
London School of Tropical Medicine, collected from dog and ferret
by Dr H. Bayon (no further data) ; (N. 885 h) o from goat, (N. 876
and 695) from dog, Mpumu, Chagwe, collected in 1909 by Sir David
Bruce ; (E. 300) from dog, Lango, vi., coll. Dr McConnell ; (E. 480 h)
$ from grass, Upper Kafu River, Unzoro, xii. 1911, coll. S. A. Neave ;
(E. 692 = N. 3153) from lion, Busuju, 12. xii. 1912, coll. C. C. Govvdey;
(N. 873 c) from Buyago District, South Buddu, Lake Shore, viii.
1909, (N. 775 c) from cattle, Masaka, in Buddu, vi. 1909, coll. Sir
David Bruce; (E. 472 a.) from grass, same locality, IX. 1911, coll.
S. A. Nea^ve; (N. 874 6) from Kumi, Bukedi, viii. 1909, coll.
E. G. Morris; (N. 682) from dog, Bussu-Bussoga, 1909, coll. Dr H.
Bayon; (N. 532c) from Entebbe, 1909, coll. R. P. Riches; (E. 289a
and 556) from dog, Entebbe, ix. 1910, coll. C. C. Gowdey ; (Liverpool
191) from lion, Kamsala, 21. xii. 1912, coll. Capt. G. J.Keane ; (N. 507)
from Sesse Islands, Victoria Nyanza, 1908, coll. Dr H. Bayon ; we have
recently identified 8 lots presented by Dr Bayon to the Genoa Museum
in 1909, the specimens having been found on cattle and Bos indicus
at Bujata, Sesse Islands, and on unrecorded hosts at Entebbe, Kyetume
and Bussu-Bussoga. German East Africa: (E. 164 (i) from cattle,
Ruhaha River, Uhehe District, xi. 1910, coll. S. A. Neave; (N. 2581 b)
from jackals, Kilimabindi, Ngogo, 14. vii. 1913, coll. Dr W. Bartels,
received from Dr Kudicke of Daressalam ; (N. 575) from Hyaena
crocuta, Darie, coll. Erlanger-Hilgert Expedition, presented by Hon.
N. C. Rothschild (we cannot trace Darie on the map and assume it
lies in this country). We have identified several lots of adults for
the Berlin Museum (their Nos. 22 b, 48, o4, 6o, 1417/08 = N. 2795) as
follows : from Genetta sp., Maliwe, 6. vi. 1908, and from spotted Hyaena,
Rukwasteppe, 31. viii. 1908, both collected by Capt. Fromm ; from
Anomaluriis orientalis (a squirrel), Amani, x. 1907, Prof Vosseler coll.;
R. Regner collected specimens at Daressalam, Tangani and in the
Hinterland; two lots (Nos. 221, 222) labelled Ssongea, 23. ii. 1905 and
Longra, 17. ii. 1909, are probably from this region. [Neumann, 1907 a,
p. 24, records cTs $s from Genetta suahelica Mtsch., and cfs from
Herpestes caffer (Sjostedt Exped.); os from Felis leo L., Massimani
and $ from leopard, Donje-Erok (Schillings coll.)] Portuguese
East Africa: (N. 2425) from dog, Magude, 11. viii. 1913, coll.
Dr J. B. Botelho; (E. 618 a) from Gala, Zambesi River, coll.
Dr H. Swale, received 1912; [Karsch records the tick from Mozam-
bique, according to Neumann]. Rhodesia : (North) : (N. 2169 b) from

H, leaehi 471

dog, Broken Hill, ii.-m. 1913, coll. E. A. Copeman ; (N. 1971) from dog,
Serenje, coll. Dr A. Brown; from N.W. Rhodesia: (N. 754 H. leaehi
var. indica) from Helogale varia Thos., Msofu River, Alala Plateau,
7. XI. 1905, coll. S. A. Neave ; from Southern Rhodesia: (N. 577) from
Itonyx capensis, Bulawayo, I v. 1907, coll. E. C. Chubb, presented by
Hon. N. C. Rothschild; from North Eastern Rhodesia: (E, 187) from
o?o^, Fort Jameson, vi. 1910, coll. S. A. Neave; (E. 178 a) from man,
i.e. the collector's person and from natives, Upper Luangwa Valley,
VIII. 1910, coll. S. A. Neave; (Liverpool 185) from lion, Mpamadzi
River, ix. 1911, coll, Dr A. Kinghorn. Nyasaland : collected by
Dr J. B. Davey at Karonga were (N. 1617) from terrier dog, iv. 1909 ;
(N. 742) OS from civet cat, i. 1909; (Liverpool 72 a = N. 728a) from
leopard, Kambwi near Karonga, iv. 1909 ; (Liverpool 71 = N. 738)
from native dog, Nyika Plateau, ii. 1909; (E. 135 h) from dog, Valley
of S. Rukuru River, vi. 1910. Collected by S. A. Neave were (E. 130)
from dog, N.W. shore of Lake Nyasa, vii. 1910. The following were
collected in unrecorded places in North Nyasaland : (E.) coll.
Dr J. B. Davey ; (Liverpool 107) from terrier dog, iii.-iv. 1909, coll.
J. B. Davey; (E. 689) from dog, 9. viii. 1909, coll. J. G. Morgan.
(N. 749) from native dog, Chirua River, Central Angoniland, v. 1909,
coll. E. H. A. Pask ; (E. 135) from dog, Bua River, Central Angoniland, vi.
1910, coll. S. A. Neave; (N. 2366) from Hyrax sp.. Monkey Bay,
23. V. 1913, coll. Dr W. C. Wigan ; (N. 2244) c/s $s o from dog,
Mpondas, Fort Johnston, i. 1913, coll. Dr W. C. Wigan. The following
were collected at Zomba : (E. 250 = N. 1327) from grass, ii. 1911, coll.
A. M. D. TurnbuU; (E. 76) from dog, ill. 1910; (E. 248, 249 a) from
domestic cat, x. and xii. 1910, coll. A. M. D. Turnbull ; (E. 196, 219,
220) from domestic cat, civet cat, and collector's bed, ii. 1911, xi. and
XII. 1910 respectively, coll. Dr H. S. Stannus ; (E. 78 a) from dog,
on Mlanje Road, v. 1910; (E. 252) from cattle, x. 1910, coll.
A. M. D. Turnbull. Dr Stannus also found specimens on a lion, near
Zomba, ii. 1910. Dr J. G. Morgan collected (E. 690 a, 619 a) from
dogs, at Sichenya River, Mlanje, xii. 1912 and i. 1913. Received
from Blantyre were: (E. 70) from dog, iv. 1910 ; two lots collected by
Dr J. E. S. Old, namely (E. 81 c) found on the ground, iv. 1910, and
(E. 214 a) from an unrecorded host, vi. 1910. Collected by S. A. Neave,
on the Mwanza River, Shire Valley, were (E. 717 a, 729, 732) from
dogs, IV. and vii. 1913 ; (E. 731) from dogs, Lower Ruo Valley, vii.
1913. From Chiromo : (E. 67 h) from dog, iv. 1910; from the Lower
Shire Valley (E. 65, 71 a) from dogs, iv. 1910; one lot was collected at

472 Grenus Haemapliysalis

Port Herald by Dr J. E. S. Old : (E. 727) from lion, 12. vi. 1913. The
remaining specimens which we have examined from Nyasaland are from
places we could not trace on the map or from places unrecorded by the
collectors, namely : (E. 5 c) from terrier dog, after journey from Florence
Bay (W. shore of Lake Nyasa) to Cheranya, Akamanga Country,
V. 1909; (E. 14cS) from dog, xi. 1910, coll. Dr J. E. S. Old; (E. 216)
from cat, i. 1911, coll. Dr H. S. Stannus ; (N. 727 = Liverpool 57) from
native dog, Chizizi, near Howe River, ii. 1909, coll. Dr J. B. Davey.
Since the above was written we have received (E. 851 h, 852 h, 853 h,
854 = N. 2841) from dogs., S.W. shore of Lake Chilwa, i. 1914, coll.
S. A. Neave.

Transvaal: [Neumann, 1901, p. 263, records the tick from the
Transvaal, without mentioning either hosts or localities.] C. W.
Howard, 1908, pp. 162, 166-168, states that he has found the
tick in nearly every part of the Transvaal, upon almost all carnivora,
where it is rarely found on cattle and other animals " even when
most abundant " ; he lists it as occurring on cat, Felis pardus, F. leo,
F. nigripes, Viverra civetta. Genetta sp., Cynictis penicillata, jackal
and Erinaceas sp., this author also records the tick as occurring in
Orange River Colony. Natal: [C. W. Howard, 1910, p. 162, found
OS on Avicanthus pumillis (field rat) and on tortoise at Pretoria ; Galli-
Valerio, 1909, p. 539, records the tick from man, Durban, 1908, coll.
Miss Fontaine]. Cape Colony: [Lounsbury, 1901, p. 4 et seq., iownd
it in the western part of the Colony upon dogs, the localities mentioned
are Wynberg, Rondebosch, Claremont, Stellenbosch, places where dogs
are particularly liable to acquire piroplasmosis through the agency
of this tick ; " cases among Cape Town dogs can usually be traced to
walks up the Kloof or along the mountain sides ; the ticks appear to be
almost confined to the grass veld districts. It is the common dog tick
of South Africa"; Neumann, 1901, p. 263 records it from Kafraria and
Port Elizabeth]. Specimens have reached us as follows: (N. 121) from
dog, Grahamstown, vi. 1906, coll. T. Bowhill ; (N. 894-896, and several
other lots besides) in all stages from dog, Capetown, 1906, sent alive by
C. P. Lounsbury ; (N. 578) from Saricata tetradactgla, Deelfontein,
IV. 1902, coll. C. J. B. Grant, presented by Hon. N. C. Rothschild.
From corresj)ondence, we learn that the mortality among dogs, due
to piroplasmosis, used to be excessive at Port Elizabeth until the
introduction of our trypanblue treatment. Portuguese Congo :
(N. 1949 a.) from dog, San Salvador, XL 1912, and (N. 1950, 2652
and E. 740) from dog, Kibokolo do Zombo, iv. 1911, both lots collected

//. Icachi 473

by Dr M. (Gamble ; [Neumann, 1901, record.s it from Landana]. Congo
Free State : (N. 310) from Lion, Katanga Di.stiict, liJOT.coll. Dr A. Yale
Massey ; (N. 2012) from dog, Bongandanga, Upper Congo, 1912, coll.
W. D. Armstrong, received from Dr J. H. Ash worth of Edinburgh ;
(Liverpool 118=N. 1621) from leopard, Tshumburi, no date, coil.
Mrs Billington ; (Liverpool 103) found at Lake Leopold II, Kutu, ii.
1904, coll. Dr S. Lorisets ; (N. 2475) from leopard, Kimaka, 4. vill.
1913, coll. F. Harker ; (N. 2399) from dog, place not stated, 1913, coll.
Dr A. Yale Massey; (N. 2771) from dog, Kibondo, 1914, coll.
F. Harker. [Newstead, Dutton and Todd, 1907, p. 99, record the
tick from Tshumburi, these l)ei)ig doubtless the same specimens we
examined as reported above.] French Congo: [Neumann, 1897,
records the tick from Sette-Camma, collected by Hupferden (Hamburg
Mus.)]. Cameroon : we have determined a number of specimens from
this region as follows: (Berlin Mus. 107) $s collected by Dr Waibel ;
(D. E. M. 24, 27) $ collected by Conrad t ; the following from South
Cameroon : (Berlin Mus. 99, I0(i)(^ $ fnjm ca^, also from an unrecorde(J
host, at launde, Abong, Mbang in the Dume Region, collected by
Sommerfeld. New Cameroon : (Berlin Mus. 277) $ from Pama
Quelle, V. 1913, Dr Houy coll.; [Neumann, 1901, p. 203, states the
species occurs in Cameroon ; Zieinann, 1911, p. 5(S, found it on dogs at
Dschang, in 1910]. Southern Nigeria: (E. 40a) from sheep, Ilesha,
II. 1910; (E. 149) from dog, Onitsha, viii. 1910; (E. 38a) from dog,
Meko, II. 1910, coll. J. J. Simpson; (N. 195a) from dog, Ilesha,
N.E. District of Lagos Province, 1907, coll. W. H. Best; (E. 704)
from dog, Okigwi, 29. x. 1913, coll. Dr H. R. M. Ferguson ; (E. 297)
from pig, Bende, vi. 1911, coll. Dr P. M. Macdonald. Northern
Nigeria: (E. 147) from dog, Zungeru, via. 1910, and (E. 195 6) from
dog, Teigna, near Zungeru, viii. 1910, coll. J. J. Simpson; (N. 2198)
from civet cat, Baro, 2. X. 1910, coll. Dr J. M. Dalziel, received from
Dr J. H. Ashworth who.se collection also contains specimens from
Abinsi, 23. xii. 1912 taken by the same collector. Togoland :
[Neumann, 1901, states that there are specimens from this region
at the Berlin and Hamburg Museums]. Gold Coast: (E. GG3 a)
from cattle, Jatto's Zonga, 5. ii. 1913; (E. 670 a) from horse.
Prang, 9. ii. 1913; (E. 673 a) from horse, Makongo, 15. ii. 1913; all
three lots collected by J. J. Simpson. (E. 762) from dog, 1913, coll.
A. E. Evans; (Liverpool 75) from Kumasi, collected by Sir Rubert
Boyce ; (Liverpool 67, 99 a) from cattle and dog, coll. Dr McConnell ;
(E. 423) from dog, Addah, v. 1911, coll. Dr H. T. Palmer; (N. 3085a)

474 Genus Haemaphysalis

from hedgehog, Accra, 20. x. 1914, coll. Dr J. W. S. Macfie. Sierra
Leone: (E. 565, 575, 577) from dogs, Port Lokko, v. 1912, coll.
J. J. Simpson; (E. 581a) from dog, Kaballa, v. 1912, coll. J. J. Simp-
son ; (N. 2266) gorged o from leopard, v. 1910 and (N. 2663) adults
from dogs, 20. X. 1913, both from Kaballa, coll. Dr J. Y. Wood;
(E. 564, 569) from dogs and (E. 568) from cattle, Laminaia, iv. 1912,
coll. J. J. Simpson ; (E. 563) from cattle and (E. 572) from dog,
Mussaia, iv. 1912, coll. J. J. Simpson ; (N. 2261 c) from cattle and
(N. 2264) from goats, Koinadugu, iii.-iv. 1913, coll. Dr J. Y. Wood.
Dr J. J. Simpson moreover collected the following : (E. 619} from grass,
Mongheri, 16. ix. 1912 ; (E. 621) from Dryoscopus turetii, Newton,
20. X. 1912 ; (E. 622) from dog, York, 23. x. 1912 ; (E. 570) from dog,
Komakoni, IV. 1912; (E. 571) from dog, Konta, iii. 1912; (E. 574 =
N. 1811) from dog, Botkana, v. 1912 ; (E. 582) from dog, Hangha,
VII. 1912; (E. 557) from dog, Simimaia, iv. 1912; (E. 561) from
cattle, Bafodea, iv. 1912 ; (E. 585) from dog, Yiraia, vi. 1912 ;
(E. 586, 587) from ? Panguma and Freetown, vii. 1912 ; (E. 484) from
dog, Dani Railway, vii. 1912; (E. 604 a = N. 1923) from hush-cat,
Komatendu, viii. 1912 ; (E. 588) from dog, Firiwa, vi. 1912. We
have also received the following specimens collected by Dr J. Y. Wood
(N. 2257 a) from goat, Kagbo, 19. v. 1913; (N. 2258) from cattle
Kapankuna, 24. v. 1913 ; (N. 2269) from dog, Kasuntana, 20. v. 1913
(N. 2482 h) from sheep, Kasukura, 8. ix. 1913; (N. 2486) from dog
Fulamanca," 28. vii. 1913 ; (N. 2488 h) from cattle, Mananeolu, 2. viii
1913 ; (N. 2490) from dog, Gahnia, 25. vii. 1913 ; (N. 2492) from dog
Gaenikora, 19. vii. 1913; (N. 2654) from dog, Mussaiya, 17. xi. 1913
(N. 2658) from dog, Tilia, 13. xi. 1913; (N. 2659 a) from cattle
Boulakarafia, 11. X. 1913 ; (N. 2496 a) from dogs, Yiben, 11. ix. 1913
(N. 2499 h) from cattle, Kakonta, 6. ix. 1913 ; (N. 2502 a) from dogs
Kasukura, 9. IX. 1913 ; (N. 2328) from dogs, Dunkiawallia, 28. vi. 1913
Gambia: (E. 238a) from dog, Alijamadu, iv. 1911, coll. J. J. Simpson.
We have recently received numerous specimens, </ $ , collected by
Dr J. Y. Wood in the Koinadugu District : (N. 3035 a, 3041 h, 3043 a,
3047 a) from dogs at Falaba, 16. viii. 1914; Ninkintumania, 18. Vii.
1914; Yeria, 21. vii. 1912; Kombile, 21. viii. 1912; (N. 3038) from
Felis serval, Kaballa, 17. ix. 1914; (N. 3048 e, 3053 c), from cattle,
Dunkiawallia, 17. vii. 1914, and Boalakarafia, 22. viii. 1914; (N. 3054 e)
from sheep, Ninkintumania, 11. vii. 1914. Algeria: [Neumann, 1897,
records leachi from the nightingale, at Oran, and from grass at
Sebdou].

H. leachi 475

ASIA : Transcaucasia : (N. 786) from hear, Surnabad, viii. 1903,
coll. Dr E. Dschunkowsky. Burma : Supino's types and co-types of the
following supposed species {=H. leachi) were collected by L. Fea in
Burma during 1885-1889 : Opisthodon canestrinii (Genoa Mus. 23, 43
= N. 2960) (/ s found, according to Fea, on Felis hengalensis, Gentrococcyx
intermedius, Nicoria trijuga and Testudo elongata at Bhamo ; 0. asiaticus
(Genoa Mus. 24) cT found at Meteleo ; 0. gestroi (Genoa Mus. 22, 42
= N. 2959) $ s found on Viverra zihetha and Felis nebulosa at Yado
and Caro-Cheba ; all of these ticks proved to be but H. leachi upon
examination ; we are much indebted to Prof. Gestro for lending us
Supino's ticks for purposes of study and for allowing us to retain some
of them. India : (Ind. Mus.) from Felis tigris, Burdwar, Nepal Terai ;
(Ind. Mus. 5994/10, 5995/10 = N. 1085) numerous specimens of all stages
from Canis aureus, Museum Compound, Calcutta; (Ind. Mus. 1797/17)
very small adults and o from jackal or wolf, Jogidih, Hazaribagh District,
Chota Nagpur, 12. ill. 1913, coll. Major O. A. Smith. Specimens corre-
sponding to those described as H. leachi var. indica Warburton, reached
us as follows : Collected by Mr N. B. Kinnear : in Cutch (K. 253) from
Herpestes mungo, Nokania, vii. 1911 ; (K. 289 a) from Felis caracal,
Dhonsa, 1911 ; in Kandeish (N. 1661) from Herpestes mungo, ill.
1911 ; (N. 1663) from Felis affinis, Fardapur, iv. 1911 ; (K. 54, 56, 65)
from Herpestes rnungo, Canis indicus and Felis ajffinis respectively,
Fardapur, ill. and iv. 1911 ; (K. 9) from Herpestes mungo, Parola, ill.
1911. Specimens (K. 402 a) were taken from Millardia meltada
(Muridae), at Ghodasgaon, and (K. 220) from Cyon dukhunensis, at
Pili, Sipna Valley, Berar, in 1911. (N. 2003) adults and nymphs
from Canis indicus, Alibagh, Bombay, 25. l. 1912. We are indebted
to Captain W. S. Patton for (N. 942 b) specimens from Herpestes mango,
Madras, received in 1909. Sumatra: (Schtiffner 4a) from tiger, 1912 ;
[Neumann, 1905, p. 238, records his H. leachi var. australis ((/"s) from
Felis tigris in Sumatra]. Our H. koningsbergeri types (N. 496 c) were
collected by Dr J. C. Koningsberger from Felis pardus in Java in 1908,
and we have since received various specimens of this form from Borneo :
(N. 954 o) cf ? from dog. Upper Sarawak, XL 1909, coll. J. C. Moulton;
Federated Malay States ; (N. 1060 c) ^ from tiger, Pahang, 1910, coll.
Dr A. T. Stanton ; we have moreover examined and determined
specimens collected by Dr Stanton from Paradoxurus, Viverra and
Felis cJiaus (London School of Tropical Medicine coll.); we have
more recently received the koningsbergeri form of leachi collected by
C. Strickland (N. 2954) ^ % from Ursus malayanus, Ulu Gombak,

476

Genus Haemaphysalis

Selangor, IV. 1913, and (N. 2732) </ from Felis pardus, Kampong
Batu, Negri Sembilan, 10. ill. 1914.

AUSTRALIA: New South Wales: [Neumann, 1905, p. 238,
records his var. australis from the horse, the specimens having been
received from the Dept. of Agriculture, N.S.W.].

