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Nuttall

Ticks
Ixodoidea

Farts 1-2

TICKS

A MONOGRAPH OF THE
IXODOIDEA

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^ICKS

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.)

7*1-

X

Cambridge

at the University Press

1908

&2T4-3

CambrtDgt:

PRINTED BY JOHN CLAY, M.A
AT THE UNIVERSITY PBE88

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 volume, 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, will
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

PABT 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.

v i i

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

cucumerinus .28

transgariepinus 29

bruinpti ............ 30

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

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

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

brevipes ............. 5

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

List of species 41

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

viii Contents

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

PAGE

Ornithodoros savignyi 42

„ moubata 46

„ coriaceus ........... 55

„ turicata ........... 57

„ talaje 59

„ pavimentosus 62

„ erraticus ........... 63

„ tholozani 65

„ lahorensis ........... 67

„ furcosus 70

„ megnini 71

„ canesirinii 78

„ papillipes 79

„ morbillosus 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

Effects of hite on man and animals . . . . . . 85

Economic importance 87

In relation to disease 88

Methods of destroying 90

Argas reflexus : Life history 91

Effects of bite 92

In relation to disease .94

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

The genus Ornithodoros.

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

Effects 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.

I X

LIST OF ILLUSTRATIONS

Plato I. Fig. 1. Argas refleanis (J, dorsum.
Fig. 2. „ reflexus § , venter.

Fig. 3. „ persicus £ , dorsum.

Fig. 4-5. „ vespertilionis, last stage nymph, dorsum and

venter to face p. 4

Plate II. Fig. 1-2. Ornithodoros savignyi $ , dorsum and venter .

42

Plate III. Fig. 1-2.

talaje var. capensis, dorsum and venter

to face p. 60

IN THE TEXT

FIGURE PAGE

A and B. Schematic, to show differences between Argasidae and Ixodidae 1

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

descriptions of Argasidae 7

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

to in the descriptions of Argasidae ..... 7

3-4. A. persicus $? , dorsum and venter ...... 13

5-8. A. persicus, details of integumental structure .... 14

9. A. persicus, the anus ......... 14

10. A. persicus, the spiracle IS

11-12. A. persicus, the male and female genital orifices, etc. ... 15

13-14. A. persicus, the capitulum and lateral view of the palp . . 15
15-18. A. persicus, the digits of the chelicerae, the hypostome, and

tarsus I seen in profile ....... 16

19-23. A. persicus, representing stages of development from unfed larva

to second stage nymph 16-17

24-25. A. persicus, the larva, dorsum and venter 17-18

26. A. persicus, tarsus and foot, and hypostome of larva ... 19

27-28. A. reflexus (J, dorsum and venter 24

29-32. A. reflexus, a palpal hair, digits and hypostome .... 24

33. A. reflexus, tarsi I and IV 25

34-35. A. hermanni, digit and hypostome 28

36-37. A. transgariepinus 9 , dorsum and venter 29

38-41. A.brumpti $? , dorsum and venter 30-31

List of Illustrations

FIGURE PAGE

42-43. A. brumpti, capitulum of $ , ventral and lateral aspects . . 31

44-47. A. brumpti, digit, tarsi I and IV, details of integument . . 32

48. A. vespertilionis $ , venter 35

49. A. vespertilionis, detail of peculiar structure on venter . . 36
50-51. A. vespertilionis, first and second stage nymphs .... 36
52-56. A. vespertilionis, intestinal caeca, detail of integument, capitu-
lum, hypostome, digit 37

57. A. vespertilionis, larval hypostome, digit, dorsal plate, anus . 38

58. The tarsi I and IV of 9 species of Ornithodoros compared . . 40
59-60. Ornithodoros savignyi, details of integument .... 43

61-63. 0. savignyi, palp, hypostome, digit 44

64-65. 0. savignyi, tarsi I, III and IV, larva, dorsum and venter . . ' 45

66. 0. moubata $ , dorsum and venter ...... 49

67-68. 0. moubata, part of (J venter, cast skin of nymph . . 49

69. 0. moubata, capitulum of $ , ventral aspect .... 49

70-71. 0. savignyi, legs I and IV 50

72-73. 0. moubata, legs I and IV . . 50

74-75. 0. moubata, nymph, venter and capitulum 51

76-79. 0. moubata, egg, larva and details thereof . . . . ■. 51

80. 0. moubata, capitulum and palp of larva 52

81-82. 0. coriaceus $? , dorsum and venter 56

83-84. 0. coriaceus $ , side view and venter 56

85-87. 0. turicata (J , dorsum and venter, digit 58-59

88. 0. talaje, hypostome and digit 60

89. 0. talaje var. capensis $ , anterior part of venter ... 62
90-92. 0. pavimentosus, legs I and IV, and I and II ... . 63

93-94. 0. tholozani, digits of $ and $ 66

95-97. 0. lahorensis $? , dorsum and venter 67-68

98-100. 0. lahorensis, digit, hypostome, tarsi I to IV . . . . 68

101. 0. furcosus <% , tarsi I, II and IV 70

102-103. 0. megnini £ , dorsum and venter 72

104. 0. megnini, digits of nymph and J 72

105-107. 0. megnini, young nymph, dorsum and venter .... 73-74

108. 0. megnini, capitulum of nymph in ventral aspect ... 74

109-110. 0. megnini, larva, dorsum and venter 76

111-112. 0. megnini, leg of larva, gorged larva 77

113. 0. canestrinii $, anterior part of venter, tarsus I ... 78

114. 0. papillipes $, anterior part of venter, tarsus IV ... 79

I

SECTION I

DEALING WITH THE CLASSIFICATION OP 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 :

VA/A. futm* f|G B

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

Argasidae Ixodidae

Sexual dimorphism

Slight

Marked.

Capitulum :

Base

Ventral, in camerostome, no
porose areas in ?

Anterior, porose areas in ? .

Palps

Leg-like, with sub-equal ar-

Relatively 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.
Generally large, well behind

Spiracles

Very small, more anterior

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 hahits 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 with leathery integument
and slight sexual dimorphism, with capitulum inferior in the nymphs
and adults, the palps being free 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
rejlexus 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 Gimex
lectularius, 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 Caris (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 Ornithodoros, 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, Argas1. 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, 18902.

Argasiden Koch, 1844, p. 219 ; 1847, p. 11.— Fam. Argantidae Agassiz, 1846, p. 32.
— Fam. Argasides Furstenberg, 1861, p. 208.— Subfam. Argasidae Murray, 1877,
p. 180.— Berlese, 1885, p. 131 (incl. one genus : Argas).— Riley, 1887, p. 744.—
Tribe Argasides Megnin, 1880, p. 132 (comprising but one genus : Argas). —
Laboulbene 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; 1900 b, p. 436, and numerous recent
authors. — Trouessart, 1891, p. 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 Argas 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 distinction. 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.

2 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. Argasinis Railliet, 1893, p. 715.

Genus 1. ARGAS1 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 own
length from the anterior border. On both dorsum and venter there are
numerous symmetrically arranged discs2, generally round or oval, more
or less disposed in radial lines. Elsewhere the integument is minutely
wrinkled into irregular zig-zag folds. Eyes absent.

Type species : Argas persicus3.

Synonymy and Literature: Genus ARGAS Latreille, 17964.

Argas Latreille, 1796, p. 178, obs. specimen 8 mm. long, had seen specimens from
Tuscany (A. reflexus, doubtless) ; 1804, p. 53 : type, Aearus reflexus ; 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.— Duges, 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. — Kuchenmeister and Ziirn, 1879, p. 538
et seq.— Berlese, 1885, p. 131.— Ludwig, 1886, p. 612.— Railliet, 1886, p. 499 ;

1 Often misspelt Argus, see Ant. Duges and Milne Edwards, pi. 27 ; Fullager, 1874 a ;
Francis, 1894, p. 452 ; Fuller, 1897, p. 590, etc. The derivation of " Argas " attributed to
the Greek apyi)s ( = white, glistening) by Spicer (1874 a, 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 conld 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 take persicus 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.

Fig. 1.

Fig. 2.

^> ^d*£Z$

"~*X!

■

-" .* •* -'• £ . - • . <"*

^2V- ••■••' :"•■•* "•* •■■

*

\\

Fig. 4-
Fig. 1. A. reflexus s , x 10.

Fig. 3.

Fig. 5.
Fig. 3. A . persicus i , x 9.

Fig. 2. A. reflexus ? , x 10.

(Wheler, 1906, figs. 6, 7. 39.)

Fig. 4. A vespertilionis. Last nymphal stage, dorsal aspect, x 16. (Specimen from Cape Colony,

LouiiBbury Coll.)
Fig. 5. The same, ventral aspect. (Original, C. and R. phot.)

Genus Argas

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.— Monioz, 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).—Donitz, 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.
Khynchoprion 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 lihynchoprion 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.— Bisso, 1826, p. 182.— Heyden, 1826, p. 613.— Latreille,
1835, p. 511.— Gerstaecker, 1863, p. 343. (See also Audouin, 1832, p. 413.)—
Duges, 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.)
" Bhyachoprion (sic) Hermann," of Duges, 1834 c, p. 14.

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

PAGE

1. persicus ......... 8

2. reflexus 22

3. transgariepinus 29

4. brumpti 30

5. aequalis 33

6. vespertilionis 34

There are four doubtful species :

7. cucumerinus (possibly = reflexus var. magnus) . . 28

8. hermanni (near to reflexus var. magnus) ... 27

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.)

6 Family Argasidae

The following species have been suppressed :

americanus, chinche, mauritianus, miniatus, radiatus, sanchezi =
A. persicuB ; columbae, magnus, marginatus - A. reflexus (including
var. magnus) ; kochi = A. transgariepinus ; decussata, elliptica, fischeri,
inermis, longimana, pipistrellae, pulchella = A. Jvespertilionis ; strogu-
loides Gervais, 1844, p. 231, is a purely nominal species (Nn., 1896,
p. 25). A./orskali 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 H. aegyptium according to Nn., 1896, p. 24, and 1901, p. 255.

Explanation of terms and signs used in the following diagnoses.

L. = the extreme length of the tick.

W. = its width at the widest part.

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

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

Discs=the 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 = the genital orifice of the female.

grooves = linear depressions. In Fig. 1 note pre-anal groove, post-anal groove
and post-anal median groove.

folds = prominent integumental ridges. In Fig. 1 note supra-coxal and coxal
folds.

camerostome = the cavity in which the capitulum lies.

hood— the 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 6ase = the basal ring, etc., of most authors.

hypostome = the "maxilla," "radula," "labium," or " Unterkiefer " of various
authors. The dentition is indicated by figures on either side of a vertical
line. Thus 3 1 3 means three longitudinal files of teeth on each half of the
hypostome.

chelicerae = thc "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 process^" internal apophysis" of some authors.

2. external article = " 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, prolarsus 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.

.--V

preanal groove

transversa
postanal groove

Supracoxal fold

genital orifice

eye

coxal Fold

spiracle

median
^'-postanal
groove

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

Palp article*
Art.?

S-External j of Chelicera

hypostome
-Shaft of

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

8 Genus Argas

Key for the determination of species of Argas.
(Exclusive of doubtful species q.v. p. 5)

(a)

(Body circular . . . .

\ „ oblong .

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

. . [Body sub-conical in front
\ „ rounded in front

(Dorsum with polygonal depressed areas
1 „ without „

/Body scarcely narrower in front, capitu

lum very small and posterior
Body narrower anteriorly, capitulum
large and more anterior

vespertilionis
b

PAGE

34

c

persicus
d

8

e

brumpti
aequalis

30
33

. transgariepinus

29

refleocus

22

ARGAS PERSICUS (Oken) 18181.

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-Meneville, 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 (PL I, Fig. 4, legend states it
is a (J but text states <$ is unknown. Description insufficient, 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 americanus has to lapse. Again, Amblyomma
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 nigua de Geer)

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

Argas persicm 9

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

Argas sanchezi Alf. Duges, 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 J 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"1 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) ; "Melleh"
(Dupre, 1819, p. 324), "malleh" (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. reflexus.
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,
PL IX, Fig. 8, dorsal aspect, very good considering the date when it was drawn.
— Guerin-Meneville, 1829-43, PI. VI, Fig. 3, of A. mauritianus dorsal aspect,
Fig. 3a part of capitulum. Both figures inaccurate. — Koch, 1847, PI. I, Fig. 4.
— Kollar2 (date?) is stated by Heller, 1858, to have figured the capitulum. —
Hellkr, 1858, Pis. I-IV, deals chiefly with the internal anatomy. — Packard3,
1872, Fig. 68, referred to by Neumann, 1901, p. 255 (A. americanus). — Megnin,
1880, PI. VI, Fig. 3 (poor, A. mauritianus). — Laboulbene and Megnin, 1882,
PI. 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. — Duces3,
1891, Plate referred to by Neumann, 1901, p. 255 (A. sanchezi). — Marx in
Osborne, 1896, PL III, Figs. 3-3k. Drawings by Marx of A. americanus ?
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-
stome, 11 tarsus IV ; Figs. 16-21 (A. sanchezi), details of integument, digit,
hypostome, tarsus I, larval hypostome and ambulacrum. (Here reproduced,

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

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

3 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 and 20, which duplicate the others since the species
americanus and sanchezi no longer hold.) — Woods1, 1898, Figs. 20a, b, referred
to by Salmon and Stiles under bibliography (1901, p. 402) A. americanus.—
Hassall, 1899, Coloured Plate XVI (Argas americanus) of <j> , 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 PL LXXVIII, $, reproduction of Hassall, 1899, PL 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 Halter'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 J (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.— Frogqatt, 1906,
Figs. 1 and 2, showing Argas in crevices of splintered wood, and mouth-parts
of tick.— Wheler, 1906, Fig. 39* the £, dorsum.— Donitz, 1907, PL II, Figs.
9, 10 J , 13, 14 larval leg.— Reaney, 1907, p. 401, two very bad figures of larva
and female.— Manson, 1908, PL IV, coloured figure of adult (!) dorsum.—
Nuttall, 1908, Fig. 4 <? capitulum*.— Banks, 1908, PL I, Figs. 1, 2, legs 1 and 4.

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

1818. Oken, pp. 1567-1570.— 1819. Ddpre, p. 323, refers to effects 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. Porter2.— 1823. Fischer de
Waldheim, pp. 269-273, gives brief description, dwells especially on effect of
bite.— 1828. Szovits2.— 1829. Latreille, b, p. 289. O.— 1829-43. Gderin-
Meneville, p. 17, only refers to plate and habitat (A. mauritianus).—
1833. Cuvier, cited by George, 1876, p. 224, as referring to A. persicus.—
1844. Duges, cited by Laboulbene 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. O.— 1849. Lucas, H., p. 317. O.— 1855. Kuchen-

1 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. persicus 11

meister, p. 423. O.— 1858. Heller, C, pp. 297-326. Distinguished the sexes
and describes and figures the <J 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. Laboulbene, A., p. 54, obviously compiled from Fischer de Waldheim,
1823 (q.v.). — 1872. Packard, pp. 740, 741 (A. americana), cited by Salmon
and Stiles, 1901, p. 402.— 1873. Fumouze, inaccessible1. — 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,
1). 171.— 1875. Fritsch, p. 611.— 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. — Ernouf, p. 2561. — Megnin, P., p. 134. More fully
dealt with in Laboulbene and Megnin, 1882. — Goudet, J., cited by Megnin
vl880) as having described and figured "A. americanus 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
Laboulbene, 1881. — Laboulbene, A., (27, vii). Report from Tholozan. Cited
in full in Laboulbene and Megnin, 1882, p. 327. — 1882. Bordier, p. 131, cited
by Nuttall, 1899, p. 47.— Cobbold, p. 77. — Laboulbene and Megnin, p. 327,
deal at length with A. persicus, very diffuse, omit many essentials but describe
development and sexes. — Perroncito, p. 450. O. — Tholozan, p. 15x. — 1885.
Megnin, p. 460 {A. americanus)2. — 1890. Blanchard, ii, p. 883 et seq. — 1892.
Marx, p. 234 (A. americanus)2. — 1893. Hoehr, p. 348 (A. americanus)2. —
Railliet, p. 718 (A. radiatus). — Riley and Howard, p. 267 (A. americana,
also A. americanus)2. — Webster, p. 149 {A. americanus)2, inaccessible. — 1894.
Francis, p. 452 (Argus americanus)2. — v. Linstow, p. 120. O. — 1895. Braun,
p. 258. O. — Howard, p. 41 7 (A. americanus) 2. — Lounsbury, p. 657. — Marx, p. 199
(A. americanus)2. — 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. americanus)2.
— 1899. Hart, p. 180 (A. americanus) 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)2. — Nuttall,
pp. 46, 47, cites older literature regarding effects of bite. — Taschenberg, 1900,
p. 738. O.— Utility, p. 185, inaccessible2.— 1900. Ward, H. B., p. 197, quotes
Packard's description; mentions "A. sanchezi" as another species. — Ward,
H. B., p. 437 (A. americanus). — 1901. Froggatt, p. 1349, inaccessible, cited
in Hunter and Hooker's Bibliogr. — Neumann, pp. 255, 256. — Perroncito,
p. 561. O. — 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.) —
MARCHOUxandSALiMBENi, p. 569. Regarding spirillosis in fowls.— 1904. Banks,

1 Cited in bibliography by Huber, 1899.

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

12 Genus Argas

p. 331. O. Described as "A. miniata." — Lounsbury (November. Repr., brief
mention), see Lounsbury, 1903. — Mosler and Peiper, p. 346. 0. — Neveu-
Lemaire, p. 154. O. — 1905. Borrel and Marchodx, p. 362. Regarding
spirillosis in fowl. — Neumann, p. 240. — Nuttall, p. 22. O. — Robertson, p. 561,
quoted by Hunter and Hooker, 1907. — 1906. Braun, p. 371. O.— Froggatt,
pp. 14-18 ; gives general account of life history 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. O.— Nuttall, p. 394 et seq.— Banks, p. 15.

(Not "Argas 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 Amblyomma 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, PL I, Fig. 3.)

Adults : — Body ovate or quite oval, widest towards the posterior end.
Margin (150-200 fi 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 /j.) crescentic, equal to about half the width of the
anal ring; genital opening between coxae I and II1. 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 b). Hypostome indented, 6 or 7 fine
denticles on each half distally, followed by stout teeth 2 | 2, the numbers
of teeth increasing to 3 1 3, 4 j 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 J1 — which is slightly narrower in front

(ovate, PL 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.

persicas

13

immediately behind which it lies in the ? (it is situate between
coxae II according to Laboulbene and M^gnin, 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 £ 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 7 x 5 to 10 x 6 mm. (Nn. 1896, p. 8) ; we have seen specimens
from Merced, California, measuring 10'4 x 63 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. long1.

Fig. 3. Fig. 4.

Figs. 3, 4. A. perticus. 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.

Fig. 6.