29. HAEMAPHYSALIS SPINULOSA Neumann, 1906.

Figs. 411, 412.

Lit. and Icon. : Neumann, 1906, pp. 212, 213, Figs. 13, 14 (reproduced) ; Donitz,
1907, p. 70 (quotes Neumann) ; Blanchard, 1909, p. 157, Figs. 196, 197 (species
only listed, figures taken from Neumann) ; Neumann, 1911 a, p. 115 (condensed
from his earlier description).

Male : Unknown.

Female (Figs. 411, 412): Scutum circular, 0'8 x 0"8 mm., with fine
scattered puuctations ; cervical grooves narrow, shallow, almost attaining
the posterior border. Gapitidntn : base rectangular, twice as broad as
long, cornua faintly indicated by the sharp postero-lateral angles ;
porose areas small, oval, far apart ; hypostome 4 | 4, with 8 or 9 teeth
per file ; palps short, article 2 pointed at the postero-lateral angle,
four hairs on the internal ventral border, a short blunt prominence at
the postero-ventral and postero-dorsal borders, the dorsal prominence
nearer the external angle, article 3 longer ventrally where it bears a
short retrograde spine. Venter : vulva between coxae II ; spiracle
small, transversely oval. Legs : coxae all bear at their internal angle a

Fig. 411. Fig. 412.

Fig. 411. H. spi7ndosa 9 , capitulum and scutum. Neumann, 1906, Fig. 13.
Fig. 412. H. spinulosa ? , capitulum in ventral aspect, coxae I. Neumann, 1906,
Fig. 14.

H. spinnlosa and H. ohtusa 477

pointed spur slightly longer than broad ; trocJianters unarmed ; tarsi
long, narrow, not humped; pad almost as long as the claws.

Description based on 2 % from Uganda, collected by E. Degen.

Types in the British Museum.

The species is allied to or a varietal form of H. leacJti.

30. HAEMAPHYSALIS OBTUSA Ddnitz, 1910.

Fig. 413.

Lit. and Icon. : Donitz, 1910, p. 492, PI. XVII, Figs. 11, 12 (here reproduced).

Male (Fig. 413) : Very small, colour brownish-grey. Scutum
1'5 X 0'9\ punctations few and fine, cervical grooves short, lateral
grooves absent: festoons much longer than broad. Capitulum: base
broadest in front, the antero-lateral margins broadly rounded ; cornua
slight. The palps form a very obtuse cone (hence ohtusa), thrice as
broad as long ; article 2 much longer than 3, broad basally, a very slight
spur under article 3. Hypostome 2 | 2. Legs : coxae unarmed ; tarsus
IV not humped, but narrowing abruptly.

Fig. 413. H. obtusa i . Capitulum in dorsal and ventral aspects, with front of scutum
and coxae I. Redrawn from Donitz, 1910, PI. XVII, Figs. 11 and 12.

Female : unknown.

Described originally from numerous cfs from the Island of Reunion.
H. ohtusa is a very small species of the general facies of H. leachi, but
distinguished from it by the different conformation of article 2 of the
palps, the dentition of the hypostome and by the absence of lateral
grooves and coxal armature. We are inclined to regard it as a
doubtful species (see discussion p. 467).

Types in the Berlin Museum, co-type (N. 2816) in Cambridge.

^ The measurement cited is that of Donitz. The scutums of 3 <? sent to us for study
from Berlin measured as follows in mm. :

1-46 X 0-87 1-43 X 0-82 1-36 x 074

N. I, 31

478

Genus Haemaphysalis

31. HAEMAPHYSALIS NUMIDIANA Neumann, 1905.

Figs. 414, 415.

Lit. : Neumann, 1897, p. 349 (included under H. leachi) ; 1905, p. 230 ; Donitz,
1907, p. 70 (quotes Neumann) ; Blanchard, 1909, p. 154 (merely listed from
Neur^iann) ; Neumann, 1911a, p. 112. The species has not hitherto been
figured.

H. numidiana, which we regard as a somewhat doubtful species
(see p. 467), is very large compared with the average H. leachi, the
cT measuring 3"5 x 2'0 mm. In the case of an exceptionally large
H. leachi all our experience would lead us to expect that salient
characteristics would be emphasized, that the spines on the palps for
instance, would be more pronounced than usual. Instead of this being
the case, article 2 of the palps in H. numidiana is almost spineless.
The general facies is much like that of H. leachi, the body being
elongate, the lateral grooves and punctations being similar, it differs,
however, from H. leachi in that the basis capituli is rectangular, whereas
in typical H. leachi it is trapezoidal ; the palps meet in front at a less
obtuse angle than they do in H. leachi ; for further details the reader is
referred to the figures (Figs. 414, 415).

Fig. 414. Fig. 415.

Fig. 414. H. numidiana i , dorsum, part of venter, tarsus IV aud spiracle. The finely
stippled areas indicate depressions in the scutum. Drawn from the type. Original,
G. H.F.N, del.

Fig. 41-5. H. numidiana ? , capitulum and scutum, spiracle. Drawn from the tyije.
Original, G. H.F.N, del.

H. nmnldimia and H. Welllnijtoni 479

Through the courtesy of Professor Neumann, wu have been able to
examine a </ and % from his collection (No. 999) and he has presented
us with a co-type (N. 2890 ^). The types, 3 cT 1 $, were taken from
a hedqehog, at Tebessa, Algeria, in 1894 by Fayet. Neumann, 1897,
p. 349, originally included the species under H. leachi.

Types in Toulouse (2 cT 1 $ ), co-type in Cambridge (1 ^).

32. HAEMAPHYSALIS WELLINGTONI Nuttall and
Warburton, 1907.

Figs. 416-420.

Lit. and Icon. : Nuttall and Warburton, xii. 1907, pp. 397-398, Figs. 9-11
(reproduced). Blanchard, 1909, p. 157 (species ouly listed).

Male (Fig. 416): Scutum 1"42 x l"l to 1'3 x 1 mm.\ oval, narrow in
front, dark brown, glossy, with many medium-sized punctations,
especially posteriorly on the pseudoscutum ; a non-punctate depression
on either side of the centre ; cervical grooves deep, straight, parallel ;
lateral grooves short, including first festoon. Venter: brown; spiracles
white, bluntly piriform. Capitulum proportionately large (03 mm. I.),
base finely punctate, cornua slight ; palps : article 2 projecting strongly
laterally ; article 3 with dorsal and ventral retrograde processes at inner
angle ; the outer borders of articles 2 and 3 form an unbroken line ;
hypostome broad, dentition 4 | 4, about 11 teeth per file. Legs: coxae
strong, with a single short spur on each, strongest on coxa I.

Female (Figs. 417, 418): Bodi/ (unfed), 1-5 x 11 mm., brown,
punctate, marginal grooves well marked. Scutum oval, about
0"9 X 0*7 mm.\ sides nearly parallel, many rather large punctations,
some confluent ; cervical grooves long, nearly parallel, middle field broad.
Vente?^ : brown ; spiracles white, rounded. Capitulum : proportionately

' The scutums of 10 <i and 9 ? specimens measured as follows in mm. :

<?

?

1-42 X 11

(N. 2089)

0-9 xO-91

(N. 2234)

1-42 X 105

,,

0-9 xO-8

,,

1-4 xl-06

,,

0-9 xO-79

,,

1-38 X 1-07

,,

0-9 xO-7

(N. 221)

1-38x105

,,

0-86 X 0-71

,,

1-37 X 1-08

,,

0-84 X 0-79

(N. 2234)

1-37 X 0-96

(N. 2232)

0-83 X 0-87

,,

1-36 X 1-05

»>

0-8 xO-75

)>

1-33 X 0-93

(N. 221)

0-76x0-ft

,,

1-3 X 1-0

31—2

480

Genus Haeinaphysalis

Fig. 416. Fig. 417.

Fig. 416. H. welUngtoni g , dorsum and venter.

Fig. 417. H. loellingtoni ? , dorsum. Nuttall and Warbiirton xii. 1907, Fig. %, A, B
and C. G. H.F.N, and E.W. del.

Fig. 418. H. welUngtoni ?. H. hypostome, 250 ytt 1. D. digit 100 ;t 1. Nuttall and
Warburton, xu. 1907, Figs. 10, 11. G. H. F. N. del.

Fig. 419. H. welUngtoni nymph. Capitulum in dorsal and ventral aspects, scutum,
coxae with trochanters, spiracle and tarsus IV. (N. 1057 from Selangor.) Original,
N. C. del.

H. loellmgtoni

481

very large (0"G mm. I.), with characteristics of </ (hy[)ostome, Fig.
418) ; porose areas large, oval, far apart. Legs : paler than body, coxae
as in f^.

Nymph (Fig. 419) : Scutum 0"4.5 x 0'44 mm., about as broad as
long, deeply emarginate, with distinct sub-parallel- cervical grooves
fading away towards the narrow posterior border. Gapitulum : base
broad, cornua short, lateral salience of article 2 of palps slight ; hypo-
stome 2 | 2, with 6 teeth per file. Venter : spiracle transversely ovoid.
Legs: coxae recalling those of the adult, tarsus IV short and tapering ;
pad long.

Larva (Fig. 420) : Scutum 0-25 x 0"38 mm., broader than long,
slightly emarginate, scapulae broadly rounded ; cervical grooves sub-
parallel, poorly defined. Capitulmn: base with concave posterior
margin, without cornua, lateral salience of article 2 of palps slight ;
hypostome as in the nymph. Legs : coxa I as in the nymph, coxae
II and III with protruding flange ; tarsi as in the nymph.

Fig. 420. H. tvellinrjtoni larva. Gapitulum in dorsal and ventral aspectH, scutum, coxae,
tarsus III. (N. 10-57 from Selangor.) Original, N. C. del.

Our description is based on (N. 221) 2 cf, H ? and 2 O taken from
domestic fowl at Sarawak, Borneo, collected by Dr A. R. Wellington
I. 1907. We have since received the following consignments: (N. 294),
8 </, 3 ?, 10 o (data as under N. 221), collected v. 1907. Siimatra:
(N. 495 c), 4 ?, from Bos indicus or huhalis, collected in 1908 at
Palembang by Dr J. C. Koningsberger ; (Schtiffner's coll., 4 6) a ^Z*
found on a goose.

Federated Malay States: (N. 1014), 1 ?, from domestic fowl,
Kuala Lurnpui-, 1910, and (N. 1057) 2 </, 2 O, 25 larvae from domestic
fowl, Selangor, collected ill. 1910, by Dr A. T. Stanton. Siam : Dr
R. Stokoe has sent us (N. 2232), 0 cT' 1 ? and 3 o from domestic fowl ;

482

Genus Haemaphysalis

(N. 2233), 6 ? , from buffalo ; and (N. 2234) 7 % from c?o^ ; the three
lots having been collected, iv. 1913, at Lakhon, Lampang. Dr A. F.
Kerr has sent us (N. 2089) 7 (^ from turkey, collected 10. xii. 1912, at
Chiengmai, by Mrs Harris. We determined 1 ? from the Andaman
Islands which is in the Indian Museum, Calcutta (no data or number).
Types in Cambridge (N. 221, ^ % ; ^. 1057, o L). We have
presented specimens, since compared with the types (N. 294 </ % o), to
the British Museum and (</ ? ) to the Zoological Museum, Berlin.

33. HAEMAPHYSALIS DOENITZI Warburton and

Nuttall, 1909.

Figs. 421-422.

Lit. and Icon. : Warburton and Nuttall, 1909, pp. 64, 65, Figs. 9, 10 (reproduced).

Male (Fig. 421): Scutum about 1*3 x 0*9 mm.\ long-oval, narrower
in front, glossy yellow-brown, finely punctate ; cervical grooves nearly
straight, converging posteriorly ; lateral grooves long, curved, deep,
including two festoons ; festoons long, the dividing lines curved.

o

Fig. 421.

Fig. 422.

Fig. 421. H. doenitzi <r . Dorsum. (Sketch by C. W.) Warburton and Nuttall, 1909,

Fig. 9.
Fig. 422. H. doenitzi J . Capitulum and scutum, spiracle and tarsus IV. (Sketch by

C. W.) Warburton and Nuttall, 1909, Fig. 10.

^ The scutums of our other specimens measure as follows in mm. :

i ?

1-4 xO-8 0-78 X 0-68

1-25 X 0-8 0-75 X 0-7

H. doenitzi 483

Capitulum 03 mm. long, base trapezoidal, broader in front, punctate,
with short pointed cornua ; hypostome 4 1 4 or 5 1 5 with very small
teeth ; palps long, articles 2 and 8 of" about equal length, article 2
strongly salient at the base, article 3 without dorsal process but with
a very small ventral spine. Venter : spiracle ovate, bluntly pointed
dorsally ; anal grooves rather ogival. Legs : coxae normal ; tarsus IV
of medium length, tapering rather abruptly.

Female (Fig. 422): Scutum oval, about 08 x 0"6 mm.^ deeply
emarginate, uniformly punctate with rather large punctations ; cervical
grooves sub-parallel, hardly visible for more than half the actual
length. Capitulum 0'4 mm. long, base rectangular, with very slight
cornua; porose areas long-oval, far apart; palps as in the </, but
longer ; hypostome narrow, rather spatulate, 4 j 4, very small teeth.
Dorsum strongly and coarsely punctate. Venter : spiracle sub-circular,
with blunt dorsal process ; anal grooves ogival. Legs : coxae and
tarsus IV as in the cf .

Described from (N. 635) 4 cT and 3 $ from water-hen, St John's
Island, Singapore, 1909, collected by Dr A. R. Wellington, and named
in honour of the late Geheimrath W. Donitz of Berlin.

Types in Cambridge, a co-type cT was presented to the Berlin
Museum.

A very small species closely allied to H. hoodi.

34. HAEMAPHYSALIS HOODI Warburton and
Nuttall, 1909.

Figs. 423-426.

Lit., Icon, and Syn. : Haemaphysalis hoodi Warburton and Nuttall, 1909 (June),
pp. 62-63, Figs. 7, 8 (reproduced).
Haemaphysalis africana C. W. Howard, 1909 (August), reprint 4 pp.,
PI. XXXIV, Figs. 1-10 capitulunis of (J, J, o and larva; $ dorsum
and venter ; $ scutum ; digits of (^ , ^ ^^^d o (not reproduced).

Male (Fig. 423): Scutum IS x 13 to 1-4 x 1-05 mm.'^ oval, rather

1 See note 1, p. 482.

- The scutums of 4 (? and 7 ? measured respectively in mm. :
S ?

1-8 X 1-3 1-0 xO-9

1-7 X 1-2 0-95 X 0-8

1-7 X 1-1 0-9 X 0-9 (2 specimens)

1-4 X 1-05 0-85 X 0-85

0-75 X 0-75

484

Genus Haemaphysalis

broad, narrowing in front, with many fairly large punctations ; cervical
grooves moderate, deep anteriorly, concave externally ; lateral grooves
well-marked, of medium length, including one festoon. Capituhim
rather short (O'S mm. L); base rectangular, punctate, with sharp
cornua; palps with article 2 very salient laterally at right angles
to the axis, article 2 slightly longer than article 3, article 3 with a
very small ventral process directed inwards; hypostome well covered
with equal teeth, 4 ] 4, about 10 teeth per file. Venter : spiracle sub-
rectangular, with blunt dorsal process. Legs: coxa I with blunt
internal spur; coxa II with a slight conical spur near the internal
angle ; coxa III almost unarmed, a blunt spur at the inner angle of
coxa IV ; tarsus IV of medium length, abruptly narrowing at its tip.

Fig. 423. Fig. 424.

Fig. 423. H. hoodi s . Dorsum, tarsus IV, spiracle, palp in ventral aspect. (Sketch by

C. W.) Warburton and Nuttall, 1909, Fig. 7.
Fig. 424. H. hoodi ? . Capitulum and scutum, tarsus IV, spiracle. (Sketch by C. W.)

Warburton and Nuttall, 1909, Fig. 8.

Female (Fig. 424) : Body (unfed) yellow with darker border, well-
marked off by deep marginal grooves, which are nearly parallel ;
festoons square. When gorged, the scutum generally shows yellow on
the dark body. 8cutum oval 1 x 09 to 0 75 x 0*75 mm.^ truncated
posteriorly ; cervical grooves sub-parallel, extending about two-thirds
its length ; punctations fairly large and uniformly distributed. Capitulum :
0'4 mm. long; base sub-rectangular, with posterior border somewhat
concave and cornua almost obsolete ; porose areas reniform and situated
anteriorly ; well separated ; hypostome 4 j 4, like that of j/* ; palps

1 See note 2, p. 483.

H. hoodi

485

longer than in ^ , es[)ecially article 2, which is nut so abruptly salient
laterally. Venter : spiracle rather large, ovate, the pointed end dorsal ;
anal grooves semi-circular. Legs: like those of the^/, but tarsus IV
is longer and tapers less abruptly.

Nymph (Fig. 425) : Scutum 0'() x O'G mm., sub-circular, the postero-
lateral borders slightly concave ; cervical grooves long, sub-parallel.
Gapitidum resembling that of $ ; base much broader than long, with
well-marked cornua ; palps with article 2 very salient laterally ;
hypostome dentition 2 1 2, 5 to 6 teeth per file. Legs : coxae I-IV
with very slight spurs ; tarsus IV stout.

Fig. 4^5. H. lioodi o. Capitulum and scutum, capitulum in ventral aspect, coxae and
tarsus IV. (N. 1364.) Original, G. H. F. N. del.

Larva (Fig. 426) : Scutum 0-25 x OS mm., broader than long,
cervical grooves long and parallel. Cwpitulum : base with slight cornua ;
palps conical, article 2 moderately prominent laterally ; hypostome
dentition 2 j 2, 4 to 5 teeth per file. Legs : coxa I with slight blunt
spur, coxae II and III unarmed; tarsi tapering.

c?

a

Fig. 426. H. hoodi larva. Capitulum and scutum, capitulum in ventral aspect, coxae
and tarsus III. (N. 1364.) Original, G. H. F. N. del.

486 Genus Haemaphysalis

Our description of the adults is based on numerous specimens
(N. 424, cfs $s 1 o) occurring on fowls, V. 1908, at Bathurst, Gambia,
West Africa, presented by Dr P. Hood. The nymph and larva are now
described and figured from specimens (N. 1364) found with cfs and
?s on the neck of a bird {Gentropus burchelli), shot 3. vi. 1908, at
Manhica, in the northern district of Lourenyo Marques, Province of
Mozambique, Portuguese East Africa. We are indebted to Mr C W.
Howard, formerly Entomologist for the Province, for the gift of these
specimens ; he described all the stages of this tick under the name
of Haemaphysalis africana in August, 1909. Our description was
published in June, 1909, under the name of Haemaphysalis hoodi,
which consequently has priority.

We have also received the following specimens : (N. 1093 and E. 12)
cfs $s, from the ears of partridges, Nyungwi Stream, N. Nyasaland,
British Central Africa, collected by Dr J. B. Davey, xi. 1909 ; (N. 1546
= E. 422), j/s $s, from Numida 'tneleagris (guinea-fowl), Addah, Gold
Coast, West Africa, collected by Dr H. T. Palmer, x. 1911 ; (E. 606),
1 </, from cuckoo {Gentropus senegalensis), Freetown, Sierra Leone, West
Africa, collected by Major H. Kelsall, v. 1912 ; (E. 473), 1 ^, from a
bird {Gymnoschizoi'his leopoldi Shelley), S. E. Ankole, British East
Africa, collected by S. A. Neave, x. 1911.

Types in Cambridge (N. 424, adults and o; N. 1364, o and L) ;
co-types ((/ $ from our collection) at Toulouse.

Haemaphysalis hoodi var. orientalis Nuttall and
Warburton, 1915, n. var.

Male : Differs from the type as follows : Body more parallel-sided,
narrowing less in front. Scutum^ more coarsely punctate ; emargination
slight ; scapulae blunt ; cervical gi^ooves deep and converging at first,
then shallow and diverging ; lateral grooves longer than in the type ;
festoons short. Gapitulum : smaller than in the type, the base par-
ticularly small, without cornua, narrowing posteriorly; palps shorter;

^ The scutums of the type lot measure in mm. respectively :
5c? 3?

1-74 X 11 0-93 X 0-76

1-7 xl-2 0-92 X 0-8

1-7 xl-2 0-9 xO-8
1-7 xl-1
1-65 X 1-0

H. hoodi var. orientalis 487

hypostome 4 1 4, 6 teeth per file. Venter: spiracle transversely
elongate, pointed dorsally. Legs : coxa I bluntly pointed, coxae II-IV
unarmed ; trochanter I with blunter spur.

Female : Differs from the type as follows: Scutum^ more pointed
posteriorly, more coarsely punctate ; cervical grooves shorter, deep,
straight and parallel. Capitulum : hypostome short, 4 1 4, about 7
teeth per file. Venter : spiracle very small. Legs : coxae practically
unarmed.

Described from 6 (f and 3 $ taken from Procavia inanningi
Wroughton, S.W. Shore of Lake Nyasa, British Central Africa,
III. 1910, S. A. Neave coll.

Types in Imperial Bureau of Entomology coll., London (Nos. 66 a
and 69) ; co-types in Cambridge (N. 2847 and N. 2848, 1 (/" 2 ? ).

35. HAEMAPHYSALIS BANCROFTI Nuttall and
Warburton, 1915. n. sp.

Figs. 427-430.

Male (Fig. 427): Scutum: oval, rather narrowed in front, 2 x 1-25
to 1'3 X 085 mm.-, rather coarsely, but not deeply, punctate; cervical
grooves well-marked but short, the interval between them rather wide ;
lateral grooves fairly long, including one festoon ; festoons longer than
broad. Capitulum: base broader than long, broader anteriorly than
posteriorly; cornua small but distinct and sharp; palps strongly
salient laterally at the base of article 2, article 2 longer than article 3,
their lateral contour an almost unbroken curve ; no dorsal spines ; a

1 See note 1, p. 486.

2 The scutums of 9 (? and 11 ? measured as follows in mm. :

9^

11?

2 X 1-25 (N. 2115)

0-83 X 0-83 (N. 2114)

1-8 xl-2 (N. 2691)

0-83 X 0-8 (N. 2100, type)

1-8 xl-15 (N. 2100)

0-8 xU-9 (N. 2691)

1-75 X 1-15 (N. 2691)

0-8 xU-83

1-6 xl-1 (N. 2100, type)

0-8 xO-8 (N. 2689)

l-55xl-0 (N. 2691)

0-76 X 0-86 (N. 2100)

1-5 xl-0

0-76 X 0-83

1-35 X 1-0

0-75 X 0-9 (N. 2691)

1-3 xO-85

0-74 X 0-9 (N. 2689)

0-67 X 0-7 (N. 2100)

0-62 X 0-8

488

Genus Hae^naphy sails

moderate retrograde spur under article 3 ; the lateral salience of article
2 is somewhat arched to accommodate coxa I, and its posterior border
is curvilinear both dorsally and ventrally ; hypostome dentition 4 j 4,
with 6 to 7 teeth per file, without median interval. Venter : spiracle
of medium size, ovate, the narrow end postero-dorsal. Legs: coxae

Fig. 427. H. bancrofti 3 , dorsum, capitulum in ventral aspect, coxae with trochanters,
spiracle and tarsus IV. (N. 2100 type.) Original, N. C. del.

Fig. 428. H. bancrofti ? , scutum, capitulum in dorsal and ventral aspects, coxae with
trochanters, spiracle and tarsus IV. (N. 2100 type.) Original, N. C. del.

H. hancrofti

489

with normal armature ; extremely small indications of spurs on the
trochanters ; tarsus IV short and blunt.

Female (Fig. 428) : Scutum sub-circular, 0-83 x 0-83 to 0-62 x 0-8
mm., coarsely punctate, the punctations less nunu-rous in the median
area, and often confluent at the sides; emargination deep; cervical

Fig. 429. H. hancrofti nymph. Scutum, capitulum in dorsal and ventral aspects,
coxae with trochanters, spiracle and tarsus IV. (N. 2114 co-type.) Original,
N. C. del.

Fig. 430. H. bancrofti larva. Scutum, capitulum in dorsal and ventral aspects, coxae,
tarsus III. (N. 2689 co-type.) Original, N. C. del.

490 Genus Haemaphysalls

grooves long, deep and sub-parallel, the interval rather broad. Capitu-
liim greatly resembles that of the cT ; porose areas much longer than
broad, converging strongly in front ; hypostome dentition 4 | 4. Venter :
spiracle oval, hardly pointed dorsally. Legs : as in the ^.

Nymph. (Fig. 429) : Scutum: broader than long, about 0"2 x 0*25 mm.^,
cordate ; cervical grooves long, sub-parallel, well-marked. Capitulum :
much like that of the </, except that the dentition is 2 j 2. Legs: like
those of the adult, with trochantal spurs more distinct.

Larva (Fig. 430) : Scutum: much broader than long, about 0*37 x 0*58
to 0"32 X 0"56 mm.^ rather deeply emarginate ; cervical grooves well-
marked and visible for the whole length of the scutum. Capitulum :
base twice as broad as long, with distinct, rather sharp, somewhat
outwardly directed cornua ; palps only slightly salient laterally, article 2
about equal in length to article 3, no dorsal spines, but a short retrograde
spur under article 3 ; hypostome dentition 2 | 2. Legs : coxal armature
normal ; no trochantal spurs ; tarsus III tapering rapidly to a rather
sharp point ; pad half as long as the claws.

Described from specimens derived from Queensland, Australia :
(N. 2100), 2 {/ and 5 $ from Macropus dorsalis, Burnett District, ill.
1913; (N. 2114), 1 $ and 1 o, found crawling on collector's trousers, at
Brigaton Scrub, Burnett; (N. 2115), 1 (/, from Macropus giganteus,
Burnett District, ill. 1913; (N. 2689), 2 $, with eggs and larvae, the
$s taken from Macropus dorsalis, Eidsvold, xii. 1913; (N. 2691), 6 </,
3 % and 2 o, host (?) found at Eidsvold. All collected by Dr T. L.
Bancroft, of Eidsvold, after whom the species is named. Mr L. Harrison
has recently presented us with (N. 3090) (/ o from Bettongia penicillata,
Queensland.

Types in Cambridge: (N. 2100, ^ % ; ^. 2691, o; N. 2689,
larvae).

i The scutums of 3 nymphs (N. 2114, N. 2691) and 12 larvae (N. 2689) measured as

follows in mm. :

o L

0-.36xO-5
0-36 X 0-52
0-35 X 0-54
0-35 X 0-51
0-34 X 0-52
0-33 X 0-52
0-33 X 0-51
0-32 X 0-56

0-2

xO-26

0-2

xO-25

0-18

xO-22

L

0-37

xO-58

0-36

xO-58

0-36

xO-56

0-36

xO-54

H. camjMnulata

491

36. HAEMAPHYSALIS CAMPANULATA Warburton,

1908.

Figs. 431-4.32.

Lit., Icon, and Syn. : H. campanulata Warburton, 1908, pp. .")13-514, Fig.s. .5, 6
(reproduced).
H. campanulata Warburton, 1908, wrongly stated to = H.Jlava Neumann, 1897,

in Blanchard, 1909, p. 148.
See discus.siou under H. flava, p. 408.

Male (Fig. 431) : Body oval, slightly convex, colour yellow, lighter
ventrally. Scutum about 2 x 1*4 mm.^ with numerous fine punctations;
cervical grooves deep, sub-parallel, slightly convex externally; lateral
grooves beginning about the anterior third of the body and ending
behind the spiracles ; festoons long and narrow, well-marked, the

i^uiy 0

Fig. 431. Fig. 432.

Fig. 431. H. campanulata i . Dorsum, coxae and spiracle. Sketch, Warburton, 1908

Fig. 5.
Fig. 482. H. campanulata ? . Capitulum and scutum, tarsus IV, and spiracle. Sketch,

Warburton, 1908, Fig. 6.

^ The scutums of 11 <? and 5 ? measured as follows in mm. :

2-1 xl-5 (N. 579)

2-0 xl-4 (2N. 897)

2-0 X 1-4 (N. 1251 h)

1-95x1 -35 (N. 579)

1-95 X 1-3 (N. 897)

1-9 xl-35

1-9 A 1-3

1-85 xl-4

1-8 xl-25

1-G xl-15

1-55x1-1 (N. 1251 «*)

1-25x1-1 (N. 579)

1-15x1-05

1-1 xl-0 (N. 897)

1-1 xO-95

1-0 xl-15 (N. 579)

492 Genus Haemapliy sails

intervals sometimes dark. Venter : sexual orifice broad, between
coxae II ; spiracles almost trapezoid, with blunt rounded dorsal pro-
tuberance. Capitulum : base rectangular, dorsally barely twice as
broad as long, punctate all over, and with short blunt cornua ; the
rest of the capitulum notably bell-like in shape (hence campanulata)
owing to the forwardly curved lateral projections of article 2 of the
palps ; hypostome 4 j 4, files of broad teeth, about 8 teeth per file,
the outermost teeth the stoutest ; palps without any dorsal spines, but
the ventral border of article 2 projects strongly backwards, and article 3
has a well-marked ventral retrograde spine. Legs: coxae each with a
short blunt spur ; tarsi IV very short and thick, ending in a somewhat
abrupt slope, the false articulation about the middle of the article.

Female (Fig. 432) : Body yellow, somewhat parallel-sided, some-
times attaining 7x4 mm. when distended ; marginal grooves including
two festoons. Scutum : 1*25 x I'l to I'l x 1 mm.\ glossy-brown or
yellow, cordiform, finely punctate ; cervical grooves well-marked, far
apart, at first convex outwardly, then turning outward. VeJiter :
yellow; spiracles like those of the (^, but with dorsal protuberance
even less marked. Capitulum : base broader and shorter than in the </,
with less distinct cornua ; porose areas oval, their axes converging
forward, far apart, the interval twice their diameter, a narrow longi-
tudinal pit midway between them ; the rest of the capitulum like that
of the cT, but even more markedly bell-shaped; hypostome 4 | 4, with
teeth narrower than in the ^T. Legs: coxae as in the cT, tarsus IV
short, somewhat humped.

Description based on numerous specimens (N. 897) taken from
Chinese dogs in Mongolia by Major M. L. Hearn, and received by
us from Colonel B. Skinner, R.A.M.C, in 1906. They agreed in every
respect with specimens kindly lent by Professor Neumann (3. III. 1908)
as H. flava, and with others so named by him in the British Museum.

We have also received: (N. 524a), 1 $, collected with other ticks,
from cattle and horses, at Wei-Hai-Wei, China, by Dr W. M. Muat,
X. 1908 and (Berlin Mus. No. 166) $s from Chengtu, China, collected
by Consul Weiss ; (N. 579), 2 cf 2 $ , from a dog, Yokohama, Japan,
collected by A. Owston, iv. 1902, and presented to us by Hon. N. C.
Rothschild; Dr Miyajima has sent us adults (N. 2917, 2921) found
on house rat, 11. ix. 1912, and on dog, 18. vi. 1914, at Tokio.
We include here (N. 2886) 1 ? from dog, Tokio, 1894, Janson coll.

1 See note 1, p. 491.

H. howletti

493

(a purported co-type of H.flava Nn.) presented to us by Prof. Neuniarui.
(N. 1251 b), 2 (/, from deer, Satharangapara, Travaricore, S. India,
collected by E. R. Howlett, i. 1911.

Types in Cambridge (N. 897); we have presented co-types (</ ?)to
the British Museum and London School of Tropical Medicine, also (</)
to the Museums in Paris, Berlin, Toulouse (Neumann coll.), and
Washington, D.C. (U.S. Dept. Agric).

37. HAEMAPHYSALIS HOWLETTI Warburton, 1913.

Figs. 433, 434.

Lit. and Icon. : Warburton, vii. 1913, pp. 123, 124, Figs. 2, 3 (reproduced).

Male (Fig. 433) : Of the general appearance of H. campanulata
Warburton, 1908, from which it differs, however, as follows: it is
smaller; the scutum, 175 x 1-2 mm., more densely and deeply punctate,
less rounded posteriorly ; sjyiracle sub-rectangular, with very slight
blunt, dorsal process ; hypostome 5 | 5 to 6 j 6, very small sharp teeth ;
tarsus IV longer ajid more tapering.

Fig. 433. H. howletti i , capitulum, scutum, profile of capitulum, anus with anal
grooves, coxae and spiracle. (N. C. del.) Warburton, 1913, Fig. 2.

N. I. 32

494

Genus Haemaphyscdis

Female (Fig. 434) : Scutum longer than broad, I'O x O'SS mm., long-
oval, of nearly uniform width throughout, strongly and deeply punctate,
the punctations more or less confluent on the lateral fields; cervical
grooves beginning as pits at some distance from the anterior border,
and continuing as shallow depressions for about | the scutal length.
Capitulum : much like that of H. canipanulata ; porose areas oval and
far apart, with a median depression between them ; hypostome 5 | 5,
very small teeth confined to distal portion. Dorsum closely and deeply
punctate all over. Legs : tarsi tapering.

Fig. 434. H. howletti ? , capitulum, scutum, coxae and trochanters, anus with anal
grooves, tarsus IV and spiracle. N. C. del. Warburton, 1913, Fig. 3.

Described from (N. 1979) 1 </ and 1 $ from hill pony, Rawalpindi,
Punjab, India, 1912, collected by F. M. Hewlett after whom the species
is named.

Types in Cambridge.

This species is closely allied to H. camjxinulata. It is, however,
smaller, much more punctate and its hypostome is very different, that
of campanulata being well covered with 4 { 4 large teeth. The different
shape of the $ scutum and the tapering tarsi in both sexes will also
serve to distinguish it from that species.

//. vidua

495

38. HAEMAPHYSALIS VIDUA Warburton and
Nuttall, 1909.

Fig. 435.

Lit. and Icon. : Warburton und Nuttall, 1909, pp. 66-67, Fig. 13 (reproduced).

Male (Fig. 48.5) : Scutum : IS x 1-1 mm, yellow, oval, broadest in the
middle, with very numerous shallow punctations ; cervical grooves deep
oval pits with shallow divergent posterior continuations ; no lateral
gi-ooves. Gapitulum: short; base rectangular, comparatively long,
with short blunt cornua ; hypostome with large corona followed by 4 | 4
rather scale-like teeth, few in number, about 30 in all ; palps with

3^

Fig. 435. H. vidua <? , dorsum, coxae, spiracle, tarsus IV. Sketch by C. W. Warburton
and Nuttall, 1909, Fig. 13.

articles 2 and 3 about equal in length, and with article 2 salient near
its distal end, and forming a sinuous contour with article 3 ; a ventral
retrograde spur from the outer border of article 3. Venter : spiracle
oval, with slight dorsal prominence ; anal grooves ogival. Legs : not
very strong, coxae with a moderate spur at each postero-internal angle,
strongest on coxa IV ; tarsi thick and abruptly tapering, almost
humped.

Female : unknown.

32—2

496

Genus Haemapliy sails

Described from 1 ^ taken (in company with H. leachi) from
Paradoxurus sp., in the Federated Malay States, by Dr A. T. Stanton,
III. 1909.

Type in Cambridge.

39. HAEMAPHYSALIS HUMEROSA Warburton
and Nuttall, 1909.

Figs. 436, 437.

Lit. and Icon. : AVarburtoii and Nuttall, 1909, pp. 60, 61, Figs. 4, 5 (reproduced).

Male (Fig. 436) : Livid yellow-brown, very long and narrow.
Scutum: 1-95 x 0"9 to 2'37 x 1'04 ram.}, glossy, with very uneven
surface due to more or less confluent punctations and ridges ; cervical

Fig. 436.

Fig. 437.

Fig. 436. H. humerosa j , dorsum, coxae, spiracle. Sketch by C. W. Warburton and

Nuttall, 1909, Fig. 4.

Fig. 437. H. humerosa 9 , capitulum and scutum. Sketch by C. W. Warburton and

Nuttall, 1909, Fig. 5.

1 The scutums of 16 <? gave the following measurements in mm. :

2-37 X 1-04 2-1 xl-0

2-3 xO-97 2-1 xO-84

2-2 xO-95 2-03 X 0-98

2-2 xO-95 2-0 xO-92

2-16 X 1-0 2-0 xO-9

2-16 X 0-94 2-0 xO-87

2-15x0-9 1-97 X 0-96

2-14x0-94 1-85x0-87

I

H. humerosa 497

grooves long and nearly straight, slightly diverging posteriorly; lateral
gi'ooves long and very near the border, including one festoon ; festoons
moderately long, the intervals curved, Capitulum 08 mm. long, base
rather long, with long sharp cornua proceeding from the dorsal surface.
Hypostome with very small teeth, 5 | 5. Palps long, especially article 2,
which is about four times as long as article 3 ; its sharp external angle
being much posterior to its insertion ; no dorsal spur ; a very small
sharp spur under article 3. Venter : genital aperture between coxae II,
covered by an elongate shield ; spiracle large, broader posteriorly,
without definite dorsal process. Legs : coxae very broad (antero-pos-
teriorly) and progressively increasing in size, all more or less pointed
at the internal angle, the spur on coxa IV dark and conical ; coxa I
protrudes beyond the scapular angle and rises to the level of the dorsal
surface, giving a characteristic appearance (hence humerosa).

Female (Fig. 437) : Of the same colour as the (^. Scutum oval,
0"8 X 0"6 to 1*0 X O'GQ^ ram., cervical grooves only slightly concave and
attaining the posterior border. Capitulum 0'4< mm. long, like that of
the cT, but with article 2 rather less salient, and only about thrice as
long as article 3 ; hypostome 5 | 5, very small teeth ; basis capituli
ridged laterally and anteriorly, the sub-circular porose areas are very
faintly indicated at the sides of the flat median area. Legs like those of
the rT ; tai-sus IV medium, rather thick, tapering.

Nymph : Strongly resembles the $ , having the same elongated
body and the same type o{ capitulum, which is, however, shorter in com-
parison to its breadth. The scutum about as long as broad, 0'4 x 0'4 mm.- ;
punctations few and scattered ; cervical grooves are deep broad furrows
not quite attaining the posterior border. Legs as in the % but with
shorter sharply pointed tarsi IV.

Described from (N. 669) 3^/1? and 1 o taken from Perameles
macrura, Barrow Island, N.W. Australia, and presented by the Hon.
N. C. Rothschild. The measurements appended in the footnotes relate to
specimens brought from Australia by Mr L. Harrison : 2 cT 3 $ 5 o
from Perameles macrura, Stapleton, Northern Territory, 31. XI. 1913,

1 The scutnms of 5 ? gave the following measurements in mm. :
1-0 xO-GG 0-95xO-6G

0-97 X 0-7 0-9 xO-6

0-97 X 0-65
The scutums of 5 nymphs gave the following measurements in mm. :
0-4 X 0-4 (:} specimens) 0-38 x 0-39

0-4 X 0-36

498 Greniis Haemaphysalis

G. F. Hill coll.; and 14 ^Z" 2 % 1 o taken from Pe7-ameles nasuta,
Sydney, New South Wales, III. 1913, L. Harrison coll. Mr Harrison
has kindly presented us with some of his specimens (N. 3024, 3025) ;
some of these $s and Os are fairly replete and their bodies appear
remarkably long and slender.
Types in Cambridge.

40. HAEMAPHYSALIS ELONGATA Neumann, 1897.

Figs. 438-440.

Lit. and Icon. : Neumann, 1897, pp. 354-356, Fig. 19 (reproduced) ; Donitz, 1907 a,
p. 73 (quotes Neumann) ; Blanchard, 1909, p. 150, Figs. 186, 187 (taken from
Neumann) ; Neumann, 1911 a, p. 113, Fig. 61 (condensed from Neumann, 1897,
with the same figure as before).

Male (Figs. 438, 439) : Remarkable because of the backward
protrusion of the body when replete ; the ventral festoons are folded
backward dorsally as shown in Fig. 438 ; in a fully distended specimen,
the body may project 06 mm. Scutum: long and narrow, about
2x1 mm.^ ; cervical gi'ooves broad, shallow, sub-parallel ; lateral
grooves well-marked behind a pseudo -scutum and including either
one or two festoons ; pseudo-scutum chiefly indicated by its more
uniform texture compared to that of the rest of the scutum,
which is diversified by raised longitudinal ridges, a median ridge or
carina being rather conspicuous ; punctations irregular, shallow, con-
fluent, not numerous. Capituluni : base much broader than long,
rectangular, with well-marked sharp cornua ; palps long, especially
article 2, which is thrice as long (dorsally) as article 3, its posterior
border being produced backward and hollowed to receive coxa I ;
a slight retrograde spine under article 3 ; hypostome 3 | 3, teeth small
and uniform. Venter : anal grooves ogival ; spiracle oval, slightly
pointed dorso-laterally. Legs: all the coxae bear a sharp internal
spur; in some specimens these spurs increase progressively fi-om I
to IV, in others the spur on coxa I is longest" ; a sharp ventral spur
on all the trochanters ; tarsus IV rather stout and narrowing rather
abruptly; pads long.

1 The scutums of our 5 <J measure respectively: 2-15x1, 2-1 x 1-2, 2x1-1, 1-9x1-2,
1-8x1 mm.

2 Compare our Figure 438 with that of Neumann (Fig. 439, here reproduced).

H. elongata

499

Fig. 438. II. elongata i (N. 670, gorged), dorsum .showing protrusion of abdomen
beyond the scutum, coxae and trochanters witli part of capitukim, spiracle, tarsus IV
and hypostonie. Original, G. H. F. N. del.

Fig. 439. II. elongata <j , {A) left palp in ventral aspect, x 100 ; {li) coxae I-IV with
trochanters, x 40. Neumann, 18'J7, Fig. I'J.