Fig. 7.

Fig. 5. A. persicus. Portion of integument from posterior dorsal border of female,

x 44. Original N.
Fig. 6. A. persicius. 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.

Fig. 8.

Fig. 9.

* *»• "• r lg. ».

Fig. 8. A. perticus. Integumental hair, x 500. N.
Fig. 9. A. persicus. Anus, x 95 (Nn. 1896, fig. 8, of A. 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
(b, Fig. 23) the discs are more marked, though not as distinct as in
adults ; when full-fed they measure 55 to 6'7 mm.

Fig. 11.

Fig. 12.

Fig. 10.

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

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

Fig. 13.

Fig. 13. A. persicus i capitulum, ventral aspect,
Fig. '

. 13. A. persicus i capitulum, ventral aspect, x71. Indian specimen. C
. 14. A. persicus ? right palp in profile, external aspect, x 75. American
Original N.

Fig. 14.

Indian specimen. Original N.
specimen.

16

Genus Argas

Fig. 15. Fig. 16. Fig. 17.

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

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

fig. 10 of A. americanus respectively).

Fig. 18. A.persicus. Adult, tarsus I (A. miniattis). Salmon and Stiles, 1901
(fig. 74, 17th Ann. Bep., B.A.I., U.S. Dep. Agr.).

Fig. 19.

Fig. 20.

Fig. 21.

Fig. 22.

Figs. 19 — 22. A. persicus 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 (1st 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.

perswus

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

18

Genus Argas

Larva: 680-770 /u, in length (minus capitulum), hexapodand 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. Rep., B.A.L, U.S. Dep. Agr.).

A. 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, -6 to '8 mm. in diameter,
apparently without external structure.

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 anmdatus 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 Argas

Geographical distribution.

A. persicus 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 (Oudemans Coll.).

Asia: Persia. Oken (1818) and Dupr6 (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. pei-sicus. 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 specimens 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 J 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 South 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. persicus 21

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

America : It (A. 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 1902a, p. 568) reports "A. sanchezi"
from Catalina Springs, A rizona (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 Merced1 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 Hart2 (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). New 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.

3 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, PL I, Figs. 1, 2.

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

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

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, N and 0, dorsal and ventral aspects, capitulum and tarsus (very
crude figures). — Duges, Ant., and Milne Edwards, PI. 27, Fig. 2 (poor). —
Gerstaecker, 1860, PI. XV, Figs. 1-6.— Pagenstecher, 1861, PI. 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., 1874 a, 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. — Megnin, 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 (?)). — Claus, 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 ? dorsum and
venter (sketchy). — Ddbreuilh and Beille, 1896, Fig. 6 (sketchy). — Neumann,
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.— Taschenberg,
1900, p. 739, dorsum and venter of adult (?) (inaccurate). — Wheler, E. G., 1906,
Figs. 6, 7, $ dorsum, $? venter, both good photographs ; here reproduced.

Literature: 1794. Fabricius, pp. 426, 427. Ac. reflexus, Ac. marginatum, Rhync.
columbae (few words). — 1804. Hermann, p. 69. — 1805. Fabricius, pp. 353, 354,
356 (Latin). Same as 1784.— (No date) Duges, Ant., and Milne Edwards
(no date).— 1837. Walckenaer, hi, p. 231. O1.— 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 ; effects 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). O.— 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. O. (b) p. 209. O.— Taschenberg, p. 171 et seq. (mostly citations
from other authors).— 1877. Canestrini, G, and Fanzago, p. 125. — 1879.
Boschulte, p. 562. — 1880. Megnin, p. 135 ; repeated verbatim in Laboulbene
and Megnin, 1882, p. 322.— Rivolta and Delprato, p. 314.— 1882. Laboulbene
and Megnin, pp. 332, 340. — Chatelin (see re effects of bite). — Contarini,
pp. 16, 161 (cited by Berlese, 1888).— ZOrn, 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.—
Megnin, 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). O.— 1895. Braun, p. 258. O.— Planchon, p. 29 (inaccessible).— 1896.
Dubreuilh and Beille, p. 76. O.— Neumann, p. 4.— Moniez, pp. 499-509. —
Gibert (thesis).— 1897. 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). O.— 1901. Neumann, p. 253.— Perroncito, p. 568.— 1904.
Mosler and Peiper, p. 345. O.— 1906. Tonnel, p. 552.— Braun, p. 369. O.—
1907. Donitz, p. 26.— 1908. Schnee, p. 32.— Schellack, p. 486 ; transmission
of spirochaetosis. — Banks, p. 16.

Adult: (Figs. 27, 28, PI. I, Figs. 1, 2). Body oval, widest toward

the posterior end. Margin irregular wrinkled into radial striae forming

a border 200 /j. wide, which is slightly turned up especially in unfed

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

1 Papers marked O 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 A. persicus.
Capitulum: 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 1 2 stout teeth merging into 3 1 3 to 6 1 6 progressively

Fig. 27. Fig. 28.

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

Fig. 29.

/

■ t\\J

0A

/~^r"~

w <_ w w 1

::_-'

:--■'■'

Fig. 30.

Fig. 31.

Fig. 32.

Fig. 29. A. reflexm. Palpal hair, x 250 (Nn. 1896, fig. 2).

Fig. 30. A. reflexws Tar. magma. Left digit, x 225 (Nn. 1896, fig. 14, modified).

Fig. 31. A. reflexus. Left digit, dorsal aspect, x 220 (Nn. 1896, fig. 1, modified).

Fig. 32. A. reflexus var. magnus. Hypostome, x 100 (Nn. 1896, fig. 15).

A. reflexus 25

smaller teeth. Legs as in A. 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 A. persicus.
The male (Figs. 27, 28) may attain 4x3 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.

J-Z

A

V V v \ v-

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

Nymphs measure 4x2 mm. (1st stage) or 55 x 42 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 Genus Argas

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. refleams 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
Donitz (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 Fiilleborn (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. reflexus 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 (1873 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. re/Saras).— 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 fi wide.
Venter: spiracles about two-thirds (150 (jl) the width of the anal ring.
Anal aperture bordered by a few short hairs. Capitulum relatively

28

Genus Argas

small (275-300 /j. 1.) ; 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. 34. Left digit, x 225 (after Nn. 1896, fig. 12, modified).
Fig. 35. 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 Raffray (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 rejlexus
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, 10x5 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. cucumerinus

29

the space between the coxae of the two sides equal to J 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.

Resembling 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. 37.

Figs. 36, 37. Argas transgariepinus S , x 4. Dorsum and venter,
specimen in Brit. Mus. Original, N.

Sketch from type

30

Genus Argas

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 10 x 7 mm.,
a smaller specimen measured 8"5 x 6 mm.)

Fig. 88. Fig. 39.

Figs.J38, 39. Argas brumpti ? , 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, Figa. 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. brumpti

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. 40. 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 hrumpti.

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

A. brumpti 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. brumpti %
(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 aequalis Neumann, 1901, p. 259.
Argas aequalis (Nn.) Neumann, 1908, p. 26.

Nymph : Body flat, 5 x 25 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 | 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
Fulleborn (Berlin Mus.). This species is unfortunately not figured.
(See p. 3, footnote 1.)

34 Genus Argas

ARGAS VESPERTILIONIS (LatreiUe), 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 Iconography).

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

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

Argus pipistrellae in Ant. Duges and Milne Edwards (no date).

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

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

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

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

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

A rgas fischeri George, 1876, pp. 223-225, in Westwood, 1877, p. lxii, referring to
George's paper. Probably nymphs1.

Argas pulchella 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. lxii.

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

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

Iconography2: 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. Duges 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. lxii), 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."

* Figures marked * are reproduced herein.

A. ve8pertiUonis 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 W.

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 from 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
integument, 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 vetpertilionis. 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.

Fig. 50. Fig. 51.

Argas vespertilionis.

Fig. 50. Nymph, ventral aspect, x28 (Nn. 1896, fig. 22).
Fig. 51. Nymph, ventral aspect, x 13 (Wheler, 1906, fig. 8).
We regard these as 1st and 2nd stage nymphs.

A. vespertilionis

37

Nymph (Figs. 50-52) : Body nearly circular, 2-4 x 2'1 mm.
(3-7 x 3-8 mm. to 495 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 vespertilionis. Nymph's intestinal caeca.
After George, 1875, pi. II, fig. 2.

Fig. 53.

Fig. 54.

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, x 585.
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^. w.), less than half the
width of the anal ring (130 x 90 /*). 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 1 2 rows of 5 or 6 teeth ; digit, see Fig. 56 ;

38

Genus Argas

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 vespertilionis. Larva: A. hypostome, x575; B. digit, x865; C. dorsal
plate, xllO; D. anus, x370 (Nn. 1896, fig. 26).

Larva (Figs. 57 A-D) : Body short oval, 1*3 x 1 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. Gapitulum
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. vespertilionis 39

It is probably Argus fischeri Audouin. (Caris elliptica Kol. was found
on Myotus murinus ; Cans longimana Kol. on Miniopterus schreibersi
and Rhinolophus clivosus ; Caris decussata. Kol. on Myotus murinus and
Rhinolophus hippocrepis ; Cans inermis Kol. on Brachyotus dasycnemus
and Synotus barbastellus.)

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. pipistrellus. 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.: Ornithodoros 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. — Lounsbury,
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 (Ornithodoros) Murray, 1877, p. 183.

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

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

Ornithodorous Koch : Morgan, 1899, p. 136 (sic).

Alectorobius Pocock, 1907, p. 189 (see Ornithodoros talaje).

Ornithodorus of numerous writers since Koch.

Generic Characters. Body flat ivhen 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 Ornithodoros

adult. Discs1 present or absent ; but when present, not arranged radially.
Certain 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
Tarsus I. Tarsus IV.

turicata

talaje

T

talaje

lahorensis

3T

Lakorerisis

megruru

^u

meg mm

V

jD

fumaai furcosus

Fig. 58. Original, N. and W.

' Structures analogous to the " discs " of Argas are probably to be found in all species
of Ornithodoros 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.

Genus Ornithodoros

41

transverse pre-anal and a transverse post-anal groove or furrow, and a
post-anal median groove1. 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 .

42
46
55
57
59
62
63
65
67
70
71

78
79
80

Doubtful species are :

12. canestrinii ........

13. papillipes

14. morbillosus

Suppressed are:

americana = turicata

miliaris = erraticus

coniceps • = talaje var. coniceps

rudis = talaje

reticulatus Gervais, 1849-51, in de Gay, Zoologie, p. 44,
PI. 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 Omithodoros

Key for the determination of species of Omithodoros.

(a)

(b)

(c)
(d)

(•)

(/)
(9)
(h)

(i)
U)

(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 discs1 ....
(without obvious discs1

(with movable cheeks to camerostome
(without movable „ „

.''coxae II larger than I ; with tarsi forked

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

I tarsi slightly humped

(tarsi II — IV with prominent dorsal humps

p. 41.)

b.
c.

d.
e.

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

pavimentosus, p. 62.
/•

moubata, p. 46.
9-

savignyi, p. 42.
coriaceus, p. 55.
h.
i.

talaje, p. 59.
3-

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 Audouin, 1827, p. 183.
Omithodoros savignyi C. L. Koch, 1844.
1 0mithodoros morbiUosus Gerstaecker, 1873 (see p. 80).
Argas schinzii Berlese, 1889.
Native names : see 0. moiibata, 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. savignyi ? . Dorsal aspect, x 8. Specimen from S. Africa.
Original, C. and B. phot.

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

O. savignyi 43

Lit. and Icon. : Savigny (1826, PI. IX, excellent figures of dorsum, venter, legs ;
less good of capitulum and mouthparts), reproduced by Walckenaer and
Gervais (1837-47, PI. XXXI, Fig. 2).-Guerin-Meneville, 1829-43, PL VI, Fig. 4,
mouthparts taken from Savigny without acknowledgement. — (J. 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). —
Laboulbene 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.
O. — 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 (PI. 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 /x in
diameter, PI. 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

<3

■■

Fig. 59. Fig. 60.

Fig. 59. O. savignyi. Granulations of integument, x 65 (Nn. 1896, fig. 27).
Fig. 60. O. savignyi. A "fossette" or disc, x23o (Nn. 1896, fig. 28).

numerous on the venter. Venter (PI. 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 IV1 ; 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 fi). Spiracles above the supra-coxal folds, 600 fj. wide,

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

44

Genus Ornithodoros

crescentic. Capitulum stout, free; palps (Fig. 61) tapering, articles
1 and 2 equal in length ; article 3 the shortest ; hypostome (Fig. 62)
short with dentition 3 1 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 p (Nn. 1896, fig. 31).
Fig. 62. 0. savignyi. Hypostome, x 65 (Nn. 1896, fig. 30).
Fig. 63. O. savignyi. Digit, 200 ju I. (Nn. 1896, fig. 29).

the two proximal near together, an interval three times as great
separating the second and third. The </ and ? resemble each other
except in the generally smaller size of the £ and of the </ genital
orifice1. Many long blunt hairs thickly beset the anterior margin
of the body giving it a downy appearance.

1 See Figs. 66, 67 of O. moubata.

0. 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 (B), I (C), x 17 (Nn. 1896, fig. 32).

Fig. 65. 0. 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. moubata 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 moubata Murray, 1877, p. 182.

Ixodes moubata, Cobbold, 1882, p. 78.

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

Native names1: bibo, Toro name in Uganda (Christy, 1903, p. 187). — Suaheli terms
are pasi (plural papasi) and Kupe (Wellman MS.). — gourat in Nubia and Soudan ;
mechger in Abyssinia ; courdoud in Galla ; oldirrho in Kilima-ndjaro (Brumpt,
1901, p. 578). — papazi in Mombassa (Karsch, 1878, p. 311, who calls it
0. savignyi). — tampan or tampao 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 mbu2

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

2 Also applied to mosquito and to malaria, according to It. Koch, in parts of E. Africa.

0. moubata 47

in Tete (misspelt " bu " in Manson, 1903, p. 714).— carrapato1 in Portuguese East
Africa (a loose term), Livingstone (1857, p. 382) spelt it carapato, often misspelt
garrapato. Wellman (in MS.) states that karapatti and garapata are Arabic and
Bantu corruptions of the Portuguese word. — tapazi at Nyangwe, Lualaba river,
Livingstone (1874, n, p. 115). — bifundikala about Leopoldville Congo (Dutton
and Todd, 1905 b). — bimpmi is the BaWke' name (Dutton and Todd, 1905 b). —
mouyata at Popokabaka (Dutton and Todd, 1905 b). — moubata in Angola
(Murray, 1877, p. 182). According to Wellman (in MS.) Dibata (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). — livmno* is the Lovale
name in E. Angola and N. W. Rhodesia (Wellman, 1908, personal communica-
tion).— kimputo, common name in Eastern Congo Free State (Dutton 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.— Dutton 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. Donitz, plate
showing tarsi I and IV*. — Wellman, p. 155, Fig. 2 ; dorsal 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, vn, 1908, Fig. 1*.

Lit.: 1857. Livingstone, pp. 382, 628.— 1874. Livingstone, Vol. n, 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. O.— 1895. Dowson, p. 1201,
sent specimens from Tete to British Museum ; describes effect of bite, fever,
etc. — 1896. Neumann, p. 30. — 1900. Pocock, p. 222, first described some of
the specific differences 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
Dowson had also sent specimens of 0. moubata from Tete, Zambesi. — 1905.
Caiman, p. 124, determined specimens sent by Wellman from Angola. — Dutton
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 moubata and partly to one of the
Ixodidae. — Kerr, p. 126, criticism of Feldmann's observations. — Newstead,
pp. 1696, 1697.— Wellman, repr.— 1906. Donitz, 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 (Ricinus communis) according to
Wellman (ms.).

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

48 Genus Ornithodoros

This species closely resembles 0. savignyi, from which, however, it may
easily be distinguished by the absence of eyes, and the details of its leg
structure {compare Figs. 70-73). The humps on theprotarsus 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 — 51. Less hairy than
0. savignyi1.

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. (Donitz, 1906 ; Wellman, 1906), or
even 12 x 10 mm. when fully grown and filled with eggs and blood
(Button and Todd)2. The colour varies from 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^

2 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. moubata

49

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 s . Anterior portion of venter. Original, N. and W.

Fig. 68. 0. 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 25 mm.,
whereas unfed nymphs of the second stage average 14 x 1*2 mm.

Fig. 70.

Fig. 72.

Fig. 71.

Fig. 73.

Figs. 70, 71. 0. 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 1-8 mm. I.

(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 J 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 ecdysis1 and may
double in size after feeding.

Fig. 74. Fig. 75.

Fig. 74. Nymph, 1st stage, (. l-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 very

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

4—2

52 Genus Omithodoros

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 stage1, 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. 0. moubata. Capitulum of larva, ventral aspect and dorsal aspect of left palp.
Whole length of capitulum including palps, 180 /i. 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. (Dutton 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 R. Koch (cited by
Dbnitz) 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 O. savignyi.

O. moubata 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 Africa 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, n. 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 Ornithodoros

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."

O. moubata

55

See further under Section II, regarding observations on the hatching
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, ? (coloured) ; Fig. 3, ventral aspect in out-
line (very good figures).— Nuttall, vn, 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, 31; long white hairs on dorsal sur-

Fig. 81. Fig. 82.

Figs. 81, 82. O. coriaceus ? , 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 Omithodoros

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.

83.

Fig. 84.

Fig.

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.1

Fig. 84. 0. coriaceus <? , 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 93 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 around2; 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

1 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.

5 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 California1 were collected
by Mr H. A. Moran at Los Olivos in August, 1904, and comprised
7 c/1,/ and 9 ? ? (Stanford University Coll.); the largest $ measured
138 x 8'2, the smallest 9'5 x 53 mm. ; the largest </ measured 8'6x46,
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 £). Both females were received alive and
the better specimen was drawn, as also the <f (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. Dug£s), 1876.

Figs. 58, 85-87.

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

Lit. and Icon.: Duges, 1876.— Megnin, 1885, p. 463, PI. XX, Figs. 1-4— Marx, 1895,
posthumously published plate, appeared in Osborn, 1896, PI. Ill, Figs. 2, 8t;
good figures of <J dorsum, venter, etc. — Neumann, 1896, p. 31, Fig. 33, digit*. —
Ward, x. 1900 a and b. O.— Braun, 1906, p. 373. O.— 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 fi broad. Spiracles circular (180
to 200 (x 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 <j . L. 3'5 mm. Dorsum and venter. Specimen
from Guanajuato, Mexico. Original, N. and W.

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 (J 230 fi,
? 300 ft) and articles 3 and 4 equal (</ 150 fj,, ? 170 /a); numerous
pennate hairs on dorsal surface 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. turicata 59

Males average 3-5x 25mm.; females 6 x 4mm.; nymphs 22 xl-5 mm.

Neumann's description, from which the above is partly taken, is
based on numerous specimens collected in Guanajuato, Mexico, by
A. Duges1.

Fig. 87. 0. turicata. Digit, 90 m 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. americanus of Marx was found in
Texas on the nostrils of the horse, in South America on the llama
and in Florida on the tortoise, Xerobates 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 (Gu(§rin-M£neville), 1849.

Fig. 88.

Synonymy : Argas talaje GueVin-Meneville, 1849, pp. 342, 343.

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

(lArgas coniceps Canestrini, 1890, p. 535. Regarded as A. reflexus <J 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 : Gudrin-Meneville, 1849, PL II ; figures inaccurate, repro-
duced by Megnin, 1885, PL XX, and by Murray, 1877, p. 183. — Canestrini,
1890, PL 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. O.— 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

Genus Ornithodoros

Body : L. 5 — 6, W. 3 — 35 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 /a 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 /m), the valves bearing two to
four hairs each. Spiracles conical, 100 fi 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. O. talaje. A, hypostome (x 160). B, digit, 60 p. I. (Nn. 1896, fig. 34).

base of capitulum finely wrinkled transversely, permitting (according to
Megnin) the mouthparts to be protruded and retracted. Chelicera :
digit 60 n I. (Fig. 88 B) with tridentate dorsal process near base of
internal article and external article bidentate. Hypostome (Fig.
88 .4) 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 /j, 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 III

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

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

O. talaje 61

Venezuela (Paris Mus.), and two specimens collected by Salle" 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 Granada by Goudot. According to Neumann
(1901, p. 258) the Paris Museum possesses specimens collected by Gay
at Santiago de Chili 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. I.; 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. reflexus, in the interstices of the mosaic of the Church of San Marco,
Venice. Birula states that several specimens taken near the Sea of Aral
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 (?ape 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 Genus Omithodoros

Museum possesses numerous specimens of nymphs and adults collected
by the Challenger Expedition, 28, vm. 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 (xn. 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*.— Dbnitz, 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, flat 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 dried1 ? 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 DSnitz). According to Donitz, who has examined many

1 This may account for the " black " eyes.

0. pavimentoms

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, 1906, pi., fig. 2.)

£%

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

ORNITHODOROS ERRATICUS (Lucas), 1849.

Syn. and Lit. : Argas erratiacs Lucas, 1849, p. 316, gives a fairly good description,
considering the date.
Ornithodoros miliaria 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 Ornithodoros

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 fi ; anus with hairs on the valves.
Spiracles semicircular (?), 100/* wide. Capitulum free, exposed ven-
trally ; palps 1, 4, 2, 31 ; 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 1 3, then 4 1 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, 35 mm. I., found by Doria in the vicinity of Tunis, one
from Nemours and two from Marnia(?). Lucas received his three
specimens (4 x 25 mm. in size) from the plain of Lake Houbeira, near
La Calle. They were found beneath stones where Bufo pantherinus was
also in hiding. Karsch's 0. miliaris 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.

1 The relative lengths of articles.

0. tholozani 65

ORNITHODOROS THOLOZANI (Laboulbene and
Megnin), 1882.

Figs. 93, 94.

Syn., Lit. and Icon. : Argas tholozani Laboulbene and Megnin, 1882, pp. 335-337,
Pis. XXII and XXIII ; the figures include : adult dorsum and venter, capitulum,
hypostomes ( $ and $ ), 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 papillipes Birula, 1895, pp. 38-44, Fig. 35 ; digits of <$ and $ ; referred here
by Neumann, 1901, p. 259. See 0. papillipes, p. 79.)

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

Adults : Body (the </" varies from 4x2 mm. to 6 x 4 mm. ; the ?
from 8x4 mm. to 10 x 5 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 fi 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 /i); anus with seven or eight long hairs on each side. Spiracles
(350 fi), crescentic. Capitulum : palps tapering, and not much longer
than the hypostome; chelicerae (digits, Figs. 93, 94). The hypostome
differs in the sexes : in the J1, 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 far apart ; then three or four rows of simple
squamiform teeth ; in the $ , it is somewhat lanceolate, and the dentition

N. I. 5

66 Genus Omithodoros

is 2 1 2 throughout, the median rows being far apart. Four long hairs
(post-hypostomal 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. Fig. 94.

Figs. 93 and 94. 0. tholozani. Left digits of g 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, -8x-6mm.; about 100 laid by each $
(Laboulbene and Megnin).

This species is allied to 0. erraticus, 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 Me*gnin'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. lahoremis

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., % 10 x 5'6 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.

O. lahoremis ? . 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. Venter : 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,

5—2

68

Genus Ornitlwdoros

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

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

Fig. 98.

Fig. 99.

r

r^rx;

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. O. lahorensis. Adult, tarsi I— IV, x 28. Nn. 1908, fig. 15.

IV

O. lahorensis 69

Capitulum ( ? 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 fi, 250 /x, 150 fi and 200 fi 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 : 5-5 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
orifice1 ; 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 £ and % . The £ 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.

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

70 Genus Ornithodoros

ORNITHODOROS PURCOSUS Neumann, 1908.

Fig. 101.
Lit. and Icon.: Neumann, 1908, p. 21, Fig. 16 (here reproduced).

Female: 10 x 5mm.; 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. 1.) ; 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— l^^^J

r

ii.

IV.

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

O. megnini 71

ORNITHODOROS MEGNINI (Duges), 1883.

Figs. 102-112.

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

" Argas americana Packard " : Townsend, 1893, p. 50.
Rhynchoprium spinosum Marx, 1895, p. 199.
Omithodoros megnini (Duges) Neumann, 1896, p. 42.
The " Spinose Ear Tick " of American authors.

Lit. and Icon.: Duges, 1883, p. 1951.— Megnin, 1885, p. 460, PI. XXI, Figs. 1-8.—
Neumann, 1888, p. 96* ; 1892a, p. 101 - ; 1892b, p. 105.— Blanchard, 1890,
p. 883 et seq. O.— Railliet, 1895, p. 718.— Townsend, 1893, p. 49.— Dolly,
1894, p. 9802.— Marx, 1895, 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, 1900 a, 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 (?) natural size and x 2),
two figures after Neumann's Fig. 36 ; Fig. 98, integument of nymph ; Figs. 103,
105-107 represent 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. O.— 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 Ornithodoros

well marked. Spiracles circular, 250 fi in diameter, stigmal plate
semilunar (resembling Fig. 10). Anus : anal frame broader than long
(150 to 200 fix 200 to 250 /*), each valve provided with four short
hairs. Eyes 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. O. megnini <r , x 8. Dorsum and venter. From specimen, loaned by
Prof. Neumann, from New Mexico, U.S.A. Original, N. and W.

Fig. 104. O. megnini. A, digit of nymph, 85 /j. 1. B, digit of ? , 75 /i 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.

0. megniiii

73

Hypostome short, broad basally, rounded distally, unarmed1. Palps thick,
300 /j. long (90 fi, 85 /*, 65 /j,, 60 //,, from first to fourth article), with
articles relatively broad and short, the second 110 fj. 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 : ? L. 5 to 6 mm., W. 3
to 4 mm. ; </" somewhat smaller. We have received 2 £ and 1 % from
Texas (W. D. Hunter), the £ £ measured 6'7 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 la. 17th Ann. Hep.
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-

1 Note observations on this point in Section II.

74

Genus Ornithodoros

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. O. megnini. Unfed. Nymph, ventral aspect. Salmon and Stiles, 1901, 6g. 100.

17th Ann. Eep. B.A.I., U.S. Dept. Agric.
Fig. 108. O. 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 /x) than long; two long hairs on each external border. Spiracles
300 to 350 fi in diameter, on salient tubercles1 (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 fM long (from base of palps to tip of hypostome) ; chelicerae (see
Fig. 104 .4) ; hypostome lanceolate, dentition 4 | 4, with 7 to 9 teeth per
file ; palps thick (articles 1 to 4 measure 140, 125, 110, 180 fi) ; second
article 110 fi 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.

Larva1 (Figs. 109-110) : hexapod, elongate oval, with long terminal
capitulum, measuring 061 mm. from tip of hypostome to posterior ex-
tremity. Integument finely striated with symmetrically arranged pits,
from which bristle-like hairs issue. Capitulum : hypostome 2 | 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. 111).

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

1 Also described by Stiles and Hassall, 1901.

76

Genus Ornithodoros

Fig. 110.

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

0. megnini

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. ill.
0. megnini.

Fig. 112.

Fig. 111. 0. megnini. Leg of larva, tarsus I in profile.
Fig. 112. 0. megnini. Larva, gorged, so-called "resting- or pupa-stage."
Both figures from Salmon and Stiles, 1901 (figs. 108 and 104.
17th Ann. Rep. 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 Ornithodoros

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 5 mm., $14x8 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 f 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 J 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. canestrinii

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, PL 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 j 2, about
as long as the base ; palps short and thick : two long hairs at base of

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

80 Genus Omithocloros

hypostome ; 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 synonym
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 Me'gnin)." 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 morbillosus 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 type, states that
it differs in the leg structure as noted below.

Species based on a single specimen, measuring 6'3 x 43 mm., 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), brumpli (p. 95).

Ornithodoros savignyi1, coriaceus (p. 102), turicata (p. 102), talaje (p. 103), pavi-
mentosus1, tholozani (p. 103), lahorensis1, 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 \\ — 2 hours to gorge2. 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 facta known regarding the biology of these species are included under
Section I : savignyi (p. 42), pavimentosus (p. 62), lahorensis (p. 67).

2 See feeding experiments in Cambridge, noted below.

N. i. 6

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 l, with hood pro-
jecting quite over the capitulum ; measured
2 x l-5mm.

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 first 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 :

103

in

5

minutes

10

>»

15

»

20

;)

25

»

30

>>

35

„

40

3)

45

»

22<

4 fed in 50 minutes

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 »j 8*5 ,, ,, ,, 10 ,,

permits them to "crawl rapidly and to secrete themselves in cracks and crevices protected
from the wily fowl." The replete larvae usually drop at night when the fowls are roosting,
consequently when they emerge as nymphs they have no difficulty in rinding 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.
natcned 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

26.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. persicus 85

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

Seasonal Prevalence: Riley and Howard (1895, p. 267) 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 animals1.

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

1 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. persicus 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. persicm 87

Economic Importance: the effect of the bites of A. persicm 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 (".4. 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 of persicus 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 persicus
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 Fowls is a disease whose cause, the Spirochaeta
marchouxi1, 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. l& = Sp. gallinarum Blanchard, 1905.

A. persicm 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. marchouxi
the latter multiplies within the body of the tick (maintained at
30 — 35° 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 off 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
Sakharoff (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 " by 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 successfully
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. reflexus 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 Fullager (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 specimens were found in an abandoned pigeon-coop after the
lapse of 2 years.

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

Feeding Habits, Economic Importance, 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 time" 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-inksied 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 ;

92 Biology

Osborn, 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. Easpail (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 persisted 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 reflexus 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 marchouxi. 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 appeared 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 Sp. marchouxi may be transmitted by two species of Argas
(jpersicus 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. brumpti 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 reflexus were discovered, and, following a thorough
disinfection of the premises the trouble ceased. It appears clearly
demonstrated by this 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. moubata ; it may
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,
O. 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. moubata 97

4 — 6 months or more by Newstead, Dutton 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,
Dutton 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 ticks1 and eggs. Wellman (1906 and 1907) has seen Phonergates
bicoloripes Stal. attack and suck the blood out of 0. moubata, 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. Dutton 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 District, Angola, he observed what appeared to be their
destruction by a parasitic mould.

Feeding: Dutton 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 off. According
to Newstead, Dutton 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 moubata feeds for £ to 4 hours. Fed
on laboratory animals in Cambridge during the day-time, the adults

1 Wellman (in sis.) once saw a swarm of driver ants (Dorylus nigricans Illiger) bearing
away moubata from a native kraal in Angola.

n. i. 7

98 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 l-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. moubata 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 Nyafigwe\ 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 moubata 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 Africa1. 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 6%
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. moubata, 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.

O. moubata 101

Africa)1. When a female 0. moubata 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. Button and
Todd, in addition, found that the spirochaetes persist in the gut of the
tick up to 5 weeks after it has fed. The tick, once infected, may harbour
the parasite for months and transmit it when it has occasion to feed.
Finally, Mollers (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.
moubata is capable of transmitting Filaria perstans to man. In this
disease the filarial embryos circulate in the blood. Feldmann (1905,
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 Fowls. It is interesting to note that Ftilleborn 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: Me'gnin (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
original papers.

Aclogue, A. (1905), Les Argas. Le Cosmos, N.S., liii. 536 (inaccessible).

Agassiz, L. (1846), Nomenclatoris zoologici Index Universalis, continens nomina
systematica cla-ssium, ordinum, familiarum et generum animalium omnium,
tarn viventium quam fossilium, secundum ordinem alphabeticum unicum disposita
adjectis homonymiis plantarum, nee non variis aduotationibus et emetidationibus,
pp. viii + 393, 4°, Soloduri. (Only lists genera : Argas, Ixodes, etc.)

Ajutolo, G. d' (1898), Nuovi casi di Argas reflexus parassita dell' uomo. Rend.
Accad. 1st. Bologna, n. 222-223.

Alt, K. (1892a), Die Taubenzecke als Parasit des Menschen. MUnchen. med.
Wochenschr., xxxix. 531-533, 2 Figs. (Cited by Nuttall, 1899, p. 45, A.
reflexus.)

(1892b), Die Taubenzecke als Parasit des Menschen. Thiermed. Rundschau,

Halle a. S., vi. 265-269. (1 Sept. ; A. reflexus.)

(1893), Die Taubenzecke als Parasit des Menschen. Centralbl. f. Bacteriol.,

xiv. 468. (Rev. by Kniippel.)

Audouin, J. V. (1827), Description de l'Egypte, ou recueil des observations et des
rechcrches qui ont etc faites en figypte pendant Pexpedition de l'armee frangaiso,
publie par les ordres de sa majeste l'empereur Napoleon le Grand, ed. 2, xxii.
Zoologie.

(1832), Lettres pour servir de materiaux a l'histoire des Insectes. Premiere

Lettre, contenant des Recherches sur quelques Araignees parasites des genres
Pte'ropte, Caris, Argas, et Ixode, adressee a M. Leon Dufour. Ann. d. sci. not.,
Paris, xxv. 401-425, Pis. IX, Figs. 8-10 ; XIV, Figs. 1-4. (Argasidae and
Ixodidae.)

n. i. 8*

Bibliography 3

Austen, E. E. (1906), An insect enemy of the disseminator of human tick fever in
Angola. Joum. Trap. Med., Lond., DC. 1 13. (Describes Phonergates bicoloripes,
sent by Wellman.)

Banks, N. (1901a), Aoftrina in "Some Spiders and other Arachnid* from Southern
Arizona." Proc. U.S. Nat. Museum, xxm. 590. (A. pensions and Ixodes sp. not
identified.)

(1901 b), Some Arachuida from New Mexico. Proc. Acad. Nat. Sci., Phil-
adelphia, liii. 568-597. {A. persicus found at Derning.)

(1902b), Papers from the Hopkins-Stanford Galapagos Expedition, 1898-1899,

vol. vii., Entomological Results (6), The Arachnida. Proc. Wash. Acad. Sci.,
IV. 70, PL II, Fig. 9. (Argas transversa, n.sp., and mention of Amblyomma
pilosum Nn. both at Albemarle Island.)

(1904a), Some Arachnida of California. Proc. California Acad. Sci. (3), m.

331-376, PI. XLI, Fig. 57 (Zoology).

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

1-114, 201 text-figs. (Ticks : pp. 42-49, Figs. 71-85.)

(1907), A catalogue of the Acarina, or Mites, of the United States. Proc. U.S.

Nat. Mus., xxxn. 595-625.

(1908), A revision of the Ixodoidea, or Ticks of the United States. U.S. Dep't

of Agricult., Bureau of Entomology, Technical Series No. 15. 60 pp., 10 Plates.
Washington IXC. (issued 6. VI. 1908).

van Beneden, P. J. (1883), Animal Parasites and Messmates. 3rd ed., 83 Figs., 274 pp.

London : Kegan Paul, Trench and Co. (brief mention of Ticks).
Berlese, A. (1885), Acarorum systematis specimen. Boll, della Soc. Ent. Italiana,

xvn. 121-144.

(1888), Acari Austro-Americani. Bollettino della Soc. Entom. Italiana, xx.

193.

(1888-1892), Acari, Myriopoda et Scorpiones hucusque in Italia reperta. Padua

[Re Ticks : see Fasciculi 47 and 49 (1888), 55 (1889), 58 (1891), 61 (1892).]

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

x. 289-300, PL VII, Figs. 2-4.

Bianconi, G. A. (1867), Sid Rhynchoprion columbae Herm. o Argas reflexus Latr.

con una Tavola. Mem. dell' Accad. d. sc. dell' Istituto di Bologna (2), VII. 107-

112, 1 PL
Birula, A. (1894 and 1895), Ixodidae novi vel parum cogniti Musei Zoologici

Acaderniae Caesareae Scientiarum Petropolitanae. I. Bull, dc V Acad. Imp.

des Sc. de St Petersbourg, ser. 5, n. (4), April, pp. 353-364, Pis. I, II.
Blanchard, R. ( ), "Parasites Animaux" in Bouchard's Traite" de Pathol.

Generate, vol. II. Paris.

(1890), Traite de la Zoologie me'dicale, II. 883, Figs. 388-883. (Refers to

different Argasidae.)

Bocquillons ( ), Storia med. fra gli Aracnidi, p. 113. Cited by Bianconi, 1867,

p. 107. (Casual reference to Argas reflexus.)
Bordier, A. (1882), L'argas persicus. Joum. de The'rapeutique, ix. 131-133. (Cited

by Nuttall, 1899, p. 47.)

Bibliography r>

Borrel and Marchoux (1905), Argas et spirilles. Compt. rend. Soc. bid., lviii. :iu^

364. (A. persicus.)
Boschulte (1860), Argas rejlexus, als Parasit an Menschen. Virchovfs Archiv, xvm.

554-556. (Cited by Nuttall, 1899, p. 44.)

(1879), Ueber den Argas rejlexus. Virchow's Archiv, lxxv. 562. (Cited by

Nuttall, 1899, p. 40.)

Brandes, G. (1892a), Ueber die Argasidae. S.B. Ver. Sachs. Thiir, pp. 17 and 22.
(CSt. Zool. Record, 1892.)

(1892b), Ueber die Faniilie der Zecken. Dei: ii. d. Sitzungen d. naturforsch.

Gesellsch. zu Halle im Jahre 1892, 8-11.

(1897), Argas rejlexus als gelegentlicher Parasit des Menschen. Centralbl. f.

Bakteriol. u. Parasitenk., xxn. 747-752.

Braun, M. (1883), Die thierischen Parasiten des Menschen nebst einer Anleitung zur
praktischen Deschaftigung init der Hehninthologie fur Studirende und Aerzte,
pp. i-viii + 1-233, Figs. 72. (See Braun, 1895.)