500

Genus Haemapliysalis

Female (Fig. 440) : When unfed, the body measures 2 x 1-2 mm.
Scutum : 1 X 0"8 mm.^ ; cervical grooves long and shallow ; faint, raised
longitudinal ridges in the lateral fields ; punctations faint, irregular,
shallow, confluent. The posteiior portion of the dorsum bears two
rather conspicuous depressions. Capituhmi : base broader than in the
(/" ; porose areas elongate, wide apart ; palps as in the (/", except that
the posterior border of article 2 is more rounded dorsally ; hypostome
3 I 3, 7 to 8 teeth per file. Venter : anal grooves as in the (/ ; spiracle
sub-circular, not pointed postero-laterally. Legs : coxal and trochantal
armature as in the ^.

Fig. 440. H. elongata ? (N. 670, unfed), dorsum, part of venter, spiracle and tarsus IV.
Original, G. H. F. N. del.

Our description is based on (N. 670) 5 </ and 2 $ from Gentetes
ecaudatus, Madagascar, presented by Hon. N. C. Rothschild. Neumann
founded the species on 4 cT and 1 $ found on Gentetes madagascariensis
at Zura, 2 </ from Gentetes ecaudatus and 2 $ from Erinaceus sp.,
collected by Sikora in Madagascar.

Types in Toulouse, co-types in Cambridge (N. 2884, 2885, 1 </,
fi:om Zura, 1 $ , taken from Erinaceus, presented by the author).

41. HAEMAPHYSALIS CORNIGERA Neumann, 1897.

Figs. 441-445.

Lit., Icon, and Synon. :

Haemaphysalis cornigera Neiuiiann, 1897, pp. 350-352,Figs. 16, 17 (left palp and
coxae; figures reproduced); Neumann, 1911a, j^p. 112, 113, Figs. 57, 58

1 The scutum of our second 9 measures 0-95 xO'85 mm.

H. cornigera

501

(the same account as the foregoing, but condensed) ; Blanchard, 1909, p. 150,

Fig. 185 (species merely listed ; figures taken from Neumann).
HaemaphysaUs spiniceps Warburton and Nuttall, v. 1909, p. 68, Fig. 15 {^

reproduced). Through a typographical error the measurements of the

cj are wrongly given.
Haeviapkysalis pro.vima Warburton and Nuttall, v. 1909, p. 61, Fig. 6 (9 not

reproduced).

Male (Figs. 441-444): Scutum long-oval, 3x2 inm., broadest near
the middle, glossy yellow-brown, darker at the sides and on the
pseudo-scutum, with large deep punctations anteriorly, some of them
confluent and indicating by their distribution a pseudo-scutum ; puncta-

Fig. 441. H. cornigera 3 , dorsum, part of venter, spiracle and tarsus IV.
from Borneo, co-type (N. 2883). Original, G. H. F. N. del.

Specimen

tions very small and numerous on the hinder portion ; cervical gi'ooves
well-marked, broad, nearly parallel ; lateral grooves almost obsolete, only
recognisable in the neighbourhood of the spiracles ; festoons medium,
punctate, the intervals dark and mostly straight. Cupitulum : ()"6 mm.
long ; base broadly rectangular, slightly punctate, with convex sides
and strong cornua ; hypostome long, parallel-sided, well-covered with
5 I 5 small teeth ; palps of remarkable form ; article 2 strongly salient
at about the middle of its length and with five spatulate hairs on its
inner margin ; article 3 with a strong, sJiarp lateral spine and with
a similar ventral retrograde spine directed downwards and outwards.
Venter: spiracle very large and short comma-shaped; anal grooves
slightly ogival. Legs : strong ; coxa I prominent in front and with

502

Genus HaemapTiysalis

a strong spur ; a conical spur about the middle of the posterior border
of coxae II and III ; two long spurs on coxa IV, the inner the longer;
tarsi tapering, pad long.

Fig. 442. H. cornigera j , capitulum in dorsal and ventral aspects, coxae. Specimen
from Federated Malay States. (N. 1647. Type of H. i^jiiniceps Warb. and Nutt.)
Sketch by C. W. Warburton and Nuttall, 1909, Fig. 1.5.

Fig. 443.

Fiix. 444.

Fig. 443. H. cornigera <? , left palp in ventral aspect, x 70. Neumann, 1897, Fig. 16.
Fig. 444. II. cornigera <? , coxae I-IV, x 35. Neumann, 1897, P'ig. 17.

H. cornigera

503

Female (Fig. 445) : SciUmn conlato, 1-3 x 1'2 mm., glossy,
uniformly punctate all over with moderate sized punctations ; cervical
grooves shallow. Capitulum: base rectangular, much broader than
long; porose areas oval, ill-defined, far apart, usually with a small
depression in the interval ; cornua strong. Palps : the absence of the
lateral spine on article 3 prevents any great resemblance to those of the
f^, but the lateral contour of article 2 is somewhat similar, and it is not
continuous with that of article 3 which is slightly salient laterally, a
strong retrograde spine under article 8 ; hypostome well covered with
4|4 strong equal teeth, 10 or 12 per file. Venter: anal grooves
semi-circular ; spiracle large, oval, with hardly any dorsal process.
Legs : a strong internal spine on coxa I ; a small spur at the middle
of the posterior border of coxae II and III and at the internal angle of
coxa IV ; tarsi long, tapering gradually ; pad as long as the claws.

Pig. 44-5. H. cornigera 9 , scutum, capitulum in dorsal and ventral aspects, coxae with
trochanters, spiracle, tarsus IV. Drawn from specimens from British N. Borneo
(N. 2119 h), the tarsus drawn from (N. 2118). Original, N. C. del.

Our description of the (/ is from a specimen (N. 1647) collected in
the Federated Malay States by Mr H. C. Pratt, which was described
by us as H. sjnnicejjs (Warburton and Nuttall, 1909, p. 68, Fig. XV,
here reproduced). We are indebted to Dr Oscar Pou for (N. 2104) 5 ^,
from a tiger or wild pig, and (N. 2105) a $ from a wild pig, shot in the
jungle at Kuantan, F'. M. S., i. 1913. We have also received numerous
specimens from British North Borneo : (N. 2118) from which we
describe the $, taken with many (/s from the >Sa7)ibu7' deer, Bode River,
Sandakan Bay, iv. 1913 ; (N. 2119 b) </ ? from a luater buffalo, Labuk
River, 10. xii. 1912; (N. 2120), 2 $, from buffalo, Mekong River, iv.
1913 ; the three lots were collected by Dr H. F. Conyngham. We have

504 Genus Haemaphysdlis

determined (Berlin Mus. No. 172) 1 $ from Darvel Bay, Dr Pagel coll.
A $ (N. 1251 a) has reached us from India; it was found on deer, at
Satharangapara, Travancore, I. 1911, by E. R. Hewlett. Neumann
(1897) described both sexes from specimens (2 </) found on Cerviis sp.
in Borneo, R. Blanchard coll., 1894 ; (2 cT 1 $ ) from Singapore, Simon
coll.; (1 (/) from Bos bubalis, Sumatra \ Oudemans coll.; (1 </) from
Judea, in the Paris Museum.

The types and co-types are in Toulouse (</ $ ), Paris Museum (</),
and Cambridge, Prof. Neumann having presented us with a co-type
(N. 2888) c/ from Cervus sp., Borneo.

The lateral spine on article 3 of the palp, and the double internal
spine on coxa IV, are such striking characteristics as to make the </ of
this species easily distinguishable. Unfortunately both these character-
istics are absent in the $ , which resembles the (/ in little except in the
lateral contour of article 2 of the palp and the break between it and
article 3, and the unusually large size of the spiracle.

Haemaphysalis cornigera var. anomala
Warburton, 1913.

Fig. 446.
Lit. and Icon. : Warburton, vii. 1913, p. 128, Fig. 7 (reprochiced).

Male (Fig. 446) : Closely resembles the type except in the following
respects : the Srd article of the jKilps not projecting laterally (hence
anomala) ; basis capituli somewhat narrower compared with its length ;
spiracle more pointed dorso-laterally.

Female : unknown.

Fig. 446. H. cornigera var. anomala 3 , dorsal and ventral aspects of capitulum.
N. C. del. Warburton, 1913, Fig. 7.

^ A ? (N. 540), described by us as H. proxirna N. and W., 1909, p. 61, is now referred
to this species; the specimen was taken by Dr J. C. Koningsberger, from Bos bubalis, in
1909 in Sumatra.

H. coniigera var. anomala and H. dentipalpis 505

The most remarkable characteristics of H. cornigera — the presence
of two divergent spines under article 8 of thi- palps, and of a double
spine on coxa IV — are repeated in this variety.

Described from a single (/ taken from a wolf, four miles west
of Koderma Station, Chota Nagpur, India, viii. 1912, by Major
O, A. Smith. In its compjiny were specimens of H. bisjnnosa var.
intermedia, H. leacJii and Rhipicephalus haemaphysaloides:

Type in the Indian Museum, Calcutta (No. 1524/17 a).

42. HAEMAPHYSALIS DENTIPALPIS Warburton
and Nuttall, 1909.

Fig. 447.

Lit. and Icon. : Warburton and Nuttall, 1909, pp. 67-68, Fig. 14 (reproduced).

Male (Fig. 447) : Body long-oval, 27 x 1-3 mm., broadest towards
the hind end; colour, dull brownish-yellow, the front legs, the capitulum
and the anterior part of the scutum darker. Scutum: showing con-
spicuous pseudo-scutum, with numerous large, deep punctations ; very
fine punctations on the anterior portion ; cervical grooves deep and
convergent, then faint and divergent and visible almost to the posterior
border of the pseudo-scutum ; lateral grooves short, beginning faintly

Fig. 447. H. dentipalpis <? , capitulum in dorsal and ventral aspects. Sketch by C. W.
Warburton and Nuttall, iy09, Fig. 14.

506 Genus Haemcq)lnj salts

at about half the body-length, and including two festoons ; festoons
rather short and broad, with straight intervals. Capitulum : OG mm.
long ; base broader anteriorly, with convex sides and rather concave
anterior border ; cornua strong, rather curved ; hypostome long, parallel-
sided, well covered with 7 | 7 very small equal teeth ; palps long, conical,
their dorsal surface concave ; article 2 much longer than 3 and with
five retrograde projections, two dorsal and three ventral, best under-
stood by reference to the figures; three simple hairs on the internal
margin of article 2 ; a strong retrograde spine under article 3. Ventei':
spiracle very large, pear-shaped, with the narrower end dorsal ; anal
grooves ogival. Legs : long, hairy, coxae like those of H. leachi (p. 468,
Fig. 409), svith a short conical spur at each postero -internal angle ;
tarsi rather thick, tapering abruptly.
Female : unknow-n.

Described from 4 (/ taken (in company with H. leachi) from Felis
ckaus, by Dr A. T. Stanton in the Federated Malay States.

Types in the London School of Tropical Medicine collection and
(N. 693 ft, 2 c/) at Cambridge.

Geo(jrapliic(d Distribution aiul Hosts 507

GEOGRAPHICAL DISTRIBUTION AND HOSTS
OF THE GENUS.

Of the species and varieties we recognize, -i (jcciir in Europe, 15 in
Africa, 32 in Asia, 4 in Australasia and 3 in America. //. hispiiiosa
has ahnost to a certainty been recently imported into East Africa and
possibly into New South Wales with Indian cattle ; //. leachi may well
have reached Australia (again New South Wales) from India. If
we regard these species as not indigenous to the contin(,'nts mentioned,
the number of indigenous species would be as fi)llo\vs: Europe; (3),
Africa (14), Asia (32), Australia (2), America (3).

This is in marked contrast to what we observe in the genus Ixodes.
If we add some species described by Bishopp (1911, pp. 197 et seq.),
Nuttall (1918, pp. 131-136), and Lahille (1913, p. 2) to those listed in
Part II of this work, varieties being excluded, we find the species
of Ixodes which we recognize as valid to-day distributed as follows:
Europe (10), Afiica (13), Asia (10), Australia (10), America (25), It is
clear therefore that Asia is richest in species of Haemaphy salts and that
America possesses the greatest number of Ixodes.

There are three species of Haemaphy salis which show a particularly
wide distribution : (1) H. cinnaharina occurs in North and South
America ; its var. jmnctata is found in Europe, Northern Africa, Asia
Minor and Transcaucasia ; (2) //. bispinosa occurs in Asia [British
East Africa and New South Wales] ; (3) H. leachi occurs practically
throughout Africa, in different parts of Asia [and in New South
Wales]. The continents where these species are perhaps not indi-
genous are enclosed in [ ] brackets.

Regarded from the point of view of the hosts which they infest, we
find but one species of Haemaphy salis which, as far as our knowledge
goes, would appear to be restricted to birds, i.e. H. hoodi.

This is again in marked contrast to what we have observed in Ixodes
where certain species have been repeatedly found only on birds
(/. hrunneus, I. caledonicus, I. neuvianni, I. patas, I. signatus, I.U7iicavatus),
on Ornithorhynchus (I. ornithorliyvchi) and bats (/. vespertilionis). In
this sense, the genus Ixodes apjjears more specialized than Haema-
phy salis.

In the list which follows, the scientific names of the hosts are given
as far as possible from the data at hand.

508

Genus Haemcqjhy sails

EUROPE

cinnabarina var.
punctata

Countries where the species
occur

Great Britain, France, Hol-
land, Denmark, Germany,
Eussia, Italy, Spain, Hun-
gary, Croatia, Dalmatia,
Eoumania, Greece, Cy-
prus, Cyclades, Crete

concinna

Germany, trance, Poland

inermis

Caucasia, France

AFRICA

aciculifer

Uganda, Gold Coast

hispinosa

British East Africa

calcarata

Abyssinia, Egypt, Soudan,
Somaliland

calcarata var.

houy

New Cameroon

cinnabarina var.

Algeria, Egypt, Canary

punctata

Islands

elongata

Madagascar

hoodi

West : Gambia, Gold Coast,

hoodi var. orientalis
leachi

Sierra Leone
East : Portuguese East
Africa, Nyasaland

Nyasaland

Practically everywhere in
Africa (refer to text pp.
469-474)

Hosts

Mammals: Uvis aries, Bos taurus,
Capra hircus, Cervus daina, Equus
cabalkis, Lepus europaeus, Erl-
naceus europaeus. Bat, Plecotus
auritus

Birds : Turdus viscivorus, partridge,
Numenixis arcuatus

Beptiles : Vipera aspis, Lacerta spp.
(o. & L.)

Ovis aries, Cervus sp., Erinaceus
europaeus

Cervus sp.

Antelope, reed-buck

Bos taurus, bullock, Capra hircus

Spermosciurus sp., Sciurus sp., Xerus
rutilus

unknown

Mammals : Ovis aries, Capra hircus,

Bos taurus
Reptiles : Lacerta sp.

Centetes ecaudatus, C. madagascar-
iensis, Erinaceus sp.

Birds : Centropus burchelli, C. sene-
galensis, Gymnoschizorchis leo-
poldi, Gallus domesticus, Nuviida
meleagris, partridge

Procavia manningi

Mammals : Man (rarely)

Caruivora (chiefly): Canisfami-
liaris, C. adustus, C. mesomelas,
Felis leo, F. pardus, F. nigripes
F. serval, F. domesticus, Que-
parda jubata, bushcat, Viverra
civetta, Genetta sp., G. suahelica,
Herpestes sp., H. caffer, Cynictis
penicillata, Suricata tetradactyla,
Helogale varia Thos., Hyaena
sp. , H. crocuta, Mustela sp.

Ovis aries, Capra hircus. Bos
taurus, Bos indicus, Equus cabal-
lus, Ehinoceros, Procavia brucei,
P. abyssinica, Tachyoryctes audax

Geographical Dit>tribntion and Hosts

509

AFRICA [cont.)

Countries where the sjjeeies
occur

ASIA

numidiana

Algeria

obtusa

Island of Eeunion

parmata

Uganda, British East Africa,

Congo Free State, Gold

Coast, Sierra Leone,

Cameroon

silacea

Cape Colony

simplex

Madagascar

spinulosa

Uganda

A

aburcnsis

India (Abor Country)

aculeata

Ceylon, India

birmaniae

Burma

bispinosa

India (widely distributed).

bispinosa var. inter-
media

calvus
campanulata

cinnabarina var.
punctata

Burma, Assam, Ceylon,
Andaman Islands, Malaya,
China, Jaj^an, Borneo

India (widely distributed),
Ceylcjn

British North Borneo
India, China, Japan

Asia Minor, Transcaucasia
(? Japan, doubtful, see
text p. 387)

(mole-rat), Avicanthus jyvmillis
(os.), Anonuilurus orientalis
(squirrel), Lepus sp., Erinaceus
albiventris, Erinaceus sp.

Birds: Drijoscopns turalii and
nightingale

Beptiles: Itonyx capensis, Testudo
sp. (os.)

Erinaceus sp.

unknown

I'olaiiioclioerus porcus, Ovis arics,
Capra hircus, Bos taurus, antelope,
bushbuck, hartebeeste, Canis
familiar is

Bos tawus

Ericulus setosm, Erinaceus sp.

unknown

unknown

Tra(julus nwiiunina

Cervulus muntjac, Atlierura iiuicrura

Mammals: Canis jamiliaris, jackal,
Felis tigris, F. domesticus, Vivcr-
ricula mulaccensis, Bos bubalis,
B. taurus, Hemitraijus hylocrius,
Nemorhaedus cinereus, Ovis aries,
Capra hircus, Eqnus caballus,
Sus scrofa, Cervulus sp., Cervus
sp., mouse deer, Hijdropotes in-
ermis, Macacus sp., Talpa sp.

Birds: Gallus domesticus

iS'^aramals: Felis caracal, F.ornata,
F. domesticus, F. ajinis, Caitis
pallipes, C. vulpes, Cyon [Cuon)
dukhunensis. Hyaena hyaena. Bos
taurus, 5ws cristatus, Erinaceus
micropus, Lepus ruficaudatus, Mil-
lardia meltada

Bos bubalis

Bos taurus, Equus caballus, Cervus
sp., Canis familiaris, Mus sp.

Bos taurus, Ocis aries, Capra hircus,
Flquus caballus, Canis vulpes,
Ursus sp. , Lepus sp.

33

510

Genus Haemaphysalis

ASIA {cont.)
corniijera

cornujera var. ano-
mala

Countries wliere the species
occur

India, Federated Malay
States, Singapore, Bor-
neo, Sumatra, Judea

India

cuitpidata

Ceylon

dentipalpis

Federated Malay States

doenitzi

Singapore

fiava

Japan, India

formosensis

Formosa, Burma

howlctti

India

hy.stricts

Burma, Assam, Cey

China, Federated Malay
States, Sumatra, Borneo,
Formosa

inermis

Transcaucasia (? Japan)

inermis var. aponom-

India, Japan

vioides

japonica

Japan, China

japonica var. doag-

China

lasi

kinneari

India

leachi

Transcaucasia, India,

matra, Java, Borneo,
Federated Malay States

montgomeryi

India

Hosts

Cervus aristotelis, Bos bubalis, Felis
tigi-is, wild pig

Cants liq)us

Tragulus memmina, Pnradoxuriis
nigcr, Herpestes mungo

Fclis chaus

water-hen

Cants familiaris, Siis cristatus, Bos
taurus, Ovis aries, Eqiius caballus

Canisfantiliaris, Hystrix hengalensis
or Ursits torqnatus

Eqntis caballus

Mammals : Hystrix bengalensis,
Erinaceus sp., Ursits torquatus,
U. malayamis, Felis tigris, Ganis
familiaris, Sus scrofa, S. celeben-
sis

Reptiles : Geoemyda sjnnosa

Bos tauriis, Naemorhaediis crispus,
Canis sp. (fox)

Bibos sp. (Himalayan zebu), Eqttus
caballus

Naemorhaedus crisjnis, roebuck

roebuck

Felis tigris

Mammals (chiefly Carnivora) : Ursus
sp., U. malayamis, Felis affinis,
F. bengalensis, F. chaus, F. ne-
bulosa, F. tigris, F. caracal, F.
pardus, Viverra zibetha, Viverra
sp., Canis aureus, C. indicus, C.
familiaris, Cyon (Cuon) dukhu-
nensis, Paradoxuriis sp., Herpestes
mitngo, Centrococcyx intermedins,
Millardia meltada

Reptiles (Testudiniae) : Nicoria tri-
juga, Testudo elongata

Equus caballus, Bos taurus, Ovis
aries, Canis familiaris

I

^

Geographical Distrlbulioit and Hosts

511

ASIA (cunt.)
papuaiia

Countries where the 8i)euie8
occ in-

New Guinea, Borneo, Java,
Sumatra, Federated Ma-
lay States

parva

Ceylon, India

spiiiiijera

India. Ceyluii, Judca

spinigeni var. mivtu'- German Now Guinea,

gitineae

Queensland

Uirturis

Ceylon

vidua

Federated Malay Status

icarburloiii

China, Western Siberia,

Formosa

well'uKjtoni

Borneo, Sumatra, Federated

Malay States, Siani, Anda-

man Islands

AUSTRALASIA

bancrofti

Queensland

bispinosa

New South Wales

humerosa

Australia

leachi

New South Wales

AMERICA

cinnabariua

United States, Canada

kochi
leporis-palustris

Brazil

United States, Canada,
Mexico, Panama, Brazil,
Paraguay, Argentina

Hosts

Gania fdmiliarin, Uraus mahiyanns,
Felis thjris, F, pardtis, Sux sp.