(1895), Die thierischen Parasiten des Menschen: ein Handbuch fur Studirende

und Aerzte, Wurzburg, ed. 2, pp. 283, Text-Figs. 147. (Ixodidae, see pp. 257-
259, Figs. 127-130, none original.)

(XI. 1906), The Animal Parasites of Man. London : John Bale, Sons and

Danielsson, Ltd., 3rd edition, 453 pp., 294 illustrations in text. 26x17 cm.
(Ticks, pp. 360-374, Figs. 236-239, none original and all poor. Numerous
mistakes.)

Brown, A. A. (1902), " Fowl Tick {Argas miniatus)." Journ. Dep. Agric. Victoria,
i. 86-90 and 209-212. (Inaccessible ; cited by Lounsbury, 1903, p. 6.)

Brumpt, E. (1901), Notes et observations sur les maladies parasitaires. Arch, de
Parasitol., iv. 561-580. (p. 578 re effects of bites of 0. moubata.)

Caiman, W. T. (15. IV. 1905), Ornithodoros savignyi var. caecum [should be caerus].

Journ. Tropical Medicine, vni. (8) 124. (0. moubata.)
Canestrini, G. (1890), Prospetto dell' Acarofauna italiana ; Fainiglie: Tetranychini,

Ixodini, Argasini, iv. 427-540, Tav. xxxvii.-xliii. Padova.
(1892a), Prospetto dell' Acarofauna italiana; Intorno alia metamorfosi degli

Ixodini (Nota). Abbozzo del sistema acarologico ; Famiglia dei Phytoptini

(Phytoptidae), v. 543-722, Tav. xliv.-lix. Padova.
Canestrini, G., and Fanzago, F. (1877), Intorno agli acari Italiani. Atti del

reale Istituto Veneto di scienze, lettere ed arti, ser. 5, IV. (1877-78) (1 [for

Nov., 1877]), 69-208, Tav. ii.-vii. (Reprint paged 1-140, citations made from

the latter.)
Chatelin (1882), [Effects of bite of A. rejlexus on man]. Soc. Entomol. de France,

Seance du 24. v. 1882. (Cited by Laboulbene and Megnin, 1882, p. 340, and

Railliet, 1895, p. 717.)
Christophers, S. R. (1906), The Anatomy and Histology of Ticks. Scientific

Memoirs by Officers of the Med. and Sanit. Dep'ts of the Gov't of India, Calcutta,

N.8., No. 23, Pis. I-VI, 55 pp.
Christy, C. (1903), "Tick Fever" in Man. (Including a note by R. I. Poeock.)

Brit. Med. Journ., II. 653 ; also appeared in Thompson and Johnston Laboratories

Report, v. 187-189, PI. XV.

Bibliography 7

Churchill, S. (1880 a), Note on Argas persicus. Proc. Entom.Soc. Lond.,\>\>. x\- xxiii.
Claus, C. (1880), Grwidztige der Zoologie, i. 821, 8°, Marburg.

(1887), Lehrbuch der Zoologic, 4te Aufl. Marburg and Leipzig : N. G. Elwert.

[pp. 470-474 give a brief description of chief features of Ixodes and reproduce
three figures from Pagenstecher (1860, 1861) : Ixodes ricinug, general internal
anatomy (Fig. 443) ; Argas, <j> and <J sexual organs (Fig. 444) ; Ixodes, rostrum
(Fig. 450).]

Cobbold, T. S. (1879), Parasites; a treatise on the Entozoa of man and animals,
including some account of the Ectozoa. Pp. xi + 508, figs. 85, London.

(1882), Human parasites. A manual of reference to all the known species of

Entozoa and Ectozoa which are found infesting man. 88 pp. London : Longmans,
Green and Co. (Brief mention of ticks on pp. 77, 78.)

Cook, A. R. (1904), Relapsing fever in Uganda. Journ. Trap. Med., 15 Jan., p. 24.

(Epidemics of the disease in Budu and Mengo (Uganda), in 1903, found spiro-

chaetes in the blood of patients : he observed the disease in 1899, at Mengo.)
Cuvier, Q. (1833), Arachnidae, with additions by Griffiths and Pigeon. (Cited by

George, 1876, p. 224, as referring to Argas persicus and A. pipistrellae (last

figured). See Duges and Edwards (no date).)

Dolly, C. (1894), Synonymatic table of the animal parasites of man and of the
animals commonly associated with him. An illustrated dictionary of medicine,
biology, and allied sciences, etc., by George M. Gould. Philadelphia, pp. 978-
1021, Figs. (Cited by Salmon and Stiles, 1901.)

Donitz, W. (1906), Ueber Afrikanische Zecken. Sitzungsber. d. Gesellsch. Naturfor-
schender Freunde, Jahrg. 1906, No. 5, pp. 143-148, 8 Figs., reprint.

(1907), Die wirtschaftlich wichtigen Zecken mit besonderer Berucksichtignng

Afrikas. 127 pp., 38 Figs, and 6 Plates. Leipzig: J. A. Barth.

(IX. 1907), Insekten als Verbreiter von Krankheiten : Die Zecken. Ber. ii. d.

XIV. Internal. Kongr.f. Hygieneu. Demogr., Berlin 23-29 Sept., 1907, a. 179-188.

Dowson, J. (1. VI. 1895), Poisonous tick. Brit. Med. Journ., I. 1201. (Followed

by a short note by B. I. Pocock.)
Dubreuilh, W. (1900), " Dermatozoaires" in Pratique dermatologique. T. I., 8°, Paris :

M asson.
Dubreuilh, W., and Beille, L. (1896), Les parasites de la peau humaine. 185 pp.,

22 text-figs. Paris : Masson et Cie. (Date taken from Huber, 1899, p. 15 ;

original undated ; ticks referred to pp. 68-84, Figs. 4-8.)
Du Buysson (1896), Observations sur les accidents produits par la piqure des Acariens

du genre Argas. Ann. Soc. Entomol., 10 pp. (Cited by Huber, 1899, p. 15.)
Dug^S, A. (1876), llepertorio de Guanajuato, Mexico. (Cited by Megnin, 1885,

pp. 463, 466, etc., who gives translated extracts and reports on subsequent

personal communications received from Duges and Aleman. Also cited by

Neumann, 1892, Malad. Parasit., p. 101.)

(1883), Natnraleza de Mexico, v. 195. (Cited by Megnin, 1885, pp. 465,

472, who gives a figure of A. megnini Duges. Cited by Neumann, 1896, p. 16.)

(1884), Turicata y garrapata de Guanajuato. La Natnraleza, Periodico

cientifico de hi Sociedad Mexicana de Historia natural, vi. (1882-1884), Mexico,
pp. 195-198, PI. IV. (MS. dated Nov., 1882.)

Bibliography 9

Duges, A. (1891), La Naturaleza (2), I. 20, 1 PI. (Cited by Neumann, 1896.)
Duges, A. L. (1834a), Recherches sur l'ordre des Acariens en general et la famille

des Trombidies en particulier ; Premiere memoire. Ann. de sci. not., Paris, ser.

2, i. Zool, pp. 5-46, PI. I. (Reprint paged 1-42.)

(1834 c), Recherches sur l'ordre des Acariens ; Troisieme memoire. Romarques

sur la famille des Gamases. Ann. de sci,. not., Paris, ser. 2, II. Zool., pp. 18-63,
Kb. VII, VIII. (Reprint paged 1-44.)

(1836), Recherches sur l'ordre des Acariens en general (i) et la famille des

Trombidies en particulier (n). Remarques sur la famille des Hydracnes (m).
Ann. des Sc. nat., 2e ser., Zool. T. I., reprint pp. 42, PI. I (i. Memoire) ; ibid.
pp. 30, Pis. X, XI (n. Mem.) ; ibid., vol. n. pp. 46, Pis. VII and VIII.

Duges, A. L., and Milne Edwards (no date), " Arachnides " in La Regne Animal, by
G. Cuvier. Paris: Masson. (Figs, illustr. Ixodoidea on PI. XXVII.)

Dupr6 (1819), Voyage en Perse fail dans les anne'es 1807, 1808 et 1809, n. 324. Paris.
(Lengthy quotations of essential parts in Fischer de Waldheim (1823), Laboulbene
(1867), and Laboulbene and Megnin (1882) : A. persicus.)

Dutton, J. E., and Todd, J. L. (XI. 1905), The Nature of Human Tick-Fever in the
Eastern part of the Congo Free State (dated 28. in. 1905). Liverpool School of
Tropical Medicine — Memoir xvn., 18 pp., 9 Charts ; see also Brit. Med. Journ.,
II. 1259, 1260.

Efisio, M. (1892), Saggio d' un catalogo metodico dei principali e piu comuni
animali invertebrati della Sardegna. Bollettino delta Societd Romana per gli
studi Zoologica, I. (6), 246-282. (Cited by Salmon and Stiles, 1901.)

Ellison, A. (1903a), Birds killed by ticks. Zoologist, Lond. (743), 4. s. (77), V. 7,
May 15, p. 194. (Refers to midges, no word of ticks!)

Ercolani, G. B. (1859), Nuovi elementi teorico-pratici di medicina veterinaria; Dei
parassiti dei morbi parassitarii degli animali domestici, pp. 550, Bologna. (Cited
in bibl. by Salmon and Stiles, 1901.)

Ernouf, Baron (1880), Le Caucase, la Perse, etc., cPapres les relations de M. de
Thielmann, p. 256. Paris, 1880. (A. persicus, cit. Huber, 1899, p. 16.)

Fabricius, J. C. (1794), Entomologia systematica, emendata et aucta, secundum classes,

ordines, genera, species adjectis synonymis, locis, observationibus, descriptionibus,

iv. pp. 472 + index, pp. 5, Hafniae. (Ticks: see pp. 425-434.)
(1805), Systema antliatorum secundum ordines, genera, species adjectis synonymis,

locis, obsereationibus, descriptioiiibas, pp. xiv + 15-372 + 30, 12°, Brunsvigae.
Fanzago. See Canestrini and Fanzago.
Feldmann (1905a), Ueber Filaria perstans im Bezirk Bukoba. n. Theil mit

1 Tafel. Arch, f Schiffs- u. Trope n- Hygiene, IX. (2), 62-65.
(1905b), On Filaria perstans in the Bukota District. (Transl. by T. S. Kerr of

the preceding : Feldmann, 1905a.) Journ. Tropical Medicine, vin. (8), 125-

126. (Should be " Feldmann " and " Bukoba.")
Fischer de Waldheim, G. (1823), Notice sur l'Argas de Perse (Malleh de Mianeh)

docrit par les voyageurs sous le noui de punaise venimeuse de Miaua. Mem. Soc.

imp. de nat. de Moscou, vi. 269-283, PI. XXIII, Figs. 1-11. (Figs. 1, 2 Ix.

camelinus, 3-7 Ix. ricinus, 8-11 Argas persicus, Fischer n.sp. Gives an account

of known species of ticks, quoting other authors.)

Bibliography 11

Fischer de Waldheim, G. (1824), Du Mallah do Minna, insecte de Perse, dit punaise

vcnimeuse. (Abstract of Fischer von Waldheim, 1823, by J. J. Virey.) Journ.

de pharm., Paris, V. 10 (5), Mai, pp. 242-245. (Note by Latreille, p. 245.)
Fischer, H. (1849). See Gene, 1849.
Francis, M. (1894), Veterinary Science. Bull. No. 30, Texas Agric. Exp. Sla.,

March, pp. 436-458, Pis. I-III. (Cited by Salmon and Stiles, 1901.)
Fritsch, G. (1875), Ueber die sogen. " Giftwanze von Mianah." Sitzungsber. d.

Geselhck. naturf. Freunde zu Berlin, pp. 61-64. (A. persicus, cit. Huber, 1899,

p. 16.)
Froggatt, W. W. (1901), The fowl tick (Argas americanus Packard). Agric. Oaz.,

JV.S. Wales, XII. 1349-1352, PI. (inaccessible).

(1906), Notes on fowl-tick and poultry. Agric. Oaz., JV.S. Wales, xvn. 14-18.

Fiilleborn and Mayer, M. (1908), Ueber die Moglichkeit der Uebertragung patho-

gener Spirochaeten durch verschiedene Zeckenarten. Arch. f. Schiffs- u. Trop.-

Hyg., xii. 31.
Fullager, J. (1874a), A Canterbury Arachnid, new to the British Fauna. Science

Gossip, 1874, pp. 121, 122, Fig. 86.
■ (1874b), The Canterbury Tick (Argas reflexus). Science Gossip, 1874, p. 234,

Fig. 164.
Fuller, C. (1897), A Fowl-infesting Tick (Argus sp.). Agric. Gaz., N.S. Wales, vn.

590.
Fumouze (1873), Sur un Acarien de lafamille des Ixodes, Argas persicus. Ann. Soc.

Entomol. (Cit. Huber, 1899, p. 16.)
Fiirstenberg, M. H. (1861), Die Kriitzmilben der Menschen und Thiere, pp. i-v + 1-240,

Taf. I-XV, 4°, Leipzig. (Cited in bibl. by Salmon and Stiles, 1901.)

Galli-Valerio, B. (IX. 1907), Les insectes comme propagateurs des maladies : Le

role des arthropodes dans la dissemination des maladies. Ber. ii. d. XIV. Internal.

Kongr. f. Hygiene u. Demogr., Berlin, 23-29 Sept., 1907, II. 189-194. (A short

review of progress to date, refers briefly to ticks.)
Gamgee, J. (1871), Our domestic animals in health and disease, II, division 3,

pp. xvi + 613, Figs. 149-237 (238). (Cited by Salmon and Stiles, 1901.)
de Gay (1849 51). See Gervais (same date).
Gene\ G. (1844), Memoria per servire alia storia naturale degli issodi. Mem. d.

li. Accad. Torino, 2d ser. ix. (1848), 751-786.

(1849), Beitrage zur Naturgeschichte der Zecken von Professor J. Gene1 in Turin.

Aus dem Italienischen, mit Anmerkungen von Dr H. Fischer, Privatdocent zu
Freiburg, pp. 34, 4°. Freiburg im Breisgau : F. X. Wangler.

George, C. F. (1876), On "The Blyborough Tick" (Argas Fischerii). Journ. Quekett
microsc. Club, iv. 223-225, Pis. XXI and XXII ; also Sc. Gossip, 1877, O.S. xm.
104.

Gerstaecker, C. E. A. (1860), Argas reflexus Latr., ein neuer Parasit des Menschen.
Vircliovi's Arch., xix. 457-470, PI. XV, Figs. 1-6.

(1863), "Arthropoden" in Handbuch der Zoologie, by Peters, Cams and

Gerstaecker. Leipzig, n. 1-421, 8°. (See Acarina, pp. 343, 344.)

(1873), GliederthieroOstafrika/svon C.v.d. Decken's Rcise, pp. 464-470. (Ambl.

'Iiurneum, Dermacentor pii/chellus, 0. savignyi.)

Bibliography 13

Gervais, P. (1844), Aceres Phryneides, Scorpionides, Solpugidcs, Phalangides et
Acarides ; Dioeres Epizoiques, Aphanipteres et Thysanoures, in vol. m. of
Walckenaer's Histoire naturelle des Insectes. Apteres, pp. i-viii, 1-476, Paris.

(1847), in vol. iv. of Walckenaer's Histoire naturelle des Insectes. Apteres,

pp. i-xvi, 1-623, Paris.

(1849-51), in de (lay's Historia fisica y politico, de Chile. Zoologie, iv. 44,

PI. V, Fig. 8.

Gervais, P., and van Beneden, P. J. (1859), Zoologie me'dicale ; expose me'thodique

du regne animal. 2 vols., 8°, Paris.
Gibert, J. M. (1896), L'Argas reflexus et son parasitisme chez l'homme. These :

Bordeaux, 60 pp., 8°, illustr. (Cit. by Mosler and Peiper (1904, p. 373) who state

author observed general symptoms after bite of A. reflexus in man.)
Gueiin-Meneville (1829-1843), Iconographie du regne animal, de G. Cuvier.

Arachnides, PI. VI, Figs. 1-6, pp. 16, 17.

(1849), Description de PArgas talaje. Rev. et Mag. de Zool., pp. 342, 343,

PI. II.

Guiart, J., and Grimbert, L. (1906), Diagnostic Chimique, Microscopique etParasito-
logique. Paris: F. R. de Rudeval, Editeur, 500 Figs., 960 pp. {Ixodidae: pp.656,
666, Figs. 401-410, of which 422 and 410 of Hyal. aegyptium and 0. moubata
are original but very sketchy. The text is compiled from various authors.)

Gulliver, G. (1872), On a Canterbury Arachnid new to the British Fauna. Quart.
Journ. Micr. Sci,, xn. N.S., 205, 206.

Gulliver and Gulliver (ca. 1871), Reports of East Kent Nat. Hist. Soc. Abstracts
of paper in Quart. Joum. Micr. Sci., April-Oct., 1872. (Cited by Fullager, 1874a,
p. 121.)

Haller, G. (1882), Beitrag zur Kenntniss der Milbenfauna Wiirttembergs. Jahreshefte

des Vereins fur vaterlcindische Naturkunde in Wiirttemberg, xxxvm., Stuttgart,

pp. 293-325, Taf. V. (See especially pp. 309-312, re Ixodidae, and Figs. 4

and 5.)
Hart, J. H. (1899), The tick of the domestic fowl and fowl fever. Bulletin of

Miscellaneous Information, Trinidad, III. Bull. 19, Pt. 11, Apr., p. 180.
Hassall, A. (1900), Note on the chicken tick (Argas americanus). \§th Annual

Report of the Bureau of Animal Industry, 1899, U.S. Dep't of Agric, pp. 496-

500, Figs. 16-22, PI. XVI. (Reprint dated 1901.)
Heller, A. (1880), Die Schmarotzer mit besonderer Beriicksichtigung der fiir den

Menschen wichtigen, 74 woodcuts, 1 Map, 230 pp. Munchen and Leipzig : R.

Oldenbourg. (P. 154, less than a page devoted to " The tick," contains nothing

original or worth noting.)
Heller, C. (1858), Zur Anatomie von Argas persicns. Sitzungsberichte d. Kaiserl.

Akad. Wien, xxx., Math.-naturwiss. Klasse, pp. 297-326, Pis. I-IV.
Hermann, J. F. (1804), Me'moire apterologique, An xn, pp. 63-71, Pis. IV, VI, VII,

Strasbourg.
Heyden, C. von (1826), Versuch einer systeinatischen Eintheilung der Acariden.

Isis von Okeu, Jena, i. Heft 6, pp. 608-613. (His system is reprinted in

Michael, 1883, pp. 32-34.)

Bibliography 15

Hinde, S. L. (1897), La Chute de la domination des Arabes die Congo. Th. Falck

and Uie, Bruxelles. Chap, xn., p. 3. (Refers to fever about Kasongo attributed

by natives to tick-bite.)
Hoehr, F. (VII. 1893), Further concerning the new chicken plague in Texas. Insect

Life, v. (5), 348.
Hooker, W. A. (1908a), Life history, habits and methods of study of the Ixodoidea.

Joum. Econ. EntomoL, I. 34-51. (Good original observations.)

(1908b), A review of the present knowledge of the role of ticks in the

transmission of disease. Joum. Econ. EntomoL, I. 65-73. (A useful digest
of some recent work.)

Howard, L. 0. (1895), Note on the chicken tick. Insect Life, vn. (5), July,
pp. 417, 418.

See Riley and Howard.

Huber, J. Ch. (1899), Bibliographie der Klinischen Entomologie, 1899 (Argas, Ixodes),

Heft 2. Jena: Fromaun.
Hubbard, H. G. (1894), The insect guests of the Florida land tortoise, Qopherus

(Xcrobates) polyphemm. U.S. Dep't Agric, vi. 306. {Ornithodoros americanus

Marx (Ms.) in the burrows and on tortoise — Amblyomma tuberculatum Marx

n.sp. on the tortoise.)
Hunter, W. D., and Hooker, W. A. (1907), Information concerning the North

American Fever Tick, with notes on other species. U.S. Dep't of Agriculture,

Bur. of Entomology, Bull. No. 72. Washington : 12 Figs., 4 Plates, 87 pp.
Husemann, T. (1862), Handbuch der Toxicologie, p. 248. (A. persicus, cit. Huber,

1899, p. 16.)

Johannessen, A. (1885b), Acute Polyurie bei einem Kinde nach dem Stiche eines
Ixodes ricinus. Archivf. Kinderheilk., VI. 337-350.

Karsch, F. (29. IV. 1878), Uebersicht der iti Mossambique gesammelten Arachniden.
Monatsbericht der K. Akademie der Wissenschaft zu Berlin, pp. 314-338, Pis. I,
II. (Ixodoidea, pp. 336-338, PI. I, Fig. 4, of Amblyomma Petersii Karsch, nat.
size, uncoloured.)

(1878), Ueber einige von Herm J. M. Hildebrandt im Zanzibargebiete erbeutete

Arachniden. Zeitschr. f. d. ges. Naturwiss., Berlin, ser. 3, III. (LI. ganzen Reihe),
pp. 311-322, Taf. VIII.

(1880), Vier neue Ixodiden des Berliner Museums. Mittheil. d. Miinc/tener

entomol. Vereins, Jahrg. iv. 141, 142.

Kerr, T. S. (15. IV. 1905), Remarks on the paper by Feldman (1905) entitled "On
Filaria perstans in the Bukota District." Joum. of Tropical Med., VIII. (8),
126.

Koch, C. L. (1844), Systematische Uebersicht iiber die Ordnung der Zecken. Arch,
f. Naturg., Berlin, Jahrg. x. Bd. I., 217-239. {N.sp. and genera from different
parts of world. Mode of copulation. Argasidae, Ixodidae, Rhipistomidae.
10 genera, 128 pp. (discussed by Marx Uf ■».)), p. 234 llaemnphysalis.)

(1847), Uebersicht des Arachnidensy stems. Niirnberg. Heft 4, pp. 136, Taf.

I-XXX.

Bibliography 1 7

Koch, L. (1875), Aegyptische und Abyssinische Arachniden gesammelt von Herrn
C. Jickeli beschrieben und abgebildet. 96 pp., 7 Pis., 4°. Niirnberg : Bauer
and Rasi>e. (0. savignyi, p. 1.)

Koch, R. (23. XI. 1905), Vorliiufige Mittheilungen iiber die Ergebnisse einer
ForschuDgsreise nach Ostafrika. Deutsche med. Wochenschr., p. 1866.

(1906), Ueber afrikanischen Recurrens. Berlin, klin. Wochenschr., xliii. 185,

10 Figs.

Kolenati, F. A. (1857), Die Parasiten der Chiroptern, pp. 51, iv. Taf., 8°, Dresden.

(1858), Synopsis prodroma der auf Chiroptern als Epizoen vorkommenden

Lausmilben, Carida Kolenati. Wien. entnmol. Monatsschr., Wien, II. pp. 4-7.

Kollar( ), Bildlichen Nalurgeschichte des Thierreiches, II. 274, Fig. 7. (Cited by

Heller, 1858, p. 297.)
Kotzebue, M. (1819), Voyage en Perse a la suite de Vambassade Russe en 1817.

Paris, viii. 180. (For lengthy quotations of essential parts see Fischer de

Waldheim (1823), also Laboulbene (1867), and Laboulbene and Mt'gnin (1882).)
Krapf (1882), Suaheli Dictionary ; Trubner and Co., London. (Cited by Dutton

and Todd, 1905, pp. 3, 18 as stating : "Pasi, PL Papasi, ticks, an insect said to

cause fever in one who is bitten by it.")
Kiichenmeister and Ziirn (1879), Parasiten, p. 538, et seq. {Argas, cit. Huber,

1899, p. 14.)
Kiichenmeister, F. (1855), Die in und an dem Korper des lebenden Menschen vorlcom-

menden Parasiten, I. Abt. : Die thierischen Parasiten, 486 pp., 9 Plates.

Leipzig: B. G. Teubner. (See pp. 421-424, re Ticks, one poor text-fig. of

" Ixodes ricinus," after Qurth, the same figure is also found in Ziirn (1872).)

Laboulbene, A. (1867), article "Argas" in Dechambre's Dictionnaire encyclop. des sci.

med., VI. 53, 54. (Quoted verbatim by Laboulbene and Mi'gnin, 1882, pp. 325-

327.)
Laboulbene, A., and M^gnin, P. (1882), Memoire sur les Argas de Perse. Jonm.

de I'anat. et de la physio/. Paris, XVIII. 317-341, Pis. XXI-XXIII.
Latreille, P. A. (1796), Precis des caracteres ge'neriques des insectes, disposes dans un

ordre naturel. An v. pp. xii+201, 12°, Brive. (Carios : p. 177.)

(1804), Histoire naturelle, ge'nerale et particuliere, des crustace's et des insectes.

An xii., viii. pp. 411, 8°, Paris. (Caris: p. 55.)

(1806), Genera crustaceorum et insectorum, I. pp. xviii + 302, Tab. i-xvi. (Caris :

p. 161.)

(1829), Crustaces arachnides et partie des insectes. Cuvier's Le regne animal,

Paris, iv. pp. 584, 8°. (See Ixodes, pp. 287-290.)

(1835), Ueber Caris vespertilionis. Isis von Oken, Jena (6), pp. 511, 512.

Leach, W. E. (1815), A tabular view of the external characters of four classes of

animals, which Linne arranged under Insecta ; with the distribution of the
genera composing three of these classes into orders, etc., and descriptions
of several new genera and species. (Read April — June, 1814.) Trans, of
the Lmnaean Society, London, xi. 306-400. (Ixodoidea : pp. 396-399.) '
Linstow, 0. von (1894), Die Giftthiere und Hire Wirkung auf den Menschen, 148 pp.,
54 Figs. Berlin : Aug. Hirschwald. (Brief reference to Ticks (Argas), pp. 120-
121.)
n. i. 9*

Bibliography 19

Livingstone, D. (1857), Missionary Travels and Researches in South Africa, etc.,

711 pp. (Ticks : pp. 382, 383, 628, 629.) London : John Murray.
(1874), The last Journals of David Livingstone, in Central Africa, from 1865 to

his death, continued by a narrative of his last moments, etc., by H. Waller.

2 vols., 360 + 346 pp. (Ticks: vol. II. pp. 33, 115.) London: J. Murray.
Lounsbury, C. P. (1895), Fowl ticks (letter from T. W. Parker answered by C. P. L).

Agric. Journ., Cape Town, vin. (25), pp. 657, 658.

(1899a), Pig lice, alias Tampans. Agric. Journ., Cape Town, xiv. (4), Feb. 16,

p. 240. (Tampan = Ornithodoros.)

(1899b), Ticks. Report of the Oov't Entomologist for the year 1898. Cape of

Good Hope Dep't of Agric, Cape Town, 1899, pp. 43-46, PI. IX.

(1900a), Notes on some South African Ticks. Proc. VZth Ann. Meeting of the

Assoc, of Econ. Entomologists. Issued by U.S. Dep't of Agric, Division of
Entomology. Bulletin No. 26, n.s. Washington, pp. 41-48. (Amblyomma
liebraeum, H. aegyptium, Rh. decoloratus, Rh. evertsi, Orn. savignyi, Arg.
persieus.)

(1900b), Insect bites and the effects thereof. Canadian Entomologist, xxxn.

17-24. (Bites of ticks, not insects. Argas, Orn. savignyi or tampan.)

(1902), (Tick investigations) Tampan not associated with heartwater (p. 65) —

Tick- Heart water investigations (pp. 2-4, 29-74) — Tick Malignant Jaundice
investigations (pp. 4-9) — Tick, catching and destruction (pp. 3, 4, 49, 71),
Pis. IV-VI. Report of the Gov't Entomologist for the year 1901, Cape of Good
Hope Dep't of Agric. Cape Town, 1902. (Tampan = Ornithodoros.)

(IX. 1903), The fowl tick. Studies on its life-cycle and habits. Agric. Journ.,

Cape Town, 15 pp., 3 Pis., repr.

(XI. 1904), External parasites of fowls. Agric. Journ., Cape of Good Hope.

No. 27, Repr., 5 pp., 4 Figs. (Includes a short account of Argas persieus ; more
fully treated in Lounsbury (IX. 1903).)

Lucas, H. (1849), Exploration scientifique de I'Alge'rie. Zoologie I. Histoire

naturellc des Animaux artieules, Part I, pp. 316, 317.
Ludwig, H. (1886), Leunis' Synopsis der Thierkunde, 3. Aufl. II. pp. xvi + 1231,

Figs. 1160, Hannover. (Cit. in bibl. by Salmon and Stiles, 1901.)

Manson, P. (1903), Tropical Diseases. A Manual of the Diseases of Warm Climates.

3rd ed. Cassell and Co., London, 756 pp. {Re Ornithodoros moubata see pp. 713,

714.)
(1908), Tropical Diseases. A Manual of Diseases of Warm Climates.

London: Cassell and Co., Ltd., 876 pp. (Ticks described, pp. 202-205 and

figured on coloured PI. IV : Boophilus (? annulatus) $> , Ixodes ricinus <j? [both

gorged], Orn. moubata, A. persieus, all in dorsal aspect.)
Marchoux, E., and Salimbeni, A. (IX. 1903), La Spirillose des Poules. Ann. d.

PInst. Pasteur, xvn. 569-580, repr.
Marlatt, C. L. (11. VIII. 1899), A Californian Tick. Canadian Entomologist, p. 229,

repr. (Ai-gasl found in Santa Lucia Mtfl.)
Marx, G. (1892a), Notes on the Classification of the Ixodidae. Proc. Entomol. Soc.

Washington, n. 232-236. (Cynorhoestea proposed as a Sub-order of Acarina to

9*-2

Bibliography 21

include all ticks (Argasidae, Ixodidae, Eschatocephalidae, Haemalastoridae,
lihipistomidae) under 2 groups : Catastomata, Antistomata.)
Marx, G. (1892b), On the morphology of the ticks. J'roc. Entomol. Soo. Washington,
n. 271-287. (Historical sketch re Antistomata; larvae, adults, detailed
morphology.)

(1895), Obituary Notice of his Work. Plate illustrating following species

published in connection with obituary, 1'roc. Ent. Soc. Wash., vol. in. pp. 195-
201 : Rhynchoprion spinosum ( = Ornithodoros megnini, nymph), Oruithodoros
americanus ( = 0. megnini, adult), Argas americanus.

Massey, A. Y. (1905), Spirillosis in Portuguese West Africa. Journ. Trop. Med.,

vin. 225. (Describes a case in man.)
M^gnin, P. (I. 1868), La fainille des Ixodes. Journ. de me'd. ve'tfr. inilit., vi. 423.
(1876a), Memoire sur l'organisation et la distribution zoologique des acariens

de la Fainille des Gamasides. Journ. de Vanat. et de la Physiol., Paris, XII.

Mai-Juin, pp. 288-336, Pis. VII, VIII.

(1880a), Les Parasites et les Maladies Parasitaires chez I'homme, les animaux

domestiques et les animaux sauvages avec lesquels Us peuvent Hre en contact; in-
secies, arachnides, crust/ice's, pp. 478, Figs. 65, Pis. XXVI, 8°, Paris. (Ticks :
pp. 106-136, 322, 344, 363, 403, 422.)

(1882), Experience sur Taction nocive des Argas de Perse. Gompt. rend. Soc. de

biol., Paris, Se>. 7, iv. 305-307.

(1885), Les Argas du Mexique. Journ. de Vanat. et de la physiol., Paris, xxi.

460-474, Pis. XX, XXI.

(1892), Les Acariens Parasites. (Encyclopedia Scientifique des Aide-memoire.)

Paris: Gauthier-Villars et Fils. 40 Figs., 182 pp., 12°. (See pp. 25-67:
Chapter n., " Famille des Ixodides.")

(1895 a), Les parasites articule's chez I'homme et les animaux utiles (maladies

qu'ils occasionnent). 2. ed. des parasites et maladies parasitaires, augmented d'un
app'endice sur les parasites des cadavres. 2 vols., 1 pi., 510 pp. ; atlas, 1 pi.,
26 1., 26 pp., 8°. Paris : G. Masson.

(15. X. 1895), Sur les prctendus r61es pathogbniques des Tiques ou Ixodes.

Bull, de Vacad. de me'd., xxxiv. 354-369.

M^gnin, P-, and Laboulbene (1882), Sur les Argas de Perse. Compt. Rendus Soc.

Biol., Paris, Ser. 7, iv. p. 577. (See full paper under Laboulbene and

Mcgnin.)
Mingazzini (1898), Zooloyia medica, pp. 245-249. (A. reflexus, cit. Huber, 1899,

p. 16.)
Mollers, B. (1907), Experimentelle Studien liber die Uebertragung des Riickfallfiebers

durch Zecken. Ztschr. f. Hyg. u. Infektionskr., lviii. 277-286.
Moniez, R. (1896), Traite de Parasitologic, pp. 499-509. (Refers especially to

A. reflexus, mentions other Argasidae; cited by Huber, 1899, Heft 2; 1900,

Heft 4.)
Moquin-Tandon (1860), Elements de Zoologie medicate. (Cited by older authors re

ticks and in Huber's bibliography (1900, Heft 4).)

(1861), Elements of Medical Zoology, translated and edited by Robert Thomas

Hulme, pp. 423, Figs. 124, London. (Cited by Salmon and Stiles, 1901.)

Bibliography 23

Morgan, H. A. (1899a), Ticks and Texas Fever. Bull. No. 56, ser. 2, Louisiana

Agric. Exp. Sta., pp. 128-141, Pis. I-IX.
Mosler, F., and Peiper, E. (1894), Thierische Parasiten, 1st ed., in NotlinagePa

Speciellc Pathol, u. Therapie, vi. 345 pp., 124 Figs., 8°. Wien : Alfred Holder.

(Pp. 320-322 brief mention of ticks, nothing original, including two figures.

See note to 2nd ed.)

(1904), Tieriwhe Parasiten, 2nd ed., 376 pp., 162 Figs. Wien : Alfred Holder.

(Brief reference to Ticks on pp. 344-346 ; one Fig. of Argas reflexus after
Pagenstecher ; contains nothing original except that a Fig. (No. 140) of Ixodes
ricinus after Kiichenmeister is described in the legend as Dermanyssus gallinae !
This correct in the 1st ed., p. 320.)

Murray, A. (1877), Economic Entomology ; Aptera, I. pp. iii.-xxiii., 3-433, with
numerous text-figures. London : Chapman and Hall (South Kensington
Museum Science Handbooks). {Sec Ixodoidea, pp. 180-204, 28 Figs., of which
9 appear original (?) ; all poor however and mostly copied from Koch, C. L., 1847.)

Neumann, L. G. (1888), Traitedes maladies parasitaires non-microbiennes des animaux
domestiques, pp. 673, Figs. 306. Paris.

(1892a), Traite" des maladies parasitaires non-microbiennes des animaux domes-
tiques, 2. ed., pp. 767, Figs. 364. Paris.

(1892b), A treatise on the parasites and parasitic diseases of the domesticated

animals, pp. 800, Figs. 364. Paris.

(1896), Revision de la famille des Ixodides. I. Argasines. Mem. Soc. Zool.

de France, IX. pp. 1-44, Figs. 1-36.

(1901), Revision de la famille des Ixodides. 4e Metnoire. Mem. Soc. Zool. de

France, xiv. 249-372, 18 Figs.

(1905), Notes sur les Ixodidds, III. Arch, de ParasitoL, ix. 225-241.

(1906), Notes sur les Ixodides, IV. Arch, de Parasitologic, x. 195-219, 17 Figs.

(1907), Notes sur les Ixodides, V. Arch, de Parasitologic, xi., pp. 215-232,

Figs. 1-14.

(V. 1907), Note XI, Quatre nouvelles especes d'lxodides. Notes from the

Leyden Museum, xxix. 88-100, 7 Figs.

(XII. 1907), Note sur les Ixodidae recueillis dans les lies de l'ocean Indien par

M. J. Stanley Gardiner. Trans. Linn. Soc, London, 2 ser., Zoology, xn. 193-
194.

(III. 1908), Notes sur les Ixodides, VI. Arch. d. Parasitologic, xn. 1-27,

16 Figs.

Neveu-Lemaire, M. (1904), Parasitologic animate, 2nd ed. {Precis de Medecine,

No. 1), 218 pp., 301 Text- Figs. Paris : F. R. de Rudeval. (Ixodoidea : 151-157,

4 Figs. ; none original.)
Newstead, R. (XI. 1905), On the External Anatomy of Omithodoros moubata.

Liverpool School of Tropical Medicine, Memoir xvn. pp. 21-26, 1 Map,

Pis. I-IV.

(30. XII. 1905), On the pathogenic Ticks concerned in the distribution of

disease in man, with special reference to- the differential characters in Ontitho-
doros moubata. Brit. Med. Journ., pp. 1695-1697 ; also Lancet, clxix. pp. 365-
385.

Bibliography 25

Nuttall, G. H. F. (1899), On the role of insects, arachnids and myriapods, as carriers
in the spread of bacterial and parasitic diseases of man and animals. A critical
and historical study. Johns Hopkins Hospital Reports, vm. 154 pp., 3 Plates.
Published also in part in Hygienische Rundschau, 1899, ix. See especially
pp. 402-408.

(1905), Ticks and tick-transmitted diseases. (Paper read 9. xu. 1904.) Trans.

Epidemiol. Soc, London, N.S., vol. xxiv., Session 1904-1905, pp. 12-32, 1 Map.