Felis viverrina, F. pardus, Lepns
I'uficaudatus, Lepus sp.

Felis tigrh, F. pdrdus, Uisus sp. ,
bullock

Kangeroo sp., I'cramclvn sp., Hydro-
mys sp.

Turtur suratemis

Paradi)xiinis sp.

Nemorhaedus (Serow goat). Bos
tauriis

Mammals : Canis familiaris. Bos

bubalis, but oftener from
Birds: fowl, goose, turkey

Macropus dorsalis, M. yiyanteus,
Bettonyia penicillata

Equus caballiis, Bos taurus

Ferameles iiuicrttra, P. nasuta

Equus caballus

Birds (chieHy) : Cliordeilis popetiie,
turkey, meadow lark, jackdaw,
red-winged blackbird, marsh-
hawk, quail, ruffed grouse (in
the United States)

Mammals: Bos taunts, Man (in
Canada)

Cervus campestris, C. ntfus

Mammals (chiefly Leporidae) : Lepus
americanus, L. auduboni, L. bairdi,
L. braziliensis, L. dalli, L. jyalus-
tris, Dasyprocta agouti, Eqttus
caballus, Felis domesticus, pine
squirrel

Birds: robin, quail, meadow lark,
chaparal cock. Brewer's blackbird,
thrush, jackdaw, blue jay, magpie,
Penelope obscura (in the United
States)

33—2

512 Genus Haemaphy sails

LIST OP

CONDEMNED AND DOUBTFUL SPECIES
OF HAEMAPHYSALIS

INCLUDING THEIR SYNONYMY AND LITERATURE

The .species are ordered according to the genera to which they have at various
times been referred. The list of good species will he found on p. 548. See also
further particulars under the Synonymy of Hae ma ph// salts on p. 349.

Haemaphysalis Koch, 1844.

afrimna C. W. Howard, viil. 1909, repr. PL XXXIV = //. hoodi Warburton
and Nuttall, vi. 1909, p. 62.

amhigua Neumann, 1901, p. 262 ; 1906, p. 217 ; 1911 o, p. 109 ; Bonnet, 1908,
p. 260, Fig. 30 ; Blanchard, 1909, p. 148, Fig. 178 (from Bonnet) = ^.
inennis Birula, 1895, p. 360.

asiatica (Supino, 1897) Neumann, 1897, p. '357 = 11. leachi (Audouin, 1827). See
further under Opisthodon asiaticus. Listed as a valid species by Blanchard,
1909, p. 148, and as a doubtful species by Neumann, 1911 a, p. 115.

" birmaniae Supino, 1897 " in Neumann, 1911 a, p. 109 = ir. bispinosa Neumann,
1897.

"bispinosa Neumann" in Neumann, 1901, p. 261 = 11. hystricis Supino, 1897.

caHes^/'iMU (Supino, 1897) Neumann, 1897, p. 'ibl = H. leachi (Audouin, 1827)
Neumann, 1897 ; Blanchard, 1909, p. 148, lists it as a valid species ;
Neumann, 1911 a, p. 115, lists it as a doubtful species.

chordeilis (Packard, 1869) = //^, dnnabarina Koch, 1844.

'■'■ chordeilis Banks, 1908" in Blanchard, 1909, p. 148 = i/. cinnahanna Koch,
1844.

'■'■chordeilis Packard" in Hunter and Bishopp, 1911, p. 229 (and subsequent
American authors) =Zr. dnnabarina Koch, 1844.

^'' cinnaberina" Koch, 1847, p. 123=^. dnnabarina Koch, 1844, p. 237.

"cocdnea" mentioned casually by Bonnet, 1906c, p. 544, doubtless meant for
IT. condnna Koch, 1844.

'■'• condnna condnna Koch" in Neumann, 1911 «, p. 110, a sub-species = i:f.
concinna Koch, 1844.

condnna kochi (Neumann) Neumann, 1911 «, p. Ill, a sub-species = if. con-
dnna Koch, 1844, and other (doubtful) species.

concinna. var. kochi Neumann, 1905, p. 239 = ff. condnna Koch, 1844, and other
(doubtful) species.

condnna longicornis (Neumami) Neumann, 1911 «,}). Ill, a sub-species ; described
as H. longicornis Neumann, 1901, p. 261, Fig. 2 (reproduced) from 2 $
fomid on cattle, at Kempsey, N. S. Wales, mounted as microscopic speci-
mens ; degraded to a variety of H. condnna Koch, 1844 (//. concinna var.

Condemned Species

513

longicornis (Neumatui)) by Neumann, 1905, j). 237, and finally raised by
that author to a sub-speuies. It is in our opinion impossible to determine
the species to which the tick belongs on the evidence of mounted speci-
mens, at any rate without other confirmatory evidence. H. mncinna 9
is scarcely to be identified in the absence of the characteristic ^, and
a purported 9 > coming from a remote country whence the ^ has not
been derived, is open to grave suspicion. Wa re[)roduce Neumann's figure
for what it is worth.

Fig. 448. H. iDiigicoriiis Neumann, 1901, Fip. 2. ? (P) palp in ventral aspect,

(//) coxa I.

JI. longicornis Neumann is listed by Rainbow, 1906, p. 165, as an Aus-
tralian species on Neumann's authority. Blanchard, 1909, p. 148, Fig. 184,
who reproduces Neumann's figure, inchides this tick in the synonymy
of H. concinna Koch. We prefer to regard it as a doul)tful species.

concinna var. longicornis (Neumann) Neumann, 1905, p. 237 ; ride supra.

crassa Warburton, 1908, p. 516 ; listed as a valid species by Blanchard, 1909,
p. 150 = ^. cinnaharina var. punctata (Can. and Fan.).

cnscobia (Canestrini, 1897) Neumann, 1897, p. 356 = merely a nominal species.
See further under Opisthodon cuscolmcs. Listed as a valid species by
Blanchard, 1909, p. 150, and as a doubtful species by Neumaim, 1911 a,
p. 115.

erinacei Pavesi, 1884, p. 484 ; merely a nominal species, the description being
insufticient. We quote from the original all that might serve for its
identification : ^Ualc : 3 x 2 mm., finely j^unctate, cervical grooves divergent,
lateral grooves present, sexual orifice facing coxae II, coxae imarmed.
Female: scutum 1 mm. long, broadening and rounded behind, anterior
borders parallel. — Found by Marquis Doria on Erinaceus algirus at
Gebel Resas, Tunisia, in 1881. Pavesi consid(>rs the tick allied to H.

514 Genus Haemaphu sails

concinna ; it may be identical with this species as Neumann states (1897,
p. 331). Neumann, 1909, p. 151, and 1911 (/, p. 116, lists the species
as doubtful.

flava as listed by Neumann since 1897, p. 333 = a confusion of species which
may have included H. flava Nn., H. campanulata Warburton, and //.
japonica Warburton, q.v.

flava armata (Neumann) Neumann, 1911 «, p. 112, a sub-species = Zr. ^aya
Neumann, 1897, p. 333.

flMva var. armata Neumann, 1905, p. ^2'il = H. flava Neumann, 1897, p. 333.

flava flava (Neumann) Neumann, 1911 «, p. 112, a sub-species = iT. japonim
Warburton, 1908.

gestroi (Supino, 1897) Neumann, 1897, p. Zbl = H. leachi (Audouin, 1827).
Neumann, 1902, p. 128, regarded the species as allied to H. leachi;
Blanehard, 1909, p. 151, lists it as valid ; Neumann, 1911a, p. 115, lists
it as a doubtful species.

hirudo L. Koch, 1877 a, p. 786, not figured. Undeterminable from the author's
description ; he may have been dealing with a HaemaphysalU ; a $ from
Japan. Neumann, 1897, p. 341, at first referred specimens of his own
to Koch's species because they also came from Japan, but afterwards
renamed them H. concinna var. kochi Neumann, 1905, p. 239, realizing
that in the absence of the type, the species hirudo was purely nominal.
We fully agree with Neumann with regard to the foregoing but do not
recognize his var. hochi as valid (see text p. 456). H. hirudo is listed
as a doubtful species by Blanehard, 1909, p. 151, and by Neumann, 1911 a,
p. 116.

" hystricis Supino " in Neumann, 1902 a, p. 128, and 1911 «, p. 109 = //. bispinosa
Neumann, 1897, p. 341.

japonnica Warburton, 1908, p. 512. Name misspelt; should read H. japonica.

lagotis (Gervais) of Neumann, 1901, p. 265 = merely a nominal species, for
Ixodes lagotis Gervais, 1849, whilst it may have been a Uaemaphysalis,
was too poorly described to make this certain. Listed as a nominal
species of Haemaphysalis by Blanehard, 1909, p. 152, and as a doubtful
species by Nevunann, 1911a, p. 116.

leachi australis (Neumann) Neumann, 1911 a, p. 115, a sub-species = ^. leachi
(Audouin, 1827).

leachi var. australis Neumann, 1905, p. 238 = //. leachi (Audouin, 1827).

leachi leachi Neumann, 1911a, p. 114, a sub-species = /T. leachi (Audouin, 1827).

" leporis Packard " in Rohr, 1909, pp. 144-146 (quotes Neumann's description)
= if. leporis-palustris (Packard, 1869).

"•leporis (Packard)" in Nemnann, 1897, p. 343; 1901, p. 262 and 1911 a, p. Ill
= H. leporis-palustris (Packard, 1869).

leporis var. proxima Aragao, 1911, p. 167, PL XI, Figs. 4, 5 {^) = H. leporis-
palustris (Packard, 1869).

'•'■leporis-palustris Packard "in Hunter and Bishopp, 1911, p. 228 = //^. leporis-
palustris (Packard, 1869).

longicornis Neumann, 1901, p. 261 ; a doubtful species, see discussion above
under H. concinna longicornis (Neumann).

Condemned SjJecles 515

longipalpis Warburtoii, xir. 1910, p. 399=//. anuleata Lavarra, 1905, p. 255.
marmorata Berlese, 1887, Fasc. 47, No. 4, PI. II {$) = Dermacentor reticulatus

(Fabricius, 1794).
micropla C'anestrini, 1887, p. 104, IM. IX, Figs. 3, 5; also in Berle.se, 1888
= Rhipicephalua microplus (Canestrini) Canostriiii, 1890, \>. ^^9^ = Boop/dlus

annulutus var. microplus according to Neuiuauu, 1901, p. 280.
Tieumanni Donitz, 1905, pp. 127-129, Figs. 4-6 = //. 6ispino«« Neumann, 1897,

p. 341.
neumanni Douitz in Blanchard, 1909, \t. 154 (merely listed); in Neumann,

1911«, p. 109; I in Yakiraoff and Kohl-Yakimoft; 1911, p. 418 = /r.

hispinosa Neumann, 1897, p. 341.
'"'■ peregrina Cambridge," listed as a doubtful sjieciesby Neumann, 1911 (/, p. 116.

See under peregrinus.
'■^ per egr huts Cambridge " in Neumann, 1897, p. 327 ; see below.
peregrinus O. Pickard-Caml)ridge, 1889, p. 406, 2 figures. Description and

figures useless ; purely a nominal species. The author informed us that

the types are lost. Included by Neumaiui, 1897, p. 327, in the .synonymy

oi H. punctata { = cinnabarina vav. pimet'it<t) without sufficient reason.
proxima Warburton and Nuttall, v. 1909, pp. 61, 62, Fig. 6=/r. comigera

Neumann, 1897, $ .
"proxima Aragao " in Rohr, 1909, p. 146, PI. I, Fig. 1(9), PI. II, Figs. 12-14;

Hooker, Bishop}) and Wood, 1912, p. 89 (merely cite Rohr) = //. leporis-

palustris (Packard, 1869).
punctata Canestrini and Fanzago, 1877, p. 188; repr. p. 120 (and other authors

since) = ^. cinnabarina var. punctata (Can. and Fan.) Nutt. and Warb.
ptaictata Can. and Fan. in Iladwen, 1912, p. 98 = JI. cinnabarina Koch, 1844.
punctata cinnaberina (Koch, 1844) in Neumann, \9\\a, p. 108, a sub-species

==H. cinnabarina var. punctata (Canestrini and Fanzago, 1877).
punctata var. cinnaberina (Koch) in Neumann, 1905, p. •1Z~ = II. ciiuKtUarina

var. punctata (Can. and Fan.).
punctata punctata Canestrini and Fanzago, 1877, in Neumann, 1911 a^ p. 107,

a sub-species=/r. cinnahtrina \ -ax. punctata (Can. and Fan.),
rhiuolophi Canestrini and Fanzago, 1877, p. 189, repr. p. 121 ; Canestrini, 1890,

p. 526 = H. rin/iabarina Kf)ch, 1844. Listed as a valid species by

Blanchard, 1909, p. 157, and as a doubtful species by Neumann, 1911a,

p. 116.
rosea Koch, 1844, p. 237; 1847, p. 121, PI. XXVI, Figs. 95, 96 ( $ ) = probably

Boophil'us annulatus (Say, 1821) judging from the original de.scription and

figures. Neumann, 1897, p. 408, and 1901, p. 276, al.so refers rosea to that

species. In the absence of the type the synonymy must remain in doubt.
" rostralis Dug5s" in Banks, 1908, p. 54 (listed) = partly //. leporis-palustris

(Packard, 1869) $ . Refer to the synonymy of the latter species in our

text p. 387 and under (Jonixodes, p. 516.
sanguinolenta Koch, 1844, p. 237; 1847, p. 124, PI. XXVII, Fig. 48 = //.

cinnabarina Koch, 1844. Type (?) examined by Nuttall in 1911. Listed

as a valid species by Blanchard, 1909, p. 157, and as a doubtful species

by Neumann, 1911a, p. 116. Rohr, 1909, p. 146, merely copies from

516 Genus Haemaphy salts

Neumann (1897, p. 332) and calls the purported species ^'' sanguinolenta

Kock" {sie).
semermis Neumann, 1901, p. 263 {^) = H. htjstricis Supino, 1897. Nuttall

examined the type.
spiniceps Warburton and Nuttall, 1909, p. 68 {^) = H. corniyera Neumann,

1897, p. 350.
'''■sulcata Koch" in Caneatrini and Fanzago, 1877, p. 189; reprint, p. 120 ; also

in Berlese, 1889. Listed as a doubtful species by Neumann, 1911 a, p. 117.

See further under the synonymy of H. cinnaharina var. punctata (Cane-

strini and Fanzago, 1877).

Gonixodes Dugfes, 1888.

rostralis Dugfes, 1888, p. 129, Fig. 2 (supposed ^) = H. leporis-pahistris (Packard,
1869) 5 i'l Neumann, 1897, p. 343, whose attribution is probably correct,
judging from Duges' figure 2 ; Figs. 1 and 3 (the supposed larva and
nymjjh) belong to another genus. See further luider synonymy of H.
Jeporis-palustris. Type : G. rostralis.

Herpetobia Canestrini, 1890.

sulcata Canestrini, 1890, pp. 486, 493, 527, PL XLI, Fig. '6 (adult, n3anph, and
\i\rya) = " JUaemaphysalis sulcata Koch" in Canestrini and Fanzago, 1877,
p. 189 = probably H. punctata Can. and Fan., 1877 as Neumann, 1897,
pp. 327, 329, supposes = /r. cinnaharina var. punctata (Can. and Fan.).
Tyjje : Herpetohia sulcata.

Ixodes Latreille, 1795.

chelifer Mdgnin, 1880, p. 132 (^) = H. concinna Koch, 1844. Wrongly included
by Canestrini, 1890, p. 526, in his synonymy of H. punctata { = 11. cinna-
harina var. punctata (Can. and Fan.)), as fii'st jjointed out by Neumann,
1897, p. 338.

chordnilis Packard, 1869, p. 67 = H. cinnaharina Koch, 1844; not H. leporis
(Packard) as stated by Neumann, 1897, p. 343.

lagotis Gervais, 1849, p. 49 = merely a nominal species, the description being
valueless but for the statement that the second palpal article is " dihxtato
a modo de espina." Stated to occur in the ears of Lagostomus. viscaccia.
See above under Haemaphy salts lagotis (Gervais) Neumann, 1901. Lahille,
1905, p. 44, notes in this connection that L. viscaccia of the Pampas does
not live in Chili, and that Gervais' record probably refers to L. peruanum
(Meyen) which occurs in the Sierras and in Argentina. Lahille has failed
to find any ticks as yet upon L. viscaccia in the Pampas.

leachii Audouin, 1827, p. 4-28 = 11. leachi (Audouin, 1827) Neumann, 1897,
p. 347.

leporis-palustris Packard, 1869, p. Q1 = H. leporis-pahistris (Packard, 1869).

testudinarius Murray, 1877, p. 192, which Murray regards &s = Ixodes inarginatiis
Leach, is referred to the .synonymy of H. punctata { = cinnaharina var.
punctata Can. and Fan.) by Neumann, 1901, p. 260. This author bases
his conclusion on a wretched figure of Murray's which in our opinion
affords no evidence i-egarding the nature of the supposed species.

Condemned Species 517

Opisthodon Canestrini, 1897.

tisidtifus Supino, 1897a ; 1897 6, p. 2h-2, IM. XI II, Fig. -li^H. harhi (Audouin,
1827). See further under .synonymy of H. learhi.

canestrinii Supino, 1897 «; 1897 6, p. 252, PI. XIII, Fig. -21 = JJ. leachi
(Audouin, 1827). See further under synonymy of //. leachi.

cuscohius Canestrini, 1897, p. 468 =Zr. cuscobia (Canestrini) Neumann, 1897
p. 356. Described by Canestrini from one ^ of which he gives the
following characters : Size 2-0 X 1 "1 mm., hypostome 5 | 5, tarsi '.i and 4 witli
two spurs; found by L. Biro on Citscus sp., at Friedrich-Wilhelmshafen,
New Guinea. This, the ti/pe of the author's genus Opisthodon, has been
lost. The species is purely nominal.

gestroi Supino, 1897, p. 252, PI. XIII, Fig. 23 = ff. leachi (Audouin, 1827). See
further under synonymy of II. leachi.

Prosopodon Canestrini, 1897.

cuscobius (Canestrini) Canestrini, 1897, y>. 417 (footnote) = /T. cuscobia (Cane-
strini) in Blanchard, 1909, p. 150. See above under Opisthodon the generic
name which the author found was preoccupied and for which he substituted
Prosopodon.

Rhipicephalus Koch, 1844.

ellipticus Koch, 1847, p. 135, PL XXX, Fig. 111. Species based on one 9
from the Cape of Good Hope. Judging from the description and figure,
we may safely conclude that the tick was H. leachi {A.\\do\\n\, 1827); we
agree in this with Neumann. See also under Rhipistoma.

expositicins L. Koch, 1877 6, p. 196 = most probably E. ciimabarina var.
punctata (Can. and Fan.), judging from the description.

Rhipistoma Koch, 1844 (misspelt " Rhipidostoma " by Karsch, 1878, and Dugfes,

1888, and " Rhiphistoma" by Osborn, 1896).
ellipticum Koch, 1844, p. 23d = Rhipicephalus ellipticus Koch, 1847, p. 135 (.see

above) = Zr. leachi (Audouin, 1827).
leachii in Koch, 1844, p. 239 (cites Savigny), and in Karsch, 1878, p. 337 = //.

leachi (Audouin, 1827). The type of the genus Rhipistoma.
leporis in Osborn, 1896, p. 261=^. leporis-palustris (Packard, 1869).

518 Genus Haemaplufsalis

NOTES ON THE BIOLOGY OF HAEMAPHYSALIS

By G. H. F. NUTTALL.

Referring to the list of 50 species and varieties enumerated in the
table at the end of this fasciculus, the table serving likewise as an
index, the reader will find that the various stages of each species and
variety known to science give the following figures : The

cT $ o Z are known in 14 species and varieties.

cf ? O „ „ „ 4

c/ O

c/ only

18
1
8
5

It follows that these species and varieties are known by 45 </ , 41 $ ,
19 o, and 14 Z.

By referring to Part II, p. 295, it will be seen that 51 species and
varieties of Ixodes are known by 21 cT, 48 $, 22 o and 14 Z. These
figures show that a marked difference exists between the two genera.
The number of c/s of Haetiufphysalis which are known is slightly
greater than that of the $s, this being evidently due to the habit
of the (/"s to remain upon the host which the $ s have abandoned ; this
habit is also indicated by certain field observations on H. cinnabarina
var. punctata, H. concinna and H. inermis, where (/s only have been at
times found upon the host. In Lvodes on the other hand, some c/s do
not attack the host (see Part II, pp. 334-345), and this accounts for the
preponderance of $s that are known to science, although the cfs of
some species frequently occur in copula upon the host and abandon the
latter together with the $ .

Reference to pp. 507-511, where we have listed the hosts upon
which Hnemaplnjsalis occur, seems to show that the members of this
genus are less .specialized in respect to hosts than are Ixodes.