(VII. 1908), The Ixodoidea or Ticks. Journ. Roy. Inst. Publ. Health, xvi.

385-403, 7 Figs., 1 Diagr. (original). Harben Lecture I.

(VIII. 1908), Spirochaetosis in Man and Animals. Journ. Roy. Inst. Publ.

Health, xvi. 449-464. Harben Lecture II. (Argasidae in relation to disease.)

Oken (1818), Sogenannte giftige Wanze in Persien. Isis, 1818, pp. 1567-1570,

PI. XIX, Figs. 1-4.
Oshorn, H. (18%), Insects affecting domestic animals : an account of the species of

importance in North America, with mention of related forms occurring in other

animals. U.S. Dep't of Agricidt., Division of Entomol., Bulletin No. 5, N.S.

(Washington D.C.), 302 pp., richly illustrated.

Packard, A. S. (1872), " Arachnida" in U.S. Geological Survey of the territories, etc.,
p. 740, Fig. 68. (Argas americanus : description reprinted by Osborn, 1896,
p. 256.)

(1893), Note on A. americanus. Insect Life, v. 267, 348.

(1895), Note on the chicken tick {Argas americanus, Pack.). Insect Life, vil.

417, 418.

Pagenstecher, H. A. (1861), A. reflexus. Virchoufs Arch., xix. p. 457. (Cited by
Pagenstecher, 1862.)

(1862), Zur Anatomie von Argas reflexus. Zeitschr. f. loiss. Zoologie, xi. 142-

155, PI. XVI.

Pavesi, P. (1884), Materiali per lo studio della fauna tunisina. II. Aracnidi. Annali
del Museo C'ivico di Storia Naturale di Oenova, xx. pp. 483-485.

Perroncito, E. (1882), / Parassiti dell' uomo e degli animali utili, p. 459. (See next
title.)

(1901), / Parassiti dell' uomo e degli animali utili e le piu comuni malattie da

essi prodotte. Profilassi e cura relativa, pp. 276, Text-Figs., 25 Plates. Milano :
F. Vallardi. (Pp. 561-568 relate to ticks, 2 Figs., one after Gurlt, the other very
similar to Blanchard's of A. persicus.)

Planchon, L. (1895), Observations sur la resistance vitale de V Argas reflexus, Latr.

Feuille des Jeun. Naturalist., (3) xxvi. 29. (Cit. Huber, 1899, p. 15 ;

inaccessible.)
Plehn, A. (1902), Tropenhygiene. (Cited by Werner, 1906, p. 779, as stating that

the natives of German E. Africa have long referred fever to bites of " papasi "

(Suaheli for tick).)
Pocock, E. I. (23. I. 1900), On a collection of insects and arachnids made in 1895

and 1897, by Mr C. V. A. Peel, F.Z.S., in.Somaliland, with descriptions of new

species. 9. Chilopoda and Arachnida. Proceedings Zool. Soc. of London, 1900,

Text, pp. 49-51 (relate to ticks), PI. Ill, Figs. 1-1 d, 2-2 f.

9*— 5

.

Bibliography 27

Pocock, R. I. (1903). See Christy, 1903, p. 188.

(1907), Article "Ticks," in A System of Medicine by many Authors, edited by

Allbutt and Rolleston (London : Macmillan), vol. II. Part u., pp. 187-203.
(Partial Bibliography, contains numerous inaccuracies.)

Pohl and Kollar (1823), Brasiliens vorziiglick liistige Insecten. Wien. (Argasidae:

title cit. by Huber, 1899, p. 14 ; inaccessible.)
Porter (1821), Travels. (Die giftigen "Wanzen in Persien. Froriep. Not. I. No. 12) :

cited thus by Huber, 1899, p. 16, as referring to A. persicus.

Railliet. A. (1886), Elements de zoologie me'dicale et agricole, pp. xv + 1053, Figs. 705,
8°. Paris.

(1893), Traite de zoologie me'dicale et agricole, fasc. 1, pp. 736, Figs. 494.

(1895), Traiti de zoologie me'dicale et agricole, 2nd ed. Paris : Asselin et

Houzeau. (Ticks: pp. 703-718, Figs. 478-488.)

Raspail, F. V. (1839), Rech. d'hist. nat. sur les insectes morbipares. Oaz. des
hSpitaux (2), I. p. 9. (Cited by Terrenzi, 1893, pp. 73-76, and Braun, 1895,
p. 258, in connection with Argas reflexus.)

"R. E." (1874), Argas reflexus. Science Gossip, 1874, p. 161.

Reaney, M. F., (V. 1907), Spirillosis in domestic fowls. Brit. Med. Journ., I. 1118.
(Only a note.)

(XI. 1907), Fowl spirillosis. Ind. Med. Oaz., xlii. 401, 402. (2 Figs., bad.)

Reclus, A. (1880), Explorations aux isthmes de Panama et de Darien, en 1876, 1877,

1878. Le Tour du Monde, 1880, xxxix. 396-398.
Riley, C. V. (1887), Poisonous Insects. Reference Handb. Med. Sci., v. 741-760,

Figs. 2971-3020. (Cited in Bibl. by Salmon and Stiles, 1901.)
Riley, C. V., and Howard, L. 0. (IV. 1895), A new chicken plague in Texas, and

the cattle tick affecting horses. Insect Life, v. 267, 348. (Further abstracts of

papers on ticks in general (unimportant), Ibid., vi. 58, 165.)
Risso, A. (1826), Histoire naturelle des principales productions de V Europe meridionale

et principalement de celles des environs de Nice et des Alpes Maritimes, vol. v.

viii + 402, Pis. I-V, Paris. (Cited by Salmon and Stiles, 1901.)
Rivolta and Delprato (1880), Ornithojatria, p. 314. (A. reflexus, cit. Huber, 1899,

p. 16.)
Robertson, F. H. (1905), Fowl tick experiments. Journ. Dep't Agric. West Australia,

xii. 561-563. (Cited by Hunter and Hooker, 1907.)
Rollinat, R., and Trouessart, E. (1897), Sur la reproduction des chauves-souris.

II. Les Rhinolophes et Note sur leur parasites dpizoiques. Mem. de la Soc.

Zool. de France, x. 114-138. {See re Ixodidae, pp. 136, 137.)
Roncisvalle, D. M. (1899), Intorno agli effetti morbosi degli Ixodidi sull' uomo.

Atti dell' Accad. Gioenia di Sc. Nat. in Catania, lxxvi. (Cited by Perroncito

(1901, p. 562) re ticks attacking man. Also cit. in Entomol. News, Philadelphia,

xi. p. 506.)

SaUi (1849), Rev. Mag. Zool., p. 342. (Description of sufferings from Argas {Orni-
thodoros) talaje, translated in Murray, A., 1877, pp. 183, 184.)

Salmon, D. E., and Stiles, C. W. (1901), Tiie cattle ticks (Ixodoidea) of the United
States. 17th Annual Report of the Bureau of Animal Industry, U.S. Dep't of
Agric, Washington, pp. 380-488, Figs. 42-257.

Bibliography • 29

Sambon, L. W. (1900), Ticks ami Tick Fevers. Journ. of Tropical Med., n. 217-223.
Sanchez, J. (1893), Acoidentes prodaoidos por his garrapatas. In Sanchez's Zool.

mdd. mexic, 8°, Mexico, 21.
Savigny, J. (1826), Description de UEgypte. Plauches de zoologie. Arachnides,

PI. IX.
Schellack, 0. (18. IV. 1908), Uebertragungsversuche der Spirochaete gallinarum

durch Argas reflexus Fabr. Centralbl. f. Bah., xlvi. pp. 486-488. (A full

account to appear in Arb. a. d. Raised. Gesundheitsamte, Berlin.)
Schlimmer, J. L. (1874), Terminologie medico-pkarmaceutique et anthropologique

francaise-persane sur les maladies ende'miques et particulieres les phis interessantes

des Imbitants de la Perse. Teheran. (Extracts cited by Bordier, 1882, p. 132.)
Schnee (1908), Ueber das Vorkoiumen von Argas in Deutschland. A rch. f. Schifs-

u. Tropen-Hyg., xu. 32,
Schneidemiihl, G (1896), Lehrbuch der Verglcichenden I'atlwlogie and Therapie des

Menschen und der Hausthiere far Thierdrzte, Arzte und Studiremle, 2. Lief.,

pp. 209-448, 8°. Leipzig.
Siebold, K. T. E. von (1850), " Parasiten," in Handwiirterb. d. Physiol, mit

Riicksieht auf physiol. Path., Braunschweig, II. pp. 641-692. (Cited in Bibl.

by Salmon and Stiles, 1901.) %

Simpson, J. C. (27. IV. 1901), Case of a parasite — " Argas (or Ornithodorus) megnini

Duges — in each ear. Lancet, 7 pp., 3 Figs. repr.
Spicer, W. W. (1874a), The Canterbury tick. Science Gossip, lxxiv. p. 185.

(1874b), Ibid., p. 209.

Stiles, 0. W. (See Salmon and Stiles.)

Stiles, C. W., and Hassall, A. (26. III. 1901). Notes on parasites, 55-57. A pupa-
like stage in the development of the spinose ear tick (Ornithodoros megnini) of

cattle. U.S. Dep't of Agr. Bureau of Animal Industry, Circular No. 34, 2 pp.,

repr.
Sundvall, C. J. (1833), Conspectus Arachnidarum Lund. In Acad. Carolina.

(Michael, 1883, p. 34, gives SundvalPs classification.)
Szovits (1828), Ueber die Wanzen von Miana. Froriep. Not., xxn. No. 478.

(A. persicus, cit. Huber, 1899, p. 16.)

Taschenberg, E. L. (1873), Mitteilung iiber die einheimische Saumzecke. Zeitschr. f.

d. gesammt. Naturwiss., xli. 381. (Cited by Brandes, 1897.)
(1874), The Canterbury tick (Argas reflexus). Science Gossip, 1874, p. 171.

(1880), Praktische Insektenkunde. 5. Theil. 238 pp., 43 Figs. Bremen :

M. Heinsius. (Ticks: pp. 150-153; describes Ixodes spec, Argas reflexus, and
gives 2 small original figures, the large adult Argas (!) has only 6 legs.)

(1900), Die lnsekten Tausendf ussier und Spinnen, being vol. ix. of Brehm's

Thierleben. Leipzig u. Wien. 764 pp. (Ixodoidea ; pp. 733-741.)

Taschenberg, 0. (date?), Einiges iiber niitzliche und sohadlicheThiere. Die Natur,
xlvi. (Cited by Lucas. Lucas, 1898, says the paper is interesting re poison
of Argas ; inaccessible.) v

Terrenzi, Q. (1893-1894), Sopra an Acaro (Argas reflexus) trovato per la prima volta
in Italia parassito sull' uomo. lliv. ital. Sci. Nat., XI v., 73-76, 97, 98.

Bibliography 31

Tholozan, J. D. (1882), Dos phenomenes morbides produits par la piqure des

parasites voisins des Ixodes, les Argas de Perse. Mem. Soc. Biol., p. 15.

(A. persicus, cit. Huber, 1899, p. 17.)
Tonnel (14. XI. 1906), L' Argas reflexus parasite do l'homme. Semaine Med., No. 46,

p. 552. (Abstract of paper which appeared in a recent number of VEcho me'd.

du Nord.)
Townsend, C. H. T. (1893), Ticks in the ears of horses. .Town. N. Y. EiUomol. Soc.,

i. (2), June, pp. 49-52. (0. megnini.)
Treviranus, G. R. (1831), Ueber den Ban der Nigua {Acarus americanus L., Acarus

nigua De Geer). Zeitschr. f. Physiologie. Heidelberg and Leipzig, iv. (2),

pp, 185-191, Taf. xv. xvi. (Cited by Salmon and Stiles, 1901.)
Trouessart (1891 and 1892), Considerations generales sur la classification des

Acariens, suivies d'un essai de classification nouvelle. Revue des sci. nat. de

Vouest, Paris, I. (4), Oct., pp. 289-308; n. (1), Jan., pp. 20-54. (Cited in

Bibl. by Salmon and Stiles, 1901.)
Trouessart, E. L. (no date, ca. 1896 ?), Les parasites des habitations humaines, 168 pp.,

53 Figs. Paris: Massou. (Ticks: pp. 122, 123, Fig. 50; little about ticks,

and nothing original.)

Utility (1899), The poultry tick. The Agric. Gaz., Tasmania, vi. (10), April, p. 185.
(Inaccessible.)

Verrill, A. E. (1870), The external parasites of domestic animals : Their effect and
remedies. Fourth Annual Report of the Secretary of the Connecticut Board of
Agriculture (1869-70), 1870, pp. 72-122, Figs. 1-47. (Cited in Bibl. by Salmon
and Stiles, 1901.)

Wagner, E. (1876), Manual of General Pathology, translated from the sixth German

edition by John van Duyn and E. C. Seguin, pp. x+728, 8°, New York, Wm.

Wood and Co. (Brief mention of ticks. )
Walckenaer, C. A. de, and Gervais, P. (1837-47), Histoire natureUe des Insectes

(Suites a Buffon). Apteres. 4 vols., 52 Pis. Paris : Roret. (First work on

general classification of Aptera. Apteres: pp. 234-251.)
Ward, H. B. (1900a), The ticks of Nebraska. Ann. Rep't Nebraska State Board of

Agricult., 1899 ; also as "Studies from the Zoological Lab. Univ. of Nebraska"

pp. 193-205 (repr.), Kigs. 1-6.
(1900b), Arachnida. Wood's Refer. Handb. of the Med. Sci. Revised edition, I.

428-437, Figs. 243-260.
Webster, F. M. (1893), Ticks on chickens. Ohio Farmer, Cleveland, lxxxiv. (8),

p. 149. (Inaccessible.)
Weed, C. M. (1888), A chicken tick. Prairie Farmer, lx. (1), Jan. 7, p. 3.

(Inaccessible. ) **

Weichselbaum, A. (1898), Parasitologic HaVdbuch der Hygiene, Jena, ix. (2 Lief.),

pp. x + 63-336, 78 Figs. (Cited in Bibl. by1 Salmon and Stiles, 1901.)

Bibliography

Wellman, F. C. (1905a;, Case of relapsing brer, witli remarks on its occurrence is
the tropic* and its relation to tick fever. Journ. Trop. Med., London, vm.
97 99.

(1905b;, Relapsing fever ; its occurrence in the tropica and ita relation to tick

fever in Africa. American Medicine, I'liila., X. 161 loo.

(1905c;, NotCH on the Tropical Diseases of the Angola Highland* The Amur.

8oc. of Trop. Mud., vol. i. 1904 1906, reprinted from : New Tori Med. Journ.
and PhUad. Utd. Journ. for Aug. 12, I!*, 26, and Sept.. 8, 1908 13 Ki^s., 4!» pp.,
reprint.

(1906a;, Human trypanosomiasis ami apirachaetoaia in Portugueae South \\'<-nt,

Africa, with suggestions for preventing their spread in the Colony. Journ. of
Hygiene, Cambridge, n. 237-245.

(1906b;, On a bemipterous inaeot which prays upon blood-aucking arthropoda

and win' nally attacks mammals (man;. Journ. Trop. Mid., Lond., IX.

(1906c;, Sohrc urn insscto hemiptero qne 08 alirnenta do propagador da Espiril-

ho.se hurnana e ataca occasionalmente o llorncm. Polytechnia, voll. in. No*. 4, r>,
7 pp., 1 Kig., reprint.

(1906d), A note on the habits of Orntihodoro* moubata (observations of two

gravid females). Journ. Trop. Med., Lond., IX. 216.

(1906 and 1907;, On (Jrnithodoro* moubata, Murray ; a disease-bearing

African tick. Medicine, Detroit, XII. 103 V.lU, and Am. Soe. Trop. Med., II.
bound reprints. 16 pp., 4 Figs.

(1907), Preliminary notes on some Isxlies found in tick* Ornith. moubata

'Murray,' fed on lilood containing embryos of /•'iloria.peritan* (Manson;. I!,*'
Mil. Journ., II. 142.

Werner (1906,, Zur Epidemiologic de* sJrikamsehen Recnrrans. Arch./. Schiff*- u.
Tropm ////,'/.. x. 770 781. (Discusses the epidemiology of African •prroobaetoeta

in Herman K. Africa and ita history, with incidental mention of ticks.;
WestWOOd, J. 0. (1877;, The President's Address. I'roe. linlomol. Sue., London,

1 J-S77, p]>. xxxvii lxxxi. (Kefers briefly to the " lilyborough tick ' on p. Ixii.)
Wheler, E. O. (1899 and 1900;, ( InaaHtmticn of I'.ritish Ticks. Science Ooeeip, I 899,

pp. S88 866 ; I960, pp. II, 12, 37 41(17 Figs.).

(1901;, Note on a remarkable stigmatic organ in the Nymph of Orwithodoro*

unijiiini Dngfee). 'i r'i^s. in text.; Journ. Quditt Microecop. Club, tut. 2, vm.
61, 02 ; also Journ. New Y<rrk Bntotnol. (foe., IH'.tt, pp. 19-62, also included under

Simpson, liXJi, q.v.

(1902;, Parasitieally inoculated diseases. Trant. of the Highland and Ayr. Sue.

of Scotland, 20 pp., repr.

(IIL 1906;, I'.ritish Ticks. Journal Agric. Hci., Cambridge, I. 100 I2!i,

PI* V X.

White, A. (1846;, List of Annulosa (principally Insects; found on the journey of
Henry H. Methuen, Ks<j., nee Appendix to H. 11. Methuen, Life in the Wilder-
net*, or Wandering* in Son//, Africa; p. 303, PL II, Kig. 4.

Wickson, E. J. 1892,, Ticks in the e*A Pacific Rural Pre**, March 19, p. 264.
(Inaccessible.)

Bibliography 35

Wood, H. C, and Fitz, R. H. (1897), The Practice of Medicine, pp. x + 1088, 8°,
Philadelphia. (Cited in Bibl. by Salmon and Stiles, 1901.)

Woods, P. T. (1898), Insects affecting poultry. Farm Poultry, ix. (6), March 15,
pp. 102, 103, Figs. 19, 20. (Cited in Bibl. by Salmon and Stiles, 1901.)

Ziirn, F. A. (1872), Die Schmarotzer aufund in dem Korper unserer Haussaugethiere,
some die durch erstere veranlassten Krankfieiten, deren Behandlung und Verhutung.
Theil I : Die thierischen Parasiten. (4 Tafeln.) 236 pp. Weimar : B. F. Voigt.
(A very short account of ticks on pp. 37, 38, only 2 species mentioned, and one
of these, " I. ricinus ? ," badly figured. No original matter.)

(1882), Die Krankheiten des Hansgeflugeh, p. 78. (A. refleants, cit. Huber, 1889,

p. 16.)