The life-history, in Haeniaphyscdis, appears to agree in the main
with that outlined for Ixodes (see Part II, p. 295) but it is impossible
to generalize since we only know the life-history of six species and
varieties of Haemaphysalis, and in H. inermis we have a species whose
life-history, in the present state of our knowledge, is unique amongst
the Ixodoidea.

Biologfi of H. cmnaharina var. punctata 519

In the following pages an account is given of all that i.s known
regarding the biology of six species :

PAGE

1. //. ciiinahurinn var. piiitctnta . . . 519

2. H. cinnabarina ...... ;J2H

3. H. leporis-palustris ..... .j30

4. H. leachi 53G

5. H. concinna 542

6. H. incrmis ....... 545

of which numbers 1 and 4 have been proved to be the carriers of
pathogenic Protozoa.

Haemaphysalis cinnabarina var. punctata^

The only authors who have occupied themselves with the biology of
this species are the writer (1908) and Stockman (1911, pp. 23-82);
a detailed study of the external anatomy has been published by Nuttall,
Cooper and Robinson (1908). The hosts upon which the tick is found are
given on pp. 508, 509. The species was first raised by me on hedgehogs,
this animal having been found to serve as a host in nature. The rabbit
and sheep were also used as hosts. The tick occurs most commonly
on sheep, and between 1902 and 1905 we received large numbers from
Kent, especially from the districts surrounding Lydd and Canterbury.
We have also received specimens found on goats and ferrets, but we
have no record of its occurrence on cattle in England. It is interesting
therefore that McFadyean and Stockman were able to transmit British
redwater, due to Piroplasina divergens, to cattle by means of this tick,
although it can but play an unimportant part in transmitting the
disease in Europe, Ixodes ricinns being ceilainly the chief vector.

Stockman raised H. cinnabarina var. punctata upon the scrotum
and ears, using cattle and sheep as hosts, the usual method of placing
bags about the scrotum and ears being employed to recover the ticks as
they dropped from the host. None of Stockman's ticks were incubated
during metamorphosis; they were maintained in corked bottles and
glass dishes in an unheated outhouse, moulds being avoided by keeping
the ticks somewhat drier than under natural conditions. The results of
his raising experiments are fully referred to and incorporated in the
following pages, the author's name being given in all cases where he is

' Reprinted (with slight modifications and certain additions duly indicated) from
Nuttall, IV. 1913, pp. 99-105.

520 Genus Haemaphy salts

cited. Stockman's records were somewhat confused and it was not
always easy to extract the desired information from them.

Seasonal occurrence on hosts. The specimens which have reached
us have been adults collected in April. According to Stockman, in
Kent and Devon, engorged females are found on sheep in April-June,
in October and occasionally at other times of the year; engorged
nymphs were found in May and August. Judging from observations at
the Alperton laboratory on material collected at various times in the
field, the ticks not being incubated during metamorphosis. Stockman
concludes that larvae which feed and moult up to May, nymphs which
feed and moult up to July, and adult females which feed and oviposit
up to August, are all derived from eggs hatched the previous year.
The larvae which hatch out, feed, and moult from July onwards, the
nymphs which feed and moult from August onwards, and the adults
which feed from October onwards are presumably all derived from eggs
of the same year. All eggs laid in February, May and June hatched in
July and August. Stockman believes that in nature the different
stages emerge and feed as follows: the larvae do so chiefly in July and
August, the nymphs in August-October, the females in October-
November, but all stages may hibernate (fed or unfed) and appear on
hosts in the spring. The females oviposit mainly in the spring.
Starting with eggs laid in the spring, the ticks, according to Stockman,
probably pass the next winter as gorged nymphs, and feed as adults in
April-May of the following year, the cycle presumably lasting about
290 days.

Further observations made in the field appear to me required before
definite conclusions can be arrived at; until such observations are made
Stockman's hypothesis, based almost purely on laboratory experience,
can scarcely be accepted.

The time H. cinnabarina var. punctata remains upon
the host. (Nuttall.)

Host main- Number of gorged

No. of Date when tained at a ticks collected on

Lot Host put on host temp, of succeeding days Renis

Larvae.

1 Hedgehog 12. vi. 1905 17° C. 18 on day 3

333 ,, 4
120 ,, 5-6
10 ,, 7

Biology of H. cinnabarina var. punctata 521

iVo. of
Lot

Host

Date wlien
put on host

Host main-
tained at a
temp, of

Number of gorged
ticks collected on
succeeding days

Remarks

Larvae.

2

Hedgehog

15, VI. 1905

17^^ C.

10 oil
151
707
330

78

(jl

1 day 4
)> 5

„ 7
,, 8
„ 9-10

3

8. vni> 1905

16^0.

30
477
142
6fi
40
15

,, -4
,, 5-6

,, 7
,, 8-9
,, 10
„ 11

4

,,

17. XI. 1905

10^ c.

10

,, 21

—

5

"

■28. XI. 1905

12° C.

21
30
18

,, 13

,, 1-5
,, 16

—

6

>»

19. XII. 1905

12° C.

•2
7
5

,, 7
,, 8-13
,, 14-19

7

Rabbit

2. VI. 1905

17° 0.

3

5

4

„ 13

,, 14
„ 15

~

8

"

6. VI. 1905

17° C.

116

119

30

,, 7

8

,, 9

—

91

Sheep

27. VI. 1914

17° C.

60

90

33

5

„ 7
,, 8
„ 9

Nymphs

1

Hedgehog

29. VII. 1905

20° C.

152 by day 12

—

2

>5

3. XII. 1905

12° C.

33

„ 25-27

—

3

"

19. XII. 1905

12° C.

4 on
1

1

day 13
„ H
„ 20

(34 lost).

4

>>

22. I. 1906

8°C.

1

1

„ 32
,, 33

(22 lost).

5

>>

17. III. 1901)

11° C.

7
11
4
4
8
11

„ 11
„ 12

M l:^

„ 14-17
„ 18-20
„ 21-29

(5 lost).

' Added to previously published record.

522

Genus Haetitaphu sails

No. of
Lot Host

Date when
put on host

Host main
tained at a
temp, of

Number of gorged
ticks collected on
succeeding days

Remarks

Nymphs.

6 Hedgehog

11. IV. 1906

12° C.

7 on

day 9

(33 lost).

7

14. V. 1006

13° C.

3

10

4

„ 7
„ 8
„ 9

(13 lost).

Adults ( ? ).

1 Hedgehog

9-11. IX. 1905

12° C.

1

„ 14

Put on with

1

„ 17

males, none

3

„ 18

lost.

Conclusions from the foregoing protocol regarding the time H. cin-
naharina var. punctata remains upon the host:

Larvae placed upon a hedgehog in June-August, the host being
maintained at 16-17° C, remain 3-11 days upon the host, the majority
dropping off gorged after 4-7 days. In November-December, the
temperature being 10-12° C, they remain upon the host 7-21 days,
the majority dropping off gorged after 13-21 days. When placed on
rabbits and sheep in June, at 17° C. they drop off gorged after 6-15
days.

Nymphs, placed on hedgehogs at 20"^ C, all dropped off by day 12;
at 11-13° C. they dropped off after 7-29 days; at 8° C. after 32-33
days.

Adults ( $ ) remained upon the hedgehog at 12° C. for 14-18 days.

Note: — Stockman (1911), who raised H. cinnaharina var. punctata
on cattle and sheep, records that.

Remain on host

Average period

Larvae

4—12 days

5 — 9 days

Nymphs

4—17

4—13

Adults

6—30

9—22

The time required by H. cinnabarina var. punctata
for metamorphosis.

In the protocols dealing with this section I include those of Stockman with my own :

Egg to Larva

Observer

No.

Date when first
eggs were laid

Time required
for meta-
Date when first morphosis
larvae appeared (days)

Tempera
ture

Nuttall

1

4. IV. 1905

25. VI. 1905

82

14° C.

Stockman

16/3

20. V. 1907

2. VIII. 1907

74

—

„

—

—

—

38

—

Remarks

The shortest time observed.

Biolofjii of II. cinnabar ilia var. punctata 523

Larva to Nymph.

Nuttall

Stockman

No.
1
2

32
33

Nymph to Adult.

Date when larvae
gorged on host

'20. XII. lyor,

1. I. 190()
30. VII. 1907
.30. VIII. 1907
30. VIII. 1907

Time reiiuired
for nieta-
Oate w)icn nH)rphosi.s 'J'enipera-

nymphs emerged

8. V.

2. VI.

15. in.

29. IV.

30. IV.

lOOC)
1900
1908
1908
1908

(<lays)
159
153
197
242
243
14

ture
10° C.

10° c.

Remarks
Host: hedgehog.

Hibernation at outside tem-
perature.

Ob.server

No.

Date when nymphs
gorged on' host

Date when
adults emerged

Nuttall

1

10. VIII. 1905

28. VIII. 1905

2

8. XII. 1905

13. I. 1900

,,

3

1. I. 1900

17. IV. 1900

"

4

28. III. 1900

14. IV. 1900

Stockman

26

27. IX. 1907

23. VII. 1908

27

30. III. 1908

15. IV. 1908

— The shortest time observed ;

the averaj^e time stated to
be 10-29 days.
Time required
for meta-
morphosis Tempera
(days) ture Hemarks

18 At room temp., Host: hedgehog,
then at 30^' C.
duringlast 3 days
30 — Host : dog.

. 107 9^C. Host: hedgehog.

17-26 At room temp., „ ,,

then at 30° C.
duringlast 14 days
229 — Hibernation at outside temp.,

a number survived winter
but died without moulting.

15 — ■

7 — The shortest time observed.

Longevity of unfed H. cinnabarina var. punctata.

The following protocols include Stockman's observations as well as my own, the latter
being but few :

Larvae.

Observer

No.

Date when
emerged from eggs

Date to which
larvae survived

Longevity
unfed (days)

Stockman

10/3

2.

VIII

.1907

1.

VI.

1908

303

lymphs.

Observer

No.

Date of
emergence

Date to which
nymphs survived

Longevity
unfed (days)

Nuttall

1

3.

VI.

1906

25.

IX.

1900

114

.>

2

12.

VII.

1914

12.

IX.

1914

02

Stockman

30

20.

X.

1907

11.

III.

1908

142

))

27

29.

IX.

1907

30.

III.

1908

182

„

53

10.

X.

1907

1.

V.

190H

197 \

,,

32

12.

X.

1907

30.

IV.

1908

200

,,

21

12.

IX.

1907

3.

IV.

1908

203

)>

29

20.

IX.

1907

30.

V.

1908

252

Remarks

This being the maximum

period in 1 out of 9 lots.

Remarks

At 10° C. Most of the lot

were dead.
At ca. 10° C; all iu 90

days'.
Hibernated at outside

temperature.
Ditto, and after starving

182 days raised to adults.

Hibernated at outside
temperature.

^ Record since added.

524

Genus Haemaphy salts

Adults.

Observer

No.

Date of
emergence

Date to which
adults survived

Longevity
unfed (days)

Remarks

Nuttall

.1

13. I. 1906

—

255

At 12° C. Lived longer,
death not noted.

,,

2

18. IV. 1906

—

160

,, „ ,,

Stockman

16/4

6. IX. 1906

18. IV. 1907

224

At outside temp., feeble
when state recorded.

Oviposition.

The manner in which the female H. cinnaharina var. punctata lays
her eggs was illustrated by me in the Harben Lectures 1908, p. 398,
but I kept no records of the time it takes for the tick to oviposit and
did not count the number of eggs laid by single females. Judging
from memory a female would lay 3000-5000. When the opportunity
arises I shall have enumerations made. Apart from this the only other
observations recorded are those of Stockman which are likewise incom-
plete. He states that the shortest and longest times which elapsed
before oviposition commenced were 10 and 211 days respectively, the
female usually ovipositing 24-29 days after abandoning the host. Some
gorged females (Lot 27) survived 216 days without ovipositing. He
gives no temperature records in his protocols; the only records with
dates are the following:

stockman
No.

16/3

16/4

Female dropped
from host

12. XI. 1906
12. X. 1906

Eggs first
laid on

20. V. 1907
15. II. 1907

Oviposition
commenced after

189 days
124 „

Stockman does not state how long oviposition lasts, but he says that
the process may be interrupted with the onset of cold weather and
resumed when the weather is warm.

The following description of the process of oviposition in H. cinna-
harina var. imnctata is quoted from Nuttall (vii. 1908, p. 398): "In
the accompanying figure (Fig. 450) I have illustrated the successive
stages I have observed in the process of oviposition in Haemaphysalis.
Prior to oviposition the capitulum is retracted so that it lies in a
depression beneath the scutum, the vulva approaching the capitu-
lum. When about to oviposit, a remarkable organ (which I named
' Gene's organ,' after its discoverer) is protruded between the scutum
and capitulum. The organ appears in the form of two vesicles
containing hyaline secretion, the vesicles being protruded and retracted
rhythmically whilst being completely extruded. When Gene's organ

2P

j2

d

3

'$

!z;

o

r=)

CO

1

34

526 Genus Haeinaphysalis

appears, the capitulum is depressed (Fig. 450 at 2) and the vuh^a is
arched upwards. The eapituhim next disappears in the depth of the
pit, and the ovipositor protrudes from the vulva through being
evaginated (8). The two halves or vesicles of Gene's organ grow in size
and show two glove-finger-like processes (4-8). The ovipositor grows
in length, forcing apart the vesicles (5), and the e^g appears (6) and is
protruded (7), after which the ovipositor is quickly retracted (8), the
egg being left between the vesicles and the capitulurn reappearing
beneath. Next (9), the vesicles begin to collapse and disappear (10),
leaving the egg resting upon the back of the capitulum, which presently
is tossed backward (11), so as to ' shovel ' the egg upward and backward
upon the scutum. The capitulum is then slowly depressed (12), and
Gene's organ reappears prior to the laying of another egg. It takes
two to four minutes to lay an egg, the intervals between ovipositions
varying somewhat in length. This process is repeated for each egg,
and entails a good deal of labour."

In Fig. 449 the position of Gene's organ (" ovipositing gland ") is
shown in a schematized longitudinal section through the capitulum of
the female.

Summary.

Haemapliysalis cinnaharina var. punctata seeks a host three times,
feeding thereon in the larval, nymphal and adult stages. It readily
attaches itself to the host 4-12 days after each ecdysis and is easily
reared under experimental conditions upon hedgehogs, rabbits, sheep,
cattle and ferrets. The larvae usually remain upon the host 4-7 days,
although they may remain attached anywhere from 3 to 19 days. The
nymphs usually remain attached for about a week, but this period of
attachment may range from 4 to 33 days. Adult females remain
attached for 6 to 22 days, the longer stay upon the host is doubtless
due to their waiting to be fertilized. Males and females both behave
like those of H. leachi, q.v. My observations suggest that when a
hedgehog (hibernating host) is maintained a t'a low temperature, the
larvae and nymphs remain somewhat longer upon the host than in
warm weather. The time required for metamorphosis is governed by
temperature, being much prolonged by cold. The larvae may hatch
from the egg after 38 days (Stockman's shortest time) to 82 days (at
14° C, Nuttall); the nymphs may emerge after 14 days (Stockman's
mininum) to 159 days (at 10" C, Nuttall) or after as long a period as
243 days (Stockman's maximum); the adults emerge after 7 to 229

B'wloijij of H. cinnubarma var. pamtata 527

Fig. 4/)0. Illustrating the mechanism of oviposition observed in H. cinnabarina var.
punctata. For description see text. (13) is a schematic sketch. Drawn from
living specimens so as to show the different stages of the process in sequence.
(Nuttall, 1908, p. 399.)

34—2

528 Greiius HaeniapJnjsalis

days (Stuckman's iiiiiiimum and maximum); my observations show a
range of 17 days (at 30" C.) to 107 days (at 9° C). The longevity of
the unfed tick is very marked; the records of Stockman and the writer
show that larvae may survive for 803 days, nymphs for 252 days, adults
255 days; all stages are therefore able to hibernate readily in this
condition. The time which elapses between the dropping of the gorged
female from the host and the commencement of oviposition is markedly
influenced by temperature, ranging accordingly from 10 to 211 days
(Stockman). Data are lacking as to the duration of oviposition and
the number of eggs laid by single females.

In nature, the tick will probably be able to complete its life cycle in
a year but it may well run into two years in our climate. Taking
average figures from the data recorded above, the cycle may be completed
in 125 da3's, but the developmental period may of course be much
prolonged :

Time required
in days

From the time egg is laid to emergence of larva ... 38

Larva hardens ... ... ... ... 7

Larva stays on host ... ... ... ... 6

Metamorphosis: Larva to Nymph ... ... 14

Nymph hardens ... ... ... ... 7

Nymph stays on host ... ,.. ... ... 7

Metamorphosis: Nymph to Adult ... ... 15

Adult hardens ... ... ... ... 7

Adult (?) stays on host ... ... ... 14

Gorged ? after dropping off host, begins to lay after ... 10

125 days

Haemaphysalis cinnabarina.

This tick requires three hosts upon which to feed as a larva, nymph
and adult. The first to raise the tick were Hooker, Bishopp and Wood
(1912, p. 98), who state that the larvae feed 5-12 days on rabbits or
guinea-pigs; they twice failed to feed on fowls. Metamorphosis from
larva to nymph takes 14 days at 79° F. or 76 days at 54° F. Some
unfed nymphs lived up to 55-75 days. Nymphs, placed on cattle, fed
for 5-8 days, and then underwent metamorphosis to adults in 26 days
at 68° F., or in 186 days at 60° F. (average temperature). Adults
survived unfed for 100-305 days. A female (probably unfertilized)
remained for 19 days upon an ox; she then dropped off replete and
survived for 172 days, but did not oviposit.

The following observations were made by me in Cambridge with the

Jiiitlotjl/ of II. riiniahariiKi ;")'JU

progeny of 2 $ (N. 2452) taken from (■(((tie in Winnipc^g, Ciuijulu, and
for which I am indebted to Mr .J. \i. N. Harrison. 'V\\v. ticks were
raised on rabbits in exj)eriniental (;ag(!s at loom tempei-ature (al)out
15" C.) in the la])oi'atory. "^I'he ticks were kipl in an incnb;itoi- at.
30" C. during nu'taniorjihosis.

The recently emer-ged larvae were placed on a rabbit mi 27. .\1.
1918:

No. of gorged larvae collected
on HUcce.<iBlve dayK

7

on flay ;">

270

,, (;

5(50

7

306

.» '^

160

,, '.)

98

10

Til

11

1.5

„ 12

1-2

i;5

The first nymplis emerged on 11. XII. I!)l'i, i.e., nietanKJiphosis from
larva to nymph lasted 10-1 1 days at 80' C. (HO F.). On 0. i. 1914 the
ii\in])lis (100) were placed on a rabbit:

-Ni). of «or«e(l iiyini>liH collettti-rl
on succeKsive days

'A oil (lay H

•t ., '.<

•A „ 11

r, ,, 12

1 „ i:j

2 „ 14
'A „ 15
2 ,, 17

The first adults emerged on 10. II. 1914, i c., metamorj)hosis from
nymph to adult lasted 2S days at 80 ' ( ■.

My residts accord witJi those of the American aiitjiois, but, the
nymphs I'cmaiiK'd longer u[jon the host.

Longevity of unfed ticks.

/j((rv(ie which emerged 7. XI. i!)l8, died after 48 days. Ni/mphs
which emerged 1 1. xii. 1918, died aftei- 84 days (at rootn temperature).

530

Genus Haemaphysalls

Haemaphysalis leporis-palustris.

~ The life-history of this species has been studied experimentally by
Hunter and Hooker (1907, pp. 53-54), Hooker (1908, pp. 47-48), Rohr
(1909, pp. 100-110), Hunter and Bishopp (1911, pp. 228-229), Hooker,
Bishopp and Wood (1912, pp. 90-96), and Hadwen (1912, pp. 97-98).
Rohr's observations were made in Brazil, his " Haemaphysalis proxima "
having been found on wild rabbits and agouti; Hadwen studied the
species in Canada, specimens having been collected in Manitoba and
British Columbia from Lepiis americanus; the remaining authors'
investigations were carried out in Dallas, Texas. In the following
pages I have sought to arrange the data supplied by these authors so
as to present them in a uniform manner. A summary will be found at
the end of the section.

Time the tick remains upon the host.