>

V

#

TICKS

A MONOGRAPH OF THE

IXODOIDEA

PART II

••**

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iLonUon: FETTEB LANE, E.C.

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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

LIBRARY

ONTARIO BFSWi FOUNDATION

MAR

26 1964

43 QUcEM'S PARK
TORONTO 5

IXODIDAE

Section I. CLASSIFICATION
Section II. THE GENUS IXODES

by

GEORGE H. F. NUTTALL

and

CECIL WARBURTON

PREFACE TO PART 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, wheu 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 the 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 of 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 movhata, 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
conjunction 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 tlie 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. Howlett I t

3 ,, ,, C. Warburton [

1 was drawn by L. E. Robinson J

Of the 107 previously 1

.,.,,„ 1-48 are from publications by Neumann

published ngures)

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 Robinson,

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 ngures 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 Preface

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, however,
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 I. angustus (types of </ , 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

(i •">

EKKATA.

p. 107, lines 1 and 26,

for Sarconyssus
read Sarconissus

p. 134, line 4 from below,

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

for Dennanyssus Kolenati, 1857, p. 20

read Dennanissus Kolenati, 1856 and 1857, p. 20

*/

CONTENTS

OF
PART II

THE IXODIDAE

SECTION I
DEALING WITH THE CLASSIFICATION OF TICKS

PAQK

Historical Review relating to Classification, with Summary and Conclusions 105

The Superfamily IXODOIDEA

Synonymy and literature 113

The superfamily characters defined 114

Ihe Family IXODIDAE

Synonymy and literature 115

The family characters defined 115

Classification of the Ixodidae, together 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

xii Contents

PAGE

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

1. ricinus$, ?,o,L 143

„ with three varieties : scapularis <J , 9 156

ovatus 9 158

calif milieus <J, 9 159

2. nigricans 9 159

3. acuminatus 9 161

4. dentatus 9 162

5. diversifossus 9 163

6. minor (J, 9 164

7. boliviensis c?> 9> ° 166

8. fuscipes 9 169

9. sculptus 9 171

10. spinicoxalis 9 171

11. marxi 9 173

12. pratti (J, $ 174

13. rubidus $?,o • 175

14. hexagonus £, 9, o, L 177

„ with one variety : cookei <J , 9 183

15. nitens 9 185

16. bicornis $ 186

17. auritulus <j> 187

18. brunneus ?fO, L 189

19. cavipalpus $, 9 192

20. angustus <J , 9 > o , L 195

21. caledonicus <j? , o , L 198

22. fossulatus 9 201

23. acutitarsus 9 202

24. gigas $ 203

25. rubicundus <J , 9 204

„ with one variety : hmbatus 9 . . . . 207

26. simplex 9 > ° ■ . 207

27. japonensis 9 208

28. canisuga <J , 9 > o , L 209

29. texanus 9 214

30. stilesi 9 216

31. neumanni 9>o 217

32. percavatus 9 220

„ with one variety : rothschildi 9,0 221

33. pilosus <J , ? , o , L 221

„ with 1 one variety : howardi $ , <j> 226

34. lunatus 9 226

35. rasus <J, 9 • • • • • . 228

Contents xiii

PAGE

36. ugaiidanus cj, ? 230

„ with one subspecies : djaronensis £ , $ . . . . 233

37. cordifer $ 233

38. holocyclus <J > ? > ° 235

39. schillingsi ^ , ? , o 238

40. omithorhynchi ? , o . . . . . . .... . . 242

41. tasmani ? . . 244

42. tenuirostris <^,9,o,L. . . . . . . . . . 246

43. fecialis ? 248

„ with one variety : aegrifossus 9 250

44. australiensis 9 ........... 250

45. vestitus $,o,L 252

46. putus (?,?,o 256

47. signatus $, o, L 261

48. unicavatus 9 > o , L . . . . . . . . . . 264

49. loricatus £ , ? 266

„ with one variety : spinoms 9 . . . . . . . 269

50. coxaefurcatus $ .......... 270

51. vespertilionis {J,$?,o,L 271

Geographical distribution of the Genus Ixodes 278

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

and literature, by G. H. F. Nuttall 280

Notes on doubtful species 291

Notes on the biology of Ixodes, by G. H. F. Nuttall .... 294

with a special index and including two appendices :

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

Nuttall and G. Merriman 318

II. On the adaptation of Ticks to the habits of their hosts, by

G. H. F. Nuttall 324

Index to valid species of Ixodes, together with a list of the collections in

which the types are to be found . 346

LIST OF ILLUSTRATIONS

PLATES

Plate IV. Ixodes ricinus. foxing P- 144

Fig. 1. £, dorsum x 12.

Fig. 2. (J, „ (mounted specimen).

Fig. 3. ?, „ x 12.

Fig. 4. 9 > venter x 9.

Fig. 5. o, dorsum x 12.

Fig. 6. L, „ x 12.

Fig. 7. Copulation.

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

Fig. 1. /. tenuirostris £, venter.

Fig. 2. „ ? , „ X 1 1 (mounted specimen).

Fig. 3. I. putus <J , dorsum x 7.

Fig. 4. „ $ , „ x 7.

Plate VI. Ixodes vespertilionis. facing 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 ? (replete), dorsum and venter . 117

117. „ „ /. cavipalpus <J, capitulum, dorsal and ventral

aspects 118

List of Illustrations

xv

118. Illustrating Ixodes: I. cavipalpus 5 , capitulum, dorsal and ventral

aspects

119. „ „ I. bicornis $?, capitulum, ventral aspect

120. „ Haemaphysalis : H. Wellington! <J, dorsum and venter

§ , dorsum

121. „ Dermacentor : D. variegatus var. hamshadalus <J, dorsum

and venter

122. „ „ D. variabilis J , capitulum and scutum

123. „ Rhipieentor : R. bicornis $ and $ , dorsal aspects .

124. „ „ R. bicornis <$ , venter ....

125. „ Rhipicephalus : Rh. bursa $, dorsum

126. „ „ Rh. sanguineus £ , venter

127. „ „ Rh. simpsoni § j capitulum and scutum

128. „ Margaropus: M. winthemi (J, venter

129. „ „ M. winthemi 9 > capitulum and scutum

130. „ Boophilus: B. decoloratus $ , dorsum and venter

131. „ „ B. annulatus 5 , capitulum and scutum.

132. „ Hyalomma: Hyal. aegyptium <$ , dorsum and venter

133. „ „ Hyal. aegyptium $ , capitulum, dorsal and

ventral aspects

134. „ Amblyomma : A. cooperi £ , dorsum and venter
135 a. „ „ A. variegatum <J , part of venter

135 6. „ „ A. cooperi 9, dorsum

136 a and 6. Digit of chelicera of Haemaphysalis punctata $?

137. Spiracles of 15 species of Ixodidae ....

138. Tarsi (4th) of 10 species of Ixodidae

139. Ixodes ricinus £, dorsum and venter

140. „ „ (J, capitulum, dorsal and ventral aspects

141. „ „ jj, tarsi 1 and 4 ....

142. „ „ <£ and $> , digits and hypostomes

143. „ „ 9 , dorsum, part of venter, etc.

144. „ „ $ , capitulum, dorsal and ventral aspects

145. „ „ o, dorsum and venter

146. „ „ o, capitulum, dorsal and ventral aspects

147. „ „ L, dorsum and venter

148. „ „ L, capitulum, dorsal and ventral aspects

149. Ixodes ricinus var. scapularis 9 , capitulum and scutum, part of

venter, etc. ....

150. Ixodes nigricans ? , essential parts ....

151. Ixodes acuminatus ? , venter .....

152. „ ,, ?, hypostome and digit

153. Ixodes dentatus $? , capitulum, ventral aspect .

154. „ „ $ , coxae and trochanters .

155. Ixodes minor <J , venter

156. „ „ c?, venter

157. „ „ t?> basis capituli, dorsal aspect .

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 FAQB

158. Ixodes minor $, hypostome 165

159. „ „ $ , essential parts 166

160. Ixodes boliviensis £ , dorsum, venter, hypostome, spiracle, tarsus 4 167

161. „ „ ?, capitulum and scutum, part of venter, spiracle . 168

162. „ „ o, dorsum, venter, tarsus 4 168

163. Ixodes fuscipes ? , capitulum and scutum, capitulum in ventral

aspect, coxae, spiracle and tarsus 4 . . 169

164. „ „ ? , capitulum and scutum, coxae with part of

capitulum 169

165. Ixodes spinicoxalis ? , basis capituli and scutum, coxae with part

of capitulum 172

166. „ „ <j>, hypostome 172

167. Ixodes marxi ? , capitulum and scutum, part of venter, etc. . . 173

168. Ixodes pratti <J, capitulum and scutum, capitulum in ventral aspect,

coxae, spiracle and tarsus 4 . . . . 174

169. Ixodes rubidus $ , dorsum, part of venter, spiracle and tarsus 4 . 176

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

and tarsus 4 176

171. Ixodes hexagonus <J, dorsum and venter, spiracle and tarsus 4 . 178

172. „ „ J, (gorged), dorsum and venter, spiracle and

tarsus 4 179

173. „ „ 9 1 capitulum and scutum, parts of venter,

spiracle and tarsus 4 . . . . 180

174. „ „ ? , capitulum, dorsal, ventral and lateral aspects . 180

175. „ „ 9 > tarsi 1 and 4> highly magnified . . . 180

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

and tarsus 4 181

177. „ „ L, dorsum, venter, tarsus 3, hypostome . . . 181

178. Ixodes nitens <J , capitulum and scutum, parts of venter, spiracle and

tarsus 4 185

179. Ixodes bieornis <j> , part of venter, hypostome 186

180. Ixodes auritulus J, capitulum in ventral and dorsal aspects, with

scutum 188

181. „ „ $, hypostome 188

182. Ixodes brunneus 9 > capitulum and scutum, coxae, capitulum in

ventral aspect 191

183. „ „ 9 1 digit, hypostome, ventral aspect of basis capituli

with coxae I 191

184. Ixodes cavipalpus £ , dorsum, capitulum in dorsal and ventral aspects 193

185. „ „ £ and ?, spiracles and digits .... 193

186. „ „ $, capitulum in dorsal and ventral aspects . 194

187. Ixodes angustus <J, dorsum and venter, spiracle and tarsus 4 . 195

188. „ „ ^ , capitulum and scutum, parts of venter, spiracle

and tarsus 4 196

189. „ „ o , capitulum and scutum, parts of venter, spiracle

and tarsus 4 197

List of Illustrations

xvn

FIGURE

190. Ixodes

191. Ixodes

192.

193.

)»

194.

Ixodes

195.

Ixodes

196.

Ixodes

197.

Ixodes

198.

jy

199.

Ixodes

200.

Ixodes

201.

»)

202.

Ixodes

203.

»*

204.

yj

205.

»»

206.

207.

209. „

210. Ixodes

211.

212. Ixodes

213. Ixodes

214.

215.

Ixodes

216.

»>

217.

Ixodes

218.

»

219.

»

220.

)>

221.

j»

222.

Ixodes

223.

»

224.

Ixodes

PAOE

angustus L, dorsum, venter, tarsus 3 197

caledonicus $> , capitulum and scutum, parts of venter, spiracle

and tarsus 4 ..... . 199

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

and tarsus 4 200

„ L, capitulum, scutum, coxae, tarsus 3, anal grooves 200
fossulatm ? , capitulum and scutum, parts of venter, spiracle

and tarsus 4 201

acutitar&us $> > dorsum, digit, coxae ..... 202

gigas <$, dorsum and venter 204

ruhicundus £, venter, hypostome ...... 205

„ $ , capitulum and scutum, parts of venter, etc . 206

simplex 9 , capitulum and scutum ...... 208

japonensis $?, capitulum in dorsal and ventral aspects. . 209
„ J , dorsal and ventral aspects of the bod}', spiracle

and tarsus 4 209

canisuga <J, dorsum and venter 210

„ <£, capitulum in dorsal and ventral aspects . . 211

„ c£, hypostome 211

6, legs 1-4 211

„ ? , capitulum and scutum, capitulum in ventral

aspect . . 212

„ <j? , (gorged), venter, spiracle and tarsus 4 . . 212
„ o , capitulum and scutum, part of venter, spiracle

and tarsus 4 213

„ L, dorsum, venter, tarsus 3 213

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

and tarsus 4 215

„ ?, capitulum and scutum, parts of venter, spiracle

and tarsus 4 215

stilesi ?, capitulum and scutum, palp in profile, digit . 217
neumanni 9 1 capitulum and scutum, parts of venter, spiracle

and tarsus 4 218

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

and tarsus 4 219

percavatus 9 > capitulum and scutum 220

,, ? , fore part of venter . . .• . . . 220

pilosus (J, dorsum and venter, spiracle and tarsus 4 . . 222
„ ? , dorsum (without capitulum), venter, spiracle and

tarsus 4 223

„ 9 > capitulum in dorsal and ventral aspects . . 224

„ o, dorsum and venter, spiracle and tarsus 4 . 224

„ L, dorsum and venter, tarsus 3 and digit . . 225

lunatus 9 > dorsum 227

„ $, venter 227

rasus (J, venter, digit, hypostome 228

win

List of Illustrations

(replete), hypo

venter, spiracl
venter, spiracle

FIOCKE

225. Ixodes rasus 9 > dorsum, venter, tarsus 4, spiracle and figures showing

variability ....

226. „ „ ? , digit, hypostome, tarsi 1 and 4

227. Ixodes ngandanus $, venter, hypostome .

228. „ „ $? , capitulum and scutum .

229. Ixodes cordifer £, venter, tarsus 4 .

230. Ixodes holocyclus ,$, venter, hypostome

231. „ „ ? , capitulum and scutum, venter

stome ....

232. „ „ o , capitulum in dorsal aspect

233. Ixodes schillingsi <$, venter, tarsus 4, digit

234. „ „ 9 , capitulum and scutum, venter, spiracle and

tarsus 4

235. „ „ o , capitulum and scutum, parts of

and tarsus 4

236. Ixodes ornithorhynchi $> , (two specimens) dorsum and

237. „ „ ?, tarsi 1 and 4
328. „ „ $ , digit, hypostome .

239. Ixodes tasmani 9 > capitulum and scutum, hypostome, end of tarsus

1 with foot .

240. „ „ ? , dorsum, venter, spiracle

241. Ixodes tenuirostris £, capitulum

242. „ „ (J, part of venter .

243. „ „ $? , capitulum and scutum

244. „ „ 9 1 ventral aspect of capitulum, with coxae I
245 a and b. Ixodes fecialis $ , capitulum and scutum, ventral aspect of

capitulum with coxae I

246. Ixodes fecialis var. aegrifossus $? , capitulum and scutum.

247. Ixodes australiensis J , capitulum and scutum, capitulum in ventral

aspect

248. „ „ $? , venter, spiracle and tarsus 4

249. „ „ ? , ventral aspect of capitulum and coxae I

250. Ixodes vestitus $? , capitulum and scutum

251. „ „ $, capitulum in ventral aspect, and essential

parts

252. „ „ o, dorsum, parts of venter, leg 4, etc.

253. „ „ L, dorsum, venter, leg 3 .

254. Ixodes putus $ , dorsum, venter, side view, hypostome, spiracle

255.

256.
257.
258.
259.
260.

<% , dorsum, venter, side view of palp, spiracle

tarsus 4

9, dorsum

9 , capitulum in dorsal and ventral aspects
9 , scutum, digit, hypostome, tarsus 4

9 , 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 Illustrations xix

FIOCRE PAOE

261. Ixodes signatus ?, capitulum and scutum, capitulum in ventral

aspect, palp in profile, coxae, spiracle,

tarsi 1 and 4, hypostome .... 262

262. „ „ ^ , 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 $ , capitulum and scutum 265

266. „ „ o , capitulum and scutum, parts of venter, spiracle

and tarsus 4 265

267. „ „ L, capitulum and scutum, parts of venter and

tarsus 3 265

268. Ixodes loricatus <J , dorsum and venter 267

269. „ „ 9 > capitulum in dorsal aspect .... 267

270. Ixodes loricatus var. spinosus 9 , capitulum, scutum, parts of

venter, etc. 269

271. Ixodes coxaefurcatus £ , dorsum, venter, spiracle, tarsus 4 . . 270

272. Ixodes vespertilionis <$ , venter 273

273. „ „ £, capitulum in dorsal aspect .... 273

274. „ „ £ , digit and hypostome 273

275. „ „ ,$, leg 1 273

276. „ „ J, 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 moubata $ , 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 Subject1.

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
Argas 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 Cynorhaestes (Ixodes)
under Acarina, but kept them far apart, not recognizing their close
relationship. Leach (1815, p. 387) called the Acarina Monomer osomata.,
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. Duges (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 (Ixodides, in English, with four genera :
Hyalomma, Haemalastor, Amblyomma and Ixodes) ; and 3. Rhipi-
stomiden (with four genera : Dermacentor, Haemaphysalis, Rhipistoma
and Rhipicephalus) ; 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

1 A number of classical authors refer to ticks. See Bibliography under Aristotle,
Cato, Varro, Columella, Pliny, Sammonius, Aldrovandi.

N. I. 8

106 Classification

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. Furstenberg
(1861, p. 208) took over the classification of ticks from C. L. Koch,
without making any change. Donnadieu (1875) included Argas under
Ixodidds. Megnin (1876 a, p. 293; 1877, p. 86; 1880, p. 117; and
1892, p. 25) gave the Ixodid6s 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. Gonipalpi (Genera
Phaulixodes1 [Berlese], Rhipicephalus, Dermacentor, Haemaphysalis
and Herpetobia1 [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 II. Antistomata, comprising three
families: 1. Haemalastoridae (with two genera: Haemalastor and

1 Since condemned, recognized as nymphal forms : Phaulixodes = Rhipicephalus,
Herpetobia = Haemaphysalis,

Classification 107

Sarconyssus), 2. Ixodidae (with three genera : Ixodes, Amblyomma and
Hyalomma), 3. Rhipistomidae (with five genera: Boophilus, Rhipi-
cephalus, Dermacentor, Rhipistoma and Haemaphysalis). Soon after-
wards, Banks (1894, p. 209) established the superfamily Ixodoidea,
which we retain in this work.