Larvae: drop off gorged beginning on the 5th day according to
Hooker (in October in Texas; host not stated). Rohr found that they
remain 6-11 days on the rabbit, he records an experiment in Brazil, as
follows:

6th day 10

lar

vae

di

opped oft

7

, 139

8

, lul

9

, 00

10

24

11

•5

12

0

no

more

collected

Total

389

Hooker, Bishopp and Wood give two experiments on rabbits and
on cattle:

larvae dropped off gorged

Rabbit I

Rabbit II

Cattle

4th day

0

5

0

5 ,,

25

77

19

6 ,,

27

71

75

7 „

33

29

27

8 ,,

7

5

11

9 ,,

5

1

2

10

1

4

0

Totals

98

192

134

Biologn of II. li'poris-palustriii 5ol

Hadwon placed larvae on a tame rabbit (in Canada 12. iv. 1912)
with the foUowinu" results:

tjtli day 10 larvae dropped off t^or^'ed
7 .. 44 „
B „ 17 „
9-10 ,, ^ „

Total 107

Nymphs: drop off gorged on the (ith day (from rabbit?) according
to Hooker. Ruhr states that they remain (J-ll days on labbits and he
records an expei-iment as folhnvs:

0th day 'd nymphs droi)pcd off gorged

7

,, 12

,,

8

,, 48

,,

9

„ 11

,,

10

■'>

,,

11

3

,,

12

0

no more

collected

Hooker, Bishop}) and Wood state that the nymphs remain 4-8 da3's
on the host, mostly 5-6 days; they carried out 4 experiments. Hadwen
placed 32 nymphson a tame rabbit and 4 dropped off gorged after 6-7
days.

Adults: accoi-ding to Rohr, the $ remains 19-25 days on the
rabbit before it drops off gorged, whereas the (/ remains on the host
long after the $ has left. Dead J's are commonly found attached to
the host's skin. Hooker, Bishopp and Wood had difficulties in
studying the adults; they only record the behaviour of 2 $ which
dropped off gorged on the 19th and 21st day respectively.

Time required for metamorphosis.

Egg to Larva: reckoned from the time the eggs are laid, according
to Rohr, the larvae emerge after

47-(31 days at ca. ■21-22'= C. (15 observations)
22-29 ,, „ 30" C. (3 „ )

none hatched out at 35° C.

Hooker, Bishopp and Wood record that larvae emerge after

22 days at ca. 32° C. (out of doors in vi.-vii. 1909)

23 ,, ,, 28° C. (in laboratory in VI.-VII.)
34 ,, ,, 28° C. ( „ ,, v.-vi.)

24 „

25° C. (

„ IX. 1907)

24 ,,

„ 25° C. ( „

„ IX. 1907)

34 ,,

„ 25° C. (

„ V.-VI. 1910)

532 ■ Genus Haem(q}]nisalis

Hadwen found that the larvae emerged 47 days after the eggs were
laid in May 1910; in 45 days after the eggs were laid in July 1911.
Hunter and Hooker (1907) state that metamorphosis lasted 24-31 days
in May-September.

Larva to Nymph: reckoned from the time the larvae drop off
gorged from the host, Rohr found that the nymphs emerged after

1.5-22 days at ca. 20-21'^ C. (5 observations)
8-12 ,, ,, 26-27° C. (5 „ )

7-9 ,, ,, 30° C. (3 „ )

the ticks died without moulting at 35° C.

Hooker, Bishopp and Wood recoid that the nymphs emerged after

18 days at ca. 20° C. (in March)
134 „ „ 16-5° C. (in November)

Hooker gives 18 days as the time required for metamorphosis in
October.

Nymph to Adult: reckoned from the time the gorged nymphs
abandon the host, Rohr found that the adults emerge after

19-2() days at ca. 20-22° C.
10-17 ,, ,, 27° C.

Hooker, Bishopp and Wood record that the adults emerge after

11 days at ca. 28° C. (.June 1908)
89 ,, „ 15° C. ( „ )

124 ,, was the longest period observed

These authors noted no difference between the j" and $ in respect
to the time required for metamorphosis. Hadwen records that from 12
gorged nymphs there emerged 2 adults after 53 days, and 1 adult after
58 days, in July-September, 1911.

Longevity of unfed ticks.

Larvae: Hooker, Bishopp and Wood found that larvae which
emerged early in the summer, died in about 60 days; others which
emerged in June, 1906, survived up to 258 days.

Nymphs: these survived, in three experiments, for 78, 246 and
307 days respectively, the ticks being kept in the laboratory. Specimens
taken from a rabbit survived for 342 days.

Adults: a ^ lived for 403 days, a % for 588 days, both having
emerged in May.

Jiioloffji of II. Irj}oris-pa/t(stris 533

Oviposition.

According to Rohr, the gorged and fecundated females begin to

oviposit after

2- 5 days, and oviposition lasts 14-21 days at ca. 21' C.

13-15 ,, ,, „ 31-57 ,, „ 15 "C.

Tlie female dies without ovipositing when maintained at 0°C.

Hooker, Bishopp and Wood state that females begin to oviposit

3-15 days after they abandon the host, i.e. after

3 days at ca. 32^ C. (June 1910)
15 „ „ 20*^0. (March li»10)
8 days in the spring and summer being the average for 25 ? s observed

According to Hadwen the female begins to oviposit aftei*

18 days (1 ? in May 1910)
8 ,, (15 „ „ 1911)
(5 ,, (1 ? „ July 1911)

The number of eggs laid per female varies as follows:
Hunter and Hooker record that a % laid 1112 eggs; they give such
small numbers of eggs as having been laid by three other females that
I conclude they must have been disturbed or that they were imperfectly
gorged specimens.

Rohr's observations are given in detail. The largest number of
eggs laid by one of five females was 2389 ; about 1820 would represent
the average number. The most replete females lay the most eggs.
The number of eggs laid per day, varied from 2 to 287 ; larger numbers
per day were laid when the females were maintained at a higher
temperature. We select two examples out of five tabulated by Rohr
as showing the number of eggs laid by two females on successive da3-s.
The first tick began to oviposit on the 5th day and died within 24
hours after oviposition had ceased as indicated by the +.

Day 12 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25

— — 46 75 79 112 158 178 148 176 172 187 118 82 134 76 49 27 19 2 +

52 298 148 105 46 171 236 161 209 92 225 135 99 159 132 58 84 44 23 22 10 — +

Totals 1830 and 23S9 respectively.

The female usually dies 0-4 days after oviposition has ceased (at
21° C), but she may survive 2-28 days at a lower temperature (15° C).

Hooker, Bishopp and Wood record that 8 females laid 59-2240 eggs
apiece, averaging 1517 ; they also note that the most gorged females
lay the most eggs. One female oviposited for 20 days, the maximum
number of eggs laid per day being 308. Females survi\i' 0-5 da^^s
after oviposition has ceased.

534 Genus Haemaplty sails

Weights and measurements.
Rohr gives a number of data which I have compressed in tabular

rm as follows :
Egg

Weiglit
in mgr.

0-07

Average of
1000

Size in mm.
0-58 X 0-49

Average of
10

Larva: uufed

0-055

200

0-72x0-52

,, replete

0-35

—

1-25x0-93

Nymph: unfed

0-2

20

1-22x0-85

,, replete

2-0

—

2-16x1-47

]\Iale: unfed

0-67

10

1-86x1-28

,, fed

0-70

—

2-0 xl-28

Female": unfed

1-10

10

3-87x1-37

,, replete

349-0

—

10x7-5x6-0

The foregoing measurements are of the length x breadth x thickness; in the case of the
gorged female only the largest size and weight attained is given.

A female which weighed 334'5 mgr. laid 1853 eggs, and, after ovi-
position had ceased, she weighed 78 mgr. Hooker, Bishopp and Wood
note that a female which laid 2240 eggs measured 11 "3 x 7 '5 x 5'3 mm.

Insect enemies.
Ixodiphagustexunns L. O. Howard (1907, p. 375) was first discovered
in engorged nymphs of H. leporis-pahistris taken from wild rabbits in
March and May, 1907, in Jackson County, Texas, by J. D. Mitchell.

Summary and Remarks.
H. leporis-pahistris requires three hosts upon which to feed in the
larval, nymphal and adult stages respectively, as was first shown by
Hunter and Hooker (1907), who state that it attacks wild rabbits in
such numbers in Montana, U.S.A., that the animals may be weakened
and thus rendered easy to capture. The larvae remain upon the host
for 4-11 days, the majority taking 7-8 days to gorge ; nymphs remain
attached for 4-11, the majority gorging in 6-8 days. The males persist
upon the host which the females have abandoned ; they may die whilst
attached to the skin. Rohr states that copulation must take place at
night upon the host as he has never succeeded in observing the process
by day. The female remains attached to the host 17-25 days. The
time required for metamorphosis is infiuenced by temperature: Larvae
emerge from the egg after 22-29 days at 30" C, or after 47-61 days at
ca. 22° C. ; nymphs emerge after 7-9 days at 30° C, or after 134 days at
16-5° C. ; adults emerge after 10-17 days at 27° C. or after 89 days at

Biology of TT. leporia-pahf^trifi 535

15° C, in one case they only emerged ut'tor 124 days. Longevity of
unfed ticks : larvae survived 258 days, nymphs up to 342 days, a male
and female lived up to 403 and 588 days respectively. Oviposition
commences 2-18 days after the female has abandoned the host, the
process lasting 14-57 days according to the surrounding temperature ;
oviposition does not occur at 0" C. The female dies 0-28 days after she
has ceased laying her quota of eggs. The normal number of eggs
appears to be 1800-2400, laid at the maxiininii rate of abi)iit 800 in 24
hours. •

Hooker states, from personal observation, that large numbers of
larvae and lesser numbers of nymphs, when replete, abandon the host
during the daytime. He regards this as a remarkable adaptation of the
tick to the habits of their usual hosts, the haie and rabbit. These
remain during the daytime in their resting places ov " forms " and roam
at night. The gorged ticks, dropping from the host in the forms,
undergo metamorphosis in a situation which favours their finding a host
when they emerge and are ready to feed. The observation is certainly
suggestive and it would be interesting to learn if other species of ticks
behave in a similar manner. The ticks usually attach themselves about
the ears and heads of rabbits.

Laboratory experiments by the authors quoted above (p. 530),
indicate that H. leporis-paUistris may, under favourable conditions,
complete its life-cycle once or twice in a year. This supposition is
strengthened by the statement of Hooker, Bishopp and Wood that all
stages occur at all seasons upon their hosts in nature in the United
States.

Judging from the foregoing data the life-cj^cle may be completed in
87 to 405 days, as follows :

Time required in days

Minimum Maximum

From the time the egg is laid to the emergence

of the larva ......

Larva hardens ......

Larva stays on host .....

Metamorphosis : Larva to Nymph .

Nymph hardens ......

Nymph stays on the host ....

Metamorphosis: Nymph to Adult .

Adult hardens .......

Adult ( ? ) stays on host

Gorged ? , having dropped from the host, begins

to lay eggs after .....

Totals 87 to 405 days

22 (at .30' C.)

f)l (at 22^C.)

7

7

4

11

7 (at 30°C.)

134 (at 10-5° C.)

7

7

4

11

10 (at27^C.)

124 (at?°C.)

7

V

17

25

2 (at 21° C.)

18(at?°C.)

536 Genus Haemcvphy sails

Haemaphysalis leachi\

Mr Charles P. Lounsbury, Union Entomologist, South Africa, was
the first to raise this tick experimentally and to demonstrate that it
transmits canine piroplasmosis in Africa. The protocols of his experi-
ments (1901, pp. 5-G; 1902, pp. 5-7; 1904, pp. 27-29) give the main
points in the life-history of the species. From his brief statements
regarding the tick, the following data are abstracted :

H. leachi is a three-host tick. The larvae and nymphs may drop off
gorged from the host as early as 48 hours after being put on. The
female remains upon the host for 9 days (minimum) to 12-15 days.
The time required for metamorphosis from egg to larva is 30-46 days
in summer and 80-110 days in winter; the nymph emerges from the
larval skin in 10-12 days in summer; the adult emerges from the
nymphal skin in 18-20 days in summer and 70-105 days in winter.
A female laid 4200 eggs. The males remain upon the host for many
weeks, they release their hold upon the skin of the host and seek the
females. By using an incubator he hastened the process of meta-
morphosis so that he was able to raise three generations in a year.
The tick abandons the dying host.

The following records relate (a) to ticks received in 1902-1906 from
Mr Lounsbury, Cape Colony, S. Africa (they were used for infection of
dogs with Piroplasma cams), and (b) to ticks received 23. ill. 1912 from
Mr R. E. Montgomery, Nairobi, B. E. Africa. The last named were
the progeny (larval stage) of two replete females collected beneath a
box in which a jackal had been kept. The first and second generations
raised in the laboratory are numbered respectively 1737 I and 1737 II.

The time H. leachi remains upon the host.

Host

No. of
Lot

Date wlien
l)ut on host

Host main-
tained at a
temperature of

Number of

gorged ticks

collected on

succeeding days

Larvae.

Hedgehog

10

10. X. 190o

10'^ C.

126 on day 4
250 „ 5

■j»

11

18. X. 1905

9"C.

104 „ 5

c> „ fi

12

28. X. 1905

9^ C.

0 „ 6

1 Reprinted, witli slight changes and additions, from Nuttall, iv. 1913, pp. 93-99.

Blolufj!/ ^'f il If'avhi

y.n

Host

Larvae.

Hc'dKebo'l

Jackal

No. of
Lot

Jackal 17:^7

"I"

Adults. (Female.

Date when
put on host

1(). XI. IDOG

Number of
Most main- Kortjed ticks
taineii at a collected on
temperature of succeeding days

Reniarli^s

24. III. 1912

Dog

1737
"II"

22.

XI. 1912

Nymphs.

Hedgehog

10

. 25.

IV. 190(5

11

11.

V. 1906

18^ C.

20^ C.

19 on

day 4

17

M 5

C

„ 6

7

„ 7

4

„ 8

8

,, 9-10

3

,. 11

1

,, 3

r.4

M 4

500

M 5

3.50

„ 6

.SO

,. 7

44

,. 3

375

,, 3

36

,, 5

20

11

14. V. 1906

11° C.

3

12

2

., 3

M 7

13

4. VI. 1906

16'^ C.

4

11 -5

1737
"I"

11. V. 1912

23° C.

13 1

y day 7

Larvae put on host 88
days (at 17° C.) after
ecdysis. Progeny of
1737 1, V 11.

SOputon, 20 recovered.
30puton, 17 recovered.

20 put on, 4 recovered.

13 nymphs put on host
15 days after ecdysis;
all recovered.

19.

v.

1912

23° C.

24 on
81
48
19

6

3

day 4
„ 5
„ 6
„ 7
,, 9
,, 11

Nymphs put on host
23 days after ecdysis.

27.

v.

1912

22° C.

12

51

5

3

7

„ 3
„ 4
„ 5
„ 7
„ 9

Nymphs put on host
31 days after ecdysis.

1

29.

v.

1902

Koom temp.

,, 12-

15

2

4.

VI.

—

,, 11-

12

—

3

18.

I.

1904

,,

—

., 11

—

4

26.

I.

,

—

„ 14

—

5

21.

X.

,,

10° C.

—

„ 12-

13

—

6

22.

X.

,,

10° c.

—

M 14

—

538

Genus Haemaph}) sails

Number of

No. of
Lot

Date when
put on liost

Host main-
tained at a
temperature of

gorged ticks

collected on

succeeding days

Remarks

Adults.

(Female.)

Jackal

1737

3. VII. 1912

20° C.

2 on

day 10

Adults (7 cf , 7 ? ) put

"I"

1
2

1

„ 12
., 14
M 16

on host 32 days after
ecdysis ; 6 ? 's re-
covered.

>>

1737

15. vn. 1912

20° C.

5

8

Adults (about 100 s

"I"

•5
18
10

7
1

1
1

9
„ 10
„ 11
„ 12
„ 13
,. 14
„ 16

and ? ) put on host
44 days after ecdysis ;
48 ? 's recovered.

The time required by H. leachi for metamorphosis.

The time required for metamorphosis is reckoned as follows: — (1) F,gg to Larva: from
the date on which the female began ovipositing to the day on which the first larvae
emerged from the egg-shells ; (2) I.arva to Nynqih : from the date on which the larvae
dropped off gorged to the day on which the first nymphs emerged ; (3) Nymph to
Adult : from the date on which the nymphs dropped off gorged to the day on which
the first adults emerged.

i

Egg to Larva.

Lot

Eggs laid on

Larvae emerged
after

Eggs main
tained at

1

15. XII. 1903

63 days

—

2

14. III. 1904

80

12^ C.

3

2. V. 1904

58

13° C.

17371

16

. vii.-l VIII. 1912

26

-37

20° C. ^

See table relating to the progeny of 39 ? 's, etc. p. 540.

Larva to Nymph.

Lot
la
lb
Ic
Id
2
3
17371

Nymph to Adult.

1

2

1737 I

Host

Date on which

gorged larvae

dropped off host

Nymphs

emerged

after

Ticks

maintained

throughout at

Ferret
Rabbit

13.
10.

VII. 1904

30 days
33

17° C.

Hedgehog

11.

,,

32

J J

Kid

20.

VI. 1904

44

,,

Dog

2.

IX. 1904

35

11° c.

Hedgehog

23-24.

X. 1905

39

10° c.

Jackal

27-30.

III. 1912

31

Adults emerged
after

24-26° C.

Rabbit

25.

VII. 1904

42 days

14° C.

Hedgehog

30.

IV. 1906

70

14° C.

Jackal

23.

V. 1912

15-16

24-26° C.

Blohijji of H. hachi ooO

Longevity of unfed H. leachi.

The longevity of the unfed tick is reckoned from the date of its emergence from the egg
in the case of the larva, from the larval skin in the case of the nymph, from the
uymphal skin in the case of the adult.

Larvae.

Lot Date of emergence Keiiiarks

1 12. VIII. 1904 StilllivelyafteroGdaysatlBC. In jar on earth.

2 '1\). VI. 1004 Dead after 44 days at 17'' C.

17371 12. vrii. 1912 Of the progeny of 32 ? 's isolated in separate corked tubes at
room-temperature in semi-darkness, 8 lots were found dead
after 322, 355, 360, 3(50, 362, 376, 376 and 400 days respec-
tively ; about 2o"/„ of the larvae, composing each of the
remaining 24 lots, survived for 354-378 days when they were
used for raising experiments*.

* Note added since the publication of the pajDcr from which this is extracted.
Nymphs.
1737 I 7. XII. 1912 Lively after 61 days at 12"-^ C. Then raised to adults.

Adults, lieceived vii. 1903 from Cape Colony, survived ca. 210 days at room temperature,
and when placed on a dog produced infection with piroplasmosis.

Observations relating to oviposition, etc.

Lot No.

Date wlien

gorged females

ilropped from host

Oviposition
began after

Ticks maintained
at temperature of

1

X. 1903

60 days

Cool room.

2

l.-iii. 1904

47

,,

3

3. III. „

24

,,

4

24. V. „

18

16° C,

5-8

iv.-v. ,,

14

16-21° C.

17371

vii.-viii. 1912

3-5

23° 0. *

* This holds for 33 out of 35 ? 's, the two remaining oviposited on days 9 and 12
respectively.

Summary.

Haeiuaphysalis leachi requires three hosts upon which t(^ feed chu-ing
its larval, nymphal and adult stages. It readily attaches itself to the
host at each stage in about a week after ecdysis. It is easily reared
under experimental conditions upon a number of different hosts (jackal,
dog, ferret, hedgehog, goat, labbit), and it ap|)ears to be inuiiaterial
upon which of these hosts it feeds. The larva and nymph i-emain
attached to the host foi- .3-7 days (2-8 days, Lounsbury), occasionally
longer; the females remain attached longer, i.e. 8-16 day.s. The males

540 Genus Haeniaphj /sails

Record relating to 39 H. leachi $ 's {Lot 1737 1) and
their progeny.

1 -S.2
<u 0) o

C » s > _

c4 o ° ° 5

0) t^'CO

1 10 3 25 2 — 27

2 10 4 26 5 — 26

3 12 4 29 5 2982 29

4 14 5 29 6 3626 28

5 14 3 31 8 4575 30

6 16 4 28 13 3362 28

7 8 4 32 5 3660 30

8 8 0 26 8 3043 29

9 8 4 21 5 — 28

10 8 9 27 8 2400 32

11 8 4 21 0 — 30

12 9 4 26 7 4803 33

13 9 3 34 15 3815 32

14 9 4 17 1 — 30

15 9 5 24 9 3595 33

16 9 5 31 12 — 32

17 10 4 22 13 — 30

18 10 3 30 31 3763 30

19 10 3 24 5 — 29

20 10 3 29 7 4817 29

21 10 3 31 15 — 29

22 10 4 32 6 — 29

23 10 4 29 3 3811 29

24 10 4 29 2 — 30

25 10 3 30 3 — 30

26 10 4 29 3 3395 32

27 10 — — — — —

28 10 3 30 15 — 30

29 10 5 30 6 — 30

30 10 — — — — —

31 10 5 28 6 — 29

32 11 — — — — —

33 12 — — — — —

34 10 3 30 6 — 30

35 10 3 26 3 — 31

36 10 12 — — * —

37 13 3 28 2 — 30

38 14 3 28 0 — 34

39 16 5 37 10 — 37

16. VII.- 10. VIII.- at 20° C. 12. viii.-

1. VIII. 12 11. IX. 12 11. IX. 12

at 22-5° C. at20°C. at20°C.

* Laid a few unfertilized eggs.

Biology of IL leachi :A\

may remain upon the host for many weeks (according to Lounsbury).
The temperature of the air within the limits observed (9-23'' C.) appears
to exert h'ttle or no influence upon the time the tick remains upon the
host, the warmth emanating from the hxtter being doubtless sufficient
to keep the ticks active. The time required for metamorphosis ' is
influenced by temperature, thus the larvae hatch after 26-37 days at
20° C, in 58 to 80 days at 12-13° C; the nymphs emerge, as a rule,
after 30 to 40 days; adults emerge after 15-16 days at 24-26 C.
whereas they may only emerge after 42-70 days at 14' C. The longevity
of the unfed tick is considerable when the conditions are favourable ; in
small corked bottles some larvae were still active after 399 days, the
nymphs after 61 days and the adults after about 210 days when
maintained at room temperature in semi-darkness and all stages fed
upon their hosts after these periods. When males and females are
simultaneously placed upon the host they scatter, but the sexes are
found attached in close proximity to each other after 2-3 days. Copu-
lation must take place upon the host, though it has never been actually
observed. (Lounsbury has seen males, which he had marked, detach
themselves and reattach themselves close to females ; a male may mate
with more than one female.) I find that the males do not seek the
females as do Ixodes when the sexes have been removed from the host.
The time which elapses before oviposition commences, after the replete
female a andons the host, is markedly influenced by temperature ; thus,
when females were placed at 23° C. they began to lay after 3-5 days, at
1 6-21° C. after 14-18 days, at lower temperatures after 24, 47 to 60 days.
Whereas an occasional female dies as soon as oviposition has ended,
others may survive for a few days or, in exceptional cases, for a month
The female lays from 2400 to 4800 eggs. H. leachi begins to abandon
its host on the approach of death in a manner that neither Lounsbury
nor myself has observed in other ticks.

In nature, this tick may doubtless run twice through its life cycle in
a year. By the use of an incubator, as first shown by Lounsbury, this
author succeeded in raising three generations in a year. Taking
average figures from my protocols of ticks raised under favourable
conditions, the cycle may be completed in 123 days, as follows :

' See reference to Lounsbury's observations on p.

536.

35

542 Genus Haemapliy sails

Time required,

in days

From the time egg is laid to emergence of larva

30

(Eggs to 20° C.)

Larva hardens

7

Larva stays on host

5

Metamorphosis : Larva to Nymph

31

(Larvae at 17° C.)

Nymph hardens

7

Nymph stays on host ...

5

Metamorphosis : Nymph to Adult

15

(Nymphs at 24°C.)

Adult hardens

7

Adult ? stays on host ...

12

Gorged ? drops from host and waits before layi

ag

4

123 days

Relation of H. leachi to disease,

H. leachi occurs all over Africa, and wherever it occurs it is the
carrier of a very fatal disease in dogs known as canine piroplasmosis or
malignant jaundice. The relation of the tick to this disease and the
pathology, parasitology and treatment of the malady, are fully treated
in the following papers, the titles of which will be found in Biblio-
graphies I and II of this work :

Nuttall, 1904, pp. 219-257 ; 1905, pp. 12-32 ; ix. 1908, pp. 513-526.
Nuttall and Graham Smith, vn. 1905, pp. 237-249 ; 1906, pp. 536-651 ;
1907, pp. 232-272; x. 1908, pp. 243-260; ix. 1909, pp. 211-214,
215-228. Nuttall and Hadwen, vn. 1909, pp. 156-191; ix. 1909,
pp. 229-235. Nuttall, x. 1913, pp. 302-320. Many of these papers
are illustrated.

Haemaphysalis concinna.

The only observations on the biology of this species are by Brumpt
in France, who has kindly supplied me with his MS. notes thereon.

Brumpt has found the adults only upon deer, attached in the
cervical region, especially along the mane, occasionally at the base of the
ears, and rarely elsewhere. When placed experimentally upon dogs,
they attached themselves anywhere. The sexes are found attached
close together upon deer. Nymphs and larvae were found by Brumpt
on deer and hedgehog and it is probable that they occur on other
mammals ; they attach themselves anywhere upon the hosts mentioned.

Biology of H. eoncinna 543

Time the tick remains upon the host.

The Larvae, a few days after emergence, attach themselves quickly
to a variety of hosts (deer, hedgehog, dog, guinea-pig, rat, mouse) and
gorge themselves with blood or lymph in 8-10 days ; nearly all of them
abandon the host in 3-5 days.

The Nymphs, a few days after emerging, attach themselves to the
same hosts as do the larvae and feed to repletion in 8-12 days, mostly
abandoning the host in 4-5 days.

The Adults, after fasting for weeks or months, attach themselves to
the host whereon they copulate, probably after 4 days, for the sexes are
then found attached close together with their venters juxtaposed. The
fertilized female becomes replete in 8-10 days ; she is then slate-
coloured and weighs on an average, when fully gorged, 0"35 g.
Unfertilized females may stay for several weeks upon the host
remaining about a third gorged.

Time required for metamorphosis.

Egg to Larva : the larvae emerge some weeks after the eggs are
laid and they are capable of living for a long time unfed.

Larva to Nymph: nymphs emerge 17 days after they abandon the
host as gorged larvae when maintained at 25-30° C. From 64 gorged
larvae found upon a hedgehog, 64 nymphs emerged.

NympJi to Adult: adults emerge after 22 days at 25° C. From 4
gorged nymphs found on a stag at Chantilly there emerged 4 cT after 2
months: from 15 gorged nymphs found on a hedgehog there emerged
15 adults. Certain incompletely gorged nymphs undergo metamorphosis
to adults very slowly and they often die before it is completed. H. eon-
cinna differs in this respect from certain species of ticks, for instance
Amblyomma cayennense (as seen by Aragao, 1912, p. 96), Rhipicephalus
appendiculatus and R. sanguineus (Nuttall, 1913, p. 195; Cunliffe, 1914,
p. 372), Dermacentor reticulatus (as seen by Brumpt), wherein im-
perfectly gorged nymphs were found to give rise to small adults.

Oviposition.

At a temperature of 18° C, oviposition occurs in about the second
week. The eggs are larger than those of Ixodes ricinus, they are dark
salmon coloured. Each female usually lays upwards of 1000 eggs.

544

Genus Haemaphy sails

Table showing the prevalence of Haemaphy salts on Deer in France
as observed hy Dr E. Brumpt.

Number

of deer

examined

Total 29

Date
1911

3. XI.
7. XII.

16. XII.
1912

24. I.
13. II.

20. II.

27. II.

28. II.
18. III.

4. III.

30. III.
13. IV.

17. IV,
24. IV.

31. IX.
3. X.
8.x.

14.x.

21. X.
21. X.
24.x.
18-19 XI.

and XII.

1913
28. IV.
30. IV.

Locality *
Fontainebleau

Indre

Fontainebleau

Vienne

Indre

Compiegne

Fontainebleau

Indre

Fontainebleau

Chantilly

H. concinna

02
03
?14

L 1

Q3

^1

cf 18

(f - ? many

^3 . . .

<? - ? few ©1 L 3

c? 37 ? 34 o 15 L 2

c? - ? -O-L f. numerous

<J - 9 -O-L f. numerous

3 150 ? 30 ©40 L 10

<?-

<?1

. ©1
? f. nam.

. ©1
. ©1

L2

©1

L2

cT- ? -©-L many
c? 30 ? few . .

H. iiicrmU

c?-?14

c? - ? f . numerous

J-?52

cTl . . .

c?3 ?3 . .

J4 ?7 . .
c?2 ?2 . .
<? - ? many

cT - ? few . .
<? 2 ? 15 . .
c? - ? f . numerous
rf - S f . numerous
3l

H. cinnabar ina
var. punctata

Jl

Note : the signs <? , ? , ©, L denote male, female, nymph and larva respectively. The
abbreviation "f. num." or " f . numerous" stands for "fairly numerous."

* Fontainebleau (Dept. Seine-et-Marne) ; Dept. Indre ; D^pt. Yienne ; Compiegne
(D6pt. Oise) ; Chantilly (Dt^pt. Oise).

Season.

From the above table relating to Brumpt's observations on the
seasonal occurrence of H. concinna (and H. inermis) in France, it will
be seen that all stages were encountered on deer in February-April
and October, none were found in January and September, 1912, and none

Biology of H. concinna 545

were encountered in November-December of 1912 and 1913. All stages
were most numerous in March and April, usually few or none being
encountered later. Observations for the 4 months May-August are
lacking. In one instance (April, 1912) the number of </s greatly
exceeded that of the $s encountered on one host.

On two hedgehogs, examined at Chantilly (27. vi. 1911 and 6. vi. 1912),
he found immature stages only: 15 nymphs and 64 larvae, and
19 nymphs and 174 larvae respectively.

Insect parasites of H. concinna.

Brumpt has several times found the nymphs of H. concinna at
Chantilly and Fontainebleau to be infested with Ixodiphagus caucurtei
du Buysson (see p. 547). Under experimental ccmditions, the nymphs
were readily attacked by the chalcidid but only about 2-4 per cent,
became parasitized.

Haemaphysalis inermis.

The biology of this species has been studied by Dr E. Brumpt of
Paris who has kindly placed his notes at my disposal thus enabling me
to compile the following account of the life-history of H. inermis.

Brumpt finds that the adults, which are very active, occur commonly
on deer in France during the months November to April inclusive.
They quickly abandon the host and readily reattach themselves to the
hedgehog or dog in the laboratory. The ticks occur in all situations
upon the host but they attach themselves by preference, as does
/. ricinus, in places where the skin is thin. On deer, they occur in the
axillary and inguinal regions and occasionally upon the ears.

H. inermis requires three hosts upon which to feed in the larval,
nymplial and adult stages respectively. Both sexes occur upon the
host on whom copulation takes place. Brumpt has never found the
sexes attached at the same spot on the host as is usually seen in
H. concinna.

Time the tick remains upon the host (Laboratory
experiments).

The Lai-vae, when a week old, attach themselves immediately in any
situation upon the host, be it either a mammal (hedgehog, mouse, rat)
or reptile (lizard) ; they gorge very rapidly, some being replete in

35—3

546 Genus Haemaphysalis

li hours; the majority, however, engorge in 2-3 hours, whilst some
feed for 20 hours. Their bodies appear bright red at first owing to the
blood ingested, but after some days they assume a bluish hue.

The Nymphs are often ready to feed a few days after emergence ;
they gorge rapidly ; some are replete within 1 hour, the majority gorge
within 2 hours, whilst others remain longer upon the host, but in no
case do they remain attached longer than 24 hours upon a mammal.
They suck blood for longer periods when attached to cold-blooded
animals (lizards) at room-temperature.

The Females may remain attached to the host for weeks whilst
awaiting the male, during which time they appear but partially fed.
In the presence of the male, i.e. when the female is fecundated, she
usually grows replete and abandons the host in about 8 days.

Time required for metamorphosis.

Egg to Larva: the larvae emerge 7-8 weeks after the eggs are
laid when the latter are kept at 18° C.

Larva to Nymph: the nymphs emerge after 17 days at 25° C, the
time being reckoned from when they dropped off the host as gorged
larvae.

Nymph to Adult: an exceptionally long time is required for
metamorphosis at this stage. When the nymphs have fully gorged
themselves they moult after 6 months at 15-25° C, some may, however,
require 10 months, and incompletely fed specimens are found to be
still active after 16 months.

Observations relating to oviposition.

Oviposition usually begins on the 5-6th week (at 18° C.) after the
replete and fecundated female has abandoned the host. The eggs are
large, dark salmon-coloured, and only number about 200 per female.

Season.

From the table on p. 544, which contains Brumpt's observations on
H, inermis and H. concinna, we learn that the adults of H. inermis
occur on deer in France during the six months November to April
inclusive. Observations for the four months May- August are lacking.
The ticks were not found in September-October.

Biology of H. inermis 547

Remarks upon the exceptional behaviour of H. inermis.

H. inermis offers an exception to all the ticks whose life-histories we
know in that it gorges with remarkable rapidity both in the; larval and
nyniphal stages. It is the rule in Ixodid ticks for all stages to remain
attached to the host for several days. In Argasidae we do not know of a
species where the larvae feed as rapidly as does the larva of H. inermis
although the nymphs, 0. megnini excepted, are rapid feeders (see
pp. 81-104, 325-345 of this work). Whilst this habit of rapid feeding
may favour the survival of the species, the long time required for the
process of oviposition and for metamorphosis from egg to larva and
from nymph to adult must exert a contrary effect. This phenomenon
observed in H. inermis by Brumpt is very interesting and worthy of
attention.

Insect parasites of H. inermis.

A chalcidid called Ixodiphagiis aiucurtei du Buysson, 1912 (p. 246)
was discovered by Brumpt in the nymphs of Ixodes ricinus in France
and raised by him to the seventh generation in the laboratory. This
insect attacks the nymphs of H. inermis. Whereas scarcely 1 "/o of
these nymphs become infested, all of the nymphs of /. ricinus, Denna-
centor venustus, D. reticulatus and Rhipicephalus sanguineus in which
the parent /. caucurtei deposits her offspring, die and yield imagines
of Ixodiphagiis.

548

Genus Haeifnaphysalis

INDEX TO VALID SPECIES OF HAEMAPHYSALIS

Together with a list of collections in which the types
are to be found.

N.B. The reader should also refer to the list of condemned
and doubtful species on p. 512.

Name of Species,
authors and date

*aborensis Warburton, 1913

*aciculifer Warburton, 1913

aculeata Lavarra, 1904

bancrofti Nuttall and War-
burton, 1915
birmaniae Supine, 1897
bispinosa Neumann, 1897

bispinosa var. intermedia War-
burton and Nuttall, 1909
calcarata Neumann, 1902

calcarata var. houyi Nuttall and

Warburton, 1915
calvus Nuttall and Warburton,

1915
campanulata Warburton, 1908

cinnabarina Kocb, 1844
cinnabarina var. punctata (Cane-

strini and Fanzago, 1877)
concinna Koch, 1844
cornigera Neumann, 1897

*cornigera var. anomala War-
burton, 1918
cuspidata Warburton, 1910

dentipalpis Warburton and Nut-
tall, 1909

doenitzi Warburton and Nuttall,
1909

elongata Neumann, 1897

Collections which include
the types and co-types

Calcutta, 1 ¥

London (b), 1 <? , 1 ?

Borne, 15 ^ ; Cambridge,

1<?, 1?
Cambridge, 9 d , 11 ? , 3 o,

IL

Stages of each

species which

are known

¥

6 ¥

^ ¥

(f ¥ o L

398
411
440

487

Jlava Neumann, 1897

Genoa and Cambridge, 4 c?

<?

415

Toulouse, 1 ¥ ; Cambridge,

(f ¥ o

L

426

c? o

Cambridge, <? ¥ ; London

<f ¥

483

(c), ?

Paris (a), Toulouse, Cam-

cT ¥ o

442

bridge, 5 <f , 8 ¥ , 1 o

Berlin (a) ; Cambridge, 7 S ,

<f ¥ o

444

l¥, 2o

Cambridge, 3 <? , 1 ¥

<r ?

445

Cambridge, London (a, c),

c? ¥

491

Paris (a), Berlin (a), Tou-

louse, Washington (d)

Berlin (a), 1 ¥

<? ¥ o

L

372,

528

?lost, <? ¥

<f ¥ o

L

378,

519

Berlin (a), 1 c? , 1 ¥

<? ¥ o

L

452,

542

Toulouse, <? ¥ ; Paris, <? ;

<? ¥

500

Cambridge, <?

Calcutta, 1 d

<?

504

Cambridge, Berlin (a), Tou-

c? ¥ o

L

438

louse, 30 <? , 2 ¥ , 7 o, 4 L

Cambridge, London (c), 4 s

<?

505

Cambridge, Berlin (a), 4 c?,

J ¥

482

3¥

Toulouse, Cambridge, 6 cf ,

c? ¥

498

3¥

Toulouse, c? ¥

cT ¥

408

= species not represented in our collection in Cambridge

Index to Valid Species

i49

Name of Species,
authors and date

formosensis Neunmnu 11I13

Iwodi Warburton and Nuttall,

1909
hoodi var. orientalis Nuttall and

Warburton, 1915
howletti Warburton, 1913
humerosa Warburton and Nuttall,

1909
hystricis Supino, 1897
iiiermis Birula, 1895

inermis var. aponommoides War-
burton, 1913
*j(iponica Warburton, 1908
japonica var. douglasi Nuttall

and Warburton, 1915
kinneari Warburton, 1913
^kochi Aragao, 1908

leacJii (Audouin, 1827)
leporis-palustris (Packard, 1869)
montgomeryi Nuttall, 1912

numidiana Neumann, 1905

ohtusa Donitz, 1910
papuana Thorell, 1882
parmata Neumann, 1905

parva Neumann, 1908

silacia Robinson, 1911
simplex Neumann, 1897

spinigera Neumann, 1897
spinige7'a var. uovae-guineae

(Hirst, 1914)
*spinulosa Neumann, 1906
*turturis Nuttall and Warburton,
1915
vidua Warburton and Nuttall,

1909
tcarburtoHi Nuttall, 1912

wellingtoni Nuttall and War-
burton, 1907

Collections which include

Stages of each
8i)ecie8 which

the types and coty])e8

are known

Page

Berlin (b), Toulouse, Cam-

d- V

400

bridge, 20 tf , 1 ?

Cambridge, Toulouse, ,J s.

<? ? O

L

483

? 8, 2 o, 1 L

London (b), Cambridge, G <? ,

S ?

486

3?

Cambridge, 1 d' , 1 ?

J ?

493

Cambridge, 3 c^ , 1 ? , 1 o

cT ? O

496

Genoa, Cambridge, <? ?

<J ?

422

Petrograd, 1 ? ; Cambridge,

<? ? O

L

362, 545

(? o L

Cambridge, Berlin, Tou-

?

367

louse, 23 ?

London (a), 2 c?

if

402

London (a), Cambridge, 4 <?

<f

403

Cambridge, 1 ?

Rio de Janeiro {a, b), 2 cj ,
2?

,? lost?

? lost?

Cambridge, Berlin («), Tou-
louse, 12 cT , 4 ?

Toulouse, Cambridge, 3 c? ,
1 ?

Berlin, Cambridge, many <j

? d^ ? ; Cambridge, o L

Toulouse, (f ¥ o L ; Cam-
bridge, (f ?

Lausanne, Toulouse, Cam-
bridge

Cambridge, 4 ?

Toulouse, 12 (f, 1 o; Cam-
bridge, 1 s

Toulouse, Paris, 4 c? , 5 ?

London (a), d ? ; Cam-
bridge, (f ? o

London («), 2 ?

Berlin (b), 1 j

Cambridge, 1 s

Cambridge, Toulouse, 5 <f ,

9 ?
Cambridge, 1 <? , 2 ? , o L

9

J ?

cT ? o L

<f ? o L

<? ?

<? ?

<?

cT ? o L

d" ? o L

cf ? o L

397
413

460, 536
387, 530
395

478

477
404
418

435

?

416

<r o

458

cT ?

447

<r ? o

449

?

476

<r

410

d

495

<i ?

369

<f ? o L

479

= species not represented in our collection in Cambridge.

550

Genus Haemaphysalis

The types of the following have not been accessible : papitana <? ? , leporis-palustris ? ,
leachi c? , inermis ? , cinnaharina var. pttnctata <? ? . If we except these, all of our
specimens, that are not either types or co-types, have been carefully compared with the
types.

Note. In the second column of the foregoing list the whereabouts of the collections is
but briefly indicated ; places not referred to in the following list will be found recorded on
p. 348 of Part II :
Berlin (a) signifies Entomologische Abteilung, Zoologisches Museum, Berlin.

,, [b) ,, Deutsches Entomologisches Museum, Berlin-Dahlem.

Genoa ,, Museo Civico di Storia Naturale, Genoa.

London (a) ,, British Museum (Natural History), London.

,, {b) ,, Imperial Bureau of Entomology, London.

,, (c) ,, London School of Tropical Medicine, London, E.
Petrograd ,, Collection of Prof. A. Birula, Imperial Academy of Science,

Petrograd.
Kio de Janeiro (a) signifies Collection of H. de B. Aragao, Eio de Janeiro, Brazil.

{b) „ „ „ Dr A. Lutz,

Kome signifies E. Museo Zoologico Universitario, Kome.

i

I

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[p. T. o.

PARASITOLOGY (continued)

CONTENTS OF VOL. VIII, NO. 2 (September 1915)

BouLENGEB, Charles L. The Life History of Nematodirus Jilicollis
Rud., a Nematode Parasite of the Sheep's Intestine. (With
Plates VIII and IX and 5 Text-figures.)

Watson, E. A. Dourine and the Complement Fixation Test.

Coles, Alfred C. Multiplication-forms of Trypanosoma lewisi in the
Body of the Rat. (With Plates X and XI.)

Cooper, W. F. and Laws, H. E. Some observations on the Theory
and Practice of Dipping. (With Plate XII and 2 Text-figures.)

Nuttall, G. H. F. and Hindle, E. Experiments in the "TrypoSa-
frol" Treatment of Trypanosomiasis {T. brucei) in Guinea-pigs
and of Piroplasmosis in Dogs.

TICKS

A Monograph of the Ixodoidea

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