Neumann (1896, p. 2) placed ticks (Ixodidds) 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 ami 1899, p. 107) recognized three tribes under
Ixodinae : Tribe 1. Ixodae (= Gultripalpi of Canestrini, including
four genera: Ixodes, Hyalomma, Amblyomma and Aponomma); Tribe 2.
Rhipicephalae (= Conipalpi of Canestrini, including three genera :
Rhipicephalus, Dermacentor 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 J1 capitulum, and gives Haemalastor only
generic rank after Ixodes. He founded the genus Aponomma (1899,
p. 180) to include Amblyom ma-like forms, differing from Amblyomma
in not possessing eyes, and in having broader bodies, besides occurring
almost exclusively on Ophidia and Sauria. 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 Phaulixodes 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
Dume"ril, Cynorhaestes Hermann are referred to as "des synonymes plus
ou moins precis A' Eschatocephalus et d' Ixodes"; Pseudixodes Haller 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
annulatus (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, Amblyomma
and Hyalomma), 2. Rhipicephalae (Genera Haemaphy sails, Rhipicephalus
and Dermacentor); and II. Argasinae (Genera A rgas 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 superfamily, 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 : Superfamily
Ixodoidea comprising two families: I. Argasidae (Genera Argas
and Ornithodoros), and II. Ixodidae, including two subfamilies:
1. Rhipicephalinae1 (Genera Rhipicephalus, Boophilus, Haemaphy salis
and Dermacentor), and 2. Ixodinae (Genera Ixodes, Eschatocephalus,
Aponomma, Amblyomma 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,

1 Lahille (1905, p. 12) remarks that the subfamily should be styled Dennacentorinae
and not Rhipicephalinae, if the rules of nomenclature are to be strictly adhered to, for
Dermacentor (Koch, 1844, p. 235) has priority over Rhipkephalm (Koch, 1844, p. 238).

Classification 109

for it brings Ixodes, Aponomma and Haemaphy salts together on the
one hand, and Amblyomma, Hyalomma, Rhipicephalus and Dermacentor
on the other. Neumann considers it essential to take as a basis of
classification 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 Amblyomma 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 Amblyomma
(Aponomma1), Dermacentor and Haemaphysalis. (Amblyomma 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 Aponomma

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. Amblyommeae (for Anopli).
Neumann characterizes the sections as follows :

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
accompanied 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 : j" venter devoid of shields. Anal groove as
under 2. Eyes often present. Capitulum as under 2.

1 We regard Aponomma as simply a subgenus of Amblyomma.

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, Margaropus1, Hyalomma), and III. Amblyommeae (Amblyomma,
Dermacentor, Aponomma, Haemaphy sails). Lahille (1905, pp. 11
et seq.) places ticks in a suborder, Arpagostoma'2, comprising two
families : I. Argasidae, and II. Ixodidae ; he divides the latter into
three groups according to the <?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, Rhipicephalus and Boophilus (all possessing eyes); and,
finally, Group 3. 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, 1844 ; 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

i

Prostriata Metastriata

, i

Brevirostrata Longirostrata

-U

— I i 1

Group 1 Group 2 Group 1 Group 2

Argas Ixodes Haemaphysalis Dermacentor Hyalomma Amblyomma

Omithodoros3 Rhipicentor4 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 (viii. 1907,

1 Meaning Boophilus Curtice, and including Margaropus Karsch.

2 Should be Harpagosloma (from harpagon = a grappling iron).

3 We have not hitherto adopted the latinized transcription Ornithodorus but shall do
so henceforth in accordance with the rules laid down in The International Code of
Zoological Nomenclature. l Since added.

Classification 111

p. 62, as stated in Banks, 1908, pp. 13, 14) advanced another classifica-
tion. He divides Ixodidae into two subfamilies : I. Ixodinae {Ixodes
and Ceratixodes1), 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. RMpicephalini (including Rhipicephalus and Margaropus).

2. Haemaphysalini {Haemaphysalis).

3. Dermacentorini (Dermacentor).

4. Amblyommini (Amblyomma, 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 annulatus 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 Ceratixodes 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.

112 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 Amblyomma from Dermacentor." Donitz
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 Amblyommeae
of Neumatm. As Donitz points out, Amblyomma are not Anopli; a
number possess small chitinous plates1 (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 Boophilus
and Rhipicephalus, i.e. Tetraopli (Rhipicephaleae), and not to Poliopli
(Ixodeae). Banks (1908, p. 13) very properly criticizes the classification
of Lahille (190-5), which is based on the characteristics of the </\ and
thus separates Hyalomma widely from Amblyomma, 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 Amblyomma are not Anopli, aud 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 : Amblyomma, 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-
blyomma. The genus Rhipicentor Nuttall and Warburton, 1907, however,
directly links Rhipicephalus to Dermacentor, as is indicated by the
1 " Plaques " in our descriptions, see Fig. 135 a.

Ixodoidea 113

name, and the discovery of this intermediate type confirms the view
advanced by Warburton. Banks agrees with us in not placing Hyalomma
near Rhipicephalus, as has been done by Neumann (1907 a, p. 26).

Summary.

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 contrast 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
with 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. Claus, 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 (6), 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 capitulum, of a
special structure. It consists of a basal portion (basis capituli), a pair
of palps, protrusible chelicerae 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, are divided into the suborders Vermiformia,
Astigmata, Metastigmata, Helerostigmata, Prostigmata, Notostigmata.
Besides the Ixodoidea, the Metastigmata include the Oribatidae and the GamaMae.
The only mites in the least likely to be confounded with the Ixodoidea are the
remarkable group of Qekobiidae, 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 I) and II. the Ixodidae.

Ixodidae 115

II. Family IXODIDAE Murray, 1877.
Synonymy and Literature:

Family Ixodei Duges, 1834 a, p. 15; 1834 c, p. 33.

Family Ixodiden Koch, 1844, p. 220; 1847, p. 13. Fiirstenberg, 1861, p. 208.

Family Ixodides Gervais and van Beneden, 1859, p. 460.

Family Ixodidae Murray, 1877, p. 185. Pavesi, 1884, p. 483. Canestrini, 1890,
p. 530 ; 1892, pp. 563, 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 et seq. Banks, 1907, p. 62. Hunter and
Hooker, 1907, p. 46. Mauson, 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 Ixodides Megnin, 1880, p. 120.

Subfamily Ixodidae Berlese, 1885, p. 131.

Tribe Ixodines 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 ; (6) p. 435.
Neumann, 1904, p. 444; 1907 a, p. 26. Howard, 1908, p. 89. Blanchard,
1910, pp. 55-77. Brumpt, 1910, p. 510.

Tribe Ixodinae Neumann, 1892 b, p. 96.

Group or Tribe Antistomata Marx, 1892 a, p. 233.

Family Characters1. Scutate ticks with 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 % capitidum.
Where eyes are present they are 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, Haemaphysalis, Dermacentor,
Rhipicentor, Rhipicephalus, Margaropus, Boophilus, Hyalomrna and
Amblyomma (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 ,

Prostriata Metastriata

i

Brevirostrata Longirostrata

~i i"

Group 1 Group 2 Group 1 Group 2

I I > J

1. Ixodes 2. Haemaphysalis 3. Dermacentor 8. Hyalomma 9. Amblyomma

4. Rhipicentor (Aponomma)

5. Rhipicephalus

6. Margaropus

7. Boophilus

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 ; in 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

- *l<4/l#j&

J-/UA-

Fig. 115. J. ricimis (Linn.), 1 : in dorsal and ventral aspects (from Nuttall, 1908,

G. H.F.N, del.).

jciv&Avn-

-rjr^aJL

Fig. 116. J. hexagonus Leach, 1815, ? (replete): in dorsal and ventral aspects.

(Original, G. H.F.N, del.)

118

Ixodidae: Genus Ixodes

Fig. 117. I. cavipalpns Nuttall and Warburton, 1908, l : capitulum in dorsal and
ventral aspects (from Nutt. and Warb., G. H. P. N. del.).

~n^faj&*>u

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. I. bicornis Neumann, 1906, ? : capitulum in ventral aspect to show auricula.

(Neumann del.)

Ixodidae: Genus Haemaphi/mlix

119

SECTION II. METASTRIATA.

With anal groove contouring the anus behind.

Note : In two genera, Boophilus and Margaropus, 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) Brevirostrata.
Group 1.

Genus (2) Haemaphysalis : inornate, without eyes but with
festoons; with usually short conical palps 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

,-;*^S*/**£*4

£a/A*f<*<tZ*c&

fatt- «*u/«</i

Fig. 120. //. wellinfjtoni Nnttall and Warburton, 1908, s in dorsal and ventral aspects
and ? in dorsal aspect (from Nutt. and 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 reticulatus (Fabricius).

DERMACENTOR

?*c£ <*-c <L*L> /^

Fig. 121. D. variegatus var. kaimhadalus Neumann, 1908, i : 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 Rhipicentor

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 iu both sexes. The <f resembles Rhipi-
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.

RHIPICENTOR

Fig. 123. R. bicornis Nuttall and Warburton, 1908, s and ? : dorsum (from Nutt. and

Warb., E. W. del.).

CeKA. -Z"

Fig. 124. R. bicornis, i : venter, etc., as under Fig. 123 (G. H. F. N. del.).

122

Ixodidae: Genus Rhipicephalus

Genus (5) Rhipicephalus : usually inornate, with eyes and festoons,
with short palps and basis capituli usually hexagonal dorsally. (The £
of one species, Rliipicephalus pulchellus, has a Dermacentor-like capitu-
lum, and both sexes are ornate.) Coxa I bifid. The £ possesses a pair
of adanal shields and usually a pair of accessory adanal shields ; some
Js, when replete, show a caudal protrusion. Spiracles bluntly or
elongate comma-shaped. Figs. 125-127.

Type species : Rhipicephalus sanguineus (Latreille).

RHIPICEPHALUS

-^aXuAa/ jiAmrt.
■fait- &A*

Fig. 125. Rh. bursa Canestrini and Fanzago, 1877, s : dorsum. (Original, L. E. R. del.)

■S/viA.a*JiL

d&UaJ.*

jurloie ablest-

3»

Fig. 126. Fig. 127.

Fig. 126. Rh. sanguineus (Latreille, 1804), <f : 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.).

Ixodidae: Genus Margaropus

123

With anal grooves obsolete.

Genus (6) Margaropus1 : inornate, with eyes, hut without festoons,
with short palps and capitulum intermediate between that of Rhipice-
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 two 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 £ shows a caudal protrusion. Anal groove obsolete.
Spiracles rounded or short-oval in both sexes. Figs. 128, 129.

Type species : Margaropus winthemi Karsch.

MARGAROPUS

/ &L£cc&

. qj^*u£a I irk^ict

-ha^fi. OLbCuA,

Fig. 129.

Fig. 128.

Fig. 128. M. winthemi Karsch, 1879, i ■ venter (from Neumann, 1907, Nn. del.).
Fig. 129. M. winthemi Karsch, 1879, ? : capitulum and scutum. (Original, sketch
by C. W.)

1 This name has become established. It should read Hargaritopns, signifying beady-
legged.

9—2

124

Ixodidae: Genus Boophilus

Genus (7) Boophilus: inornate with eyes, but without festoons ; with
very short compressed palps ridged dorsally and laterally ; basis capitidi
hexagonal dorsally ; slightly chitinized; the unfed adults of small size.
Coxa I bifid. Anal groove obsolete in ?, faintly indicated in $ . The
% with 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

,A«~&t ttiAUU

Fig. 130. B. decoloratus (Koch, 1844), <$ : 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.).

Ixodklae: Genus Hyalomma

125

(b) Longirostrata.
Group 1.

Genus (8) Hyalomma: ornamentation absent or present, at times
confined to the legs ; with eyes, with or without festoons, with long palps
(shorter in Hyalomma monstrosum £ ) and basis capituli subtriangular
dorsally. The ? approaching Amblyomma. The £ with a pair of
adanal shields, and with or without accessory adanal shields and two
•posterior abdominal protrusions capped by chitinized points. Coxa I
bifid. Spiracles comma-shaped. Figs. 132, 133.

Type species : Hyalomma aegyptium (Linnaeus).

HYALOMMA

CH/i+taJUisui

p&xU&l/gA

Fig. 132. Hyal. aegyptium (Linn.), t : dorsum and venter (from Warb. and Nutt. 1909,

G. H.F.N. andE.W. del.).

Fig. 133. Hyal. aegyptium (Linn.), ? : capitulum in dorsal and ventral aspects. (Original,

G. H.F.N, del.)

126

Ixodidae: Genus Ambly omnia

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 J1 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 : Aponomma = Amblyomma, 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

-tdjL* ca/t^i^-

^viA.astU&

avitut aA^nrt^e

Fig. 134. A. cooperi Nuttall and Warburton, 1908, s : dorsum and venter (from Nutt.

and Warb., E. W. del.).

tJLO.. fu'G-u

<i&*£~.

Fig. 135 a.

Fig. 135 b.

Fig. 135 a. A. variegatum (Fabricius, 1794), s : posterior part of venter, showing plaques.

(Original, G. H. F. N. del.)
Fig. 135 6. A. cooperi, ? : dorsum, schematized (from Tick*, 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 : —

3, $, o, L, signify male, female, nymph and larva respectively; rfs, <j?s and os
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.

W. = its width at the widest part. Where the measurements are given as xxy,
measurement x stands always for the length and y for the breadth. (I. and
w. refer to length and width of various structures.)
Capitulum = the "rostrum," "head" or "false head" of various authors.

Basis capituli, or, shortly, base = the 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 £ 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 capituli, and
bearing recurved teeth. 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. 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 tip bears a number of very minute denticles.

128

Terminology

Chelicerae = the "mandibles" or " pseudo-chelicerae " of some authors, paired
structures lying dorsally to the hypostome. The terminology usually
applied to the portions of the digit 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 :
Internal article — "median apophysis" or "immovable finger" of some authors.

Dorsal process =" internal apophysis" of some authors.
External article = " external apophysis" or "free" or "movable finger" of
some authors.

We attach little importance to the description of the chelicerae in routine
systematic work.

1.

■2.

EXTERNAL ARTICLE';— ...

DORSAL PROCESS.

FLEXOR TENDON.-
EXTBNSOR TENDON

EXTERNAL ARTICLE

Fig. 136,1.

INTERNAL ARTICLE.

Fig. 136 B.

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 Robinson, 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, especially articles 2 and 3,

Term inology 129

may be fused. In all Ixodidae the 4th 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.

Scut um = the "dorsal shield," " Kopfschild," etc., of some authors. The size is

stilted by giving the length first, then the greatest width (including the

eyes, if protruding), thus -9 x "7 mm.
Pseudoscutum = that portion of the <$ scutum which corresponds in shape and

position with the 5 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.
Scapulae — the anterior angles or "shoulders" of the scutum projecting on

either side of the emargination and included in measurements of the length.
Cervical grooves = the 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 <J 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 <J , starting at the

postero-lateral scutal border (they correspond to the <J 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 fields = 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 = the " submedian dorsal porose areas " of Salmon and Stiles, two small

spots often observable near the middle of the scutum in the <$ ; posterior

to the scutum in the ? . (Figs. 121, 123, 130.)
Marginal fold = a raised fold of the body, external to the scutum, as seen in

Ixodes (see Fig. 115).
Venter = the whole ventral surface of the body.

Genital orifice : situated anteriorly, posterior to the capitulum, and in the

median line. Called vulva for brevity's sake in the $> .
Apron : a delicate chitinous flap arising in front of the sexual orifice which it

covers.
Genital grooves: starting at either side of the genital orifice and running

130 Terminology

backward between the coxae, then externally to the anal grooves toward
the posterior body-margin which they frequently attain.

Anus 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 {Prostriata) 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 Metastriata 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 structures occurring in Ixodes ^ ,
not rising above the surface of the body (as do the shields, for instance,
in Rhipicephalus, etc.) ; the plates are bounded by the ventral 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
Amblyomma, and situated ventrally in front of the festoons. (Fig. 135.)

Shields (adanal and accessory) are salient chitinous structures occurring in the
males of Rhipicephalus, Boophilus 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 bifid, 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 {pulvillum), should also be noted. (Fig. 138.)
Punctations are circular depressions dotting the integument, and frequently

Terminology

131

bearing hairs. The parts, scutum, capitulum, 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, Amblyomma and Dermaeentor
are usually ornate.

«£*i

/ i 3 ♦ J

Fig. 137, illustrating the spiracles of 15 species of Ixodidae to show the different 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.

11—15.
11.
12.
13.
14.
15.

long axis of spiracle transverse
to body axis.

long axis normal, i.e.
directed forward.

Round as in J. ricinus $ .

Bluntly oval as in I. boliviensis ? )

Elongate sub-oval as in I. tasmani $ )

,, oval as in I. angustus ;
Ovoid, flattened posteriorly (or sub-triangular) as

in I. angustus s

With slight postero-dorsal protrusion as in Haemaphysalis hystricis s .
With distinct darkly ehitinized 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 Boophilus annulatus ? .
Short comma-shaped as in Dermacentor variabilis ? .
„ „ „ Amblyomma longirostrum ? , with frame broad-

ened dorsally.
Sub-triangular, with rounded angles and with frame broadening much

dorsally as in A mbly omnia geoemydae ? .
More or less elongate comma-shaped spiracles :
As in Rhipicentor nuttalli i .

„ Dermacentor reticulalus £ , and Rhipicentor bicornis s •
„ Rhipicejihalus sanguineus s , and Amblyomma versicolor i .
,, Dermacentor andersoni 3 .
,, Hyalomma aegyptium S , and Rhipicephalus sp. S .

The "macula" is central in (1), eccentric in the others, being situated ventrally and
anteriorly, and possessing a variable form.

132

Terminology

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. Tapering gradually as in I. ricinw.

2. ,, „ „ I- signatus.

3. ,, obliquely „ I. ornitlwrhynchi.

4. Humped prior to tapering as in I. liexagonw.

5. Tapering abruptly as in I. cordifer.

6. . to one Bpur as in Haemaphysalis cornigera.

7. Ending bluntly with two spurs as in Bhipiceplialw masseyi.

8. Tapering to a long spur as in Margaropus winthemi.

9. ,, abruptly and bearing two spurs as in Amblyomma cooperi.

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 (de Geer, 1778 ;
Fabricius, 1794 ; Panzer, 1795). See further under list of species 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. Duges, (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.
Qen6 (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. lvii ; Verrill1,
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, O2).— 1881. Haller,
(a), p. 165 ; (6), p. 380 (mouth-parts).— 1882. Haller, p. 309.— 1883. Braun,
p. 211.— 1885. Berlese, p. 131 ; Baymondaud, p. 129.— 1886. Ludwig, p. 612.
—1888. Railliet, p. 496 ; Neumann, p. 89.— 1889. Berlese, fasc. lv, 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 ; Efisio1, p. 256 ;
Trouessart, pp. 20-47 ; Lewis, p. 449 ; Marx, (b), p. 232 (mouth-parts, classi-
fication) ; Bernard, p. 289 (respiration) ; 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.; Morgan1, 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, p. 142 (mouth-parts). — 1902. Neumann, p. 115 (genus Eschatocephalus

1 Inaccessible, cited in bibliography by Salmon and Stiles, 1901.

2 O 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 : Euixodes, Ceratixodes and Eschatocephalus, 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) ; Hunter 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 ; Heyden, p. 610. — 1831, Treviranus, p. 188, after which the name
fell into synonymy. (Referred to as Cynorrhaestes Hermann by Fabricius,
1805, p. 351, and Duges, 1834 c, p. 33 ; as Cynoraesthes by Latreille, 1829, p. 287 ;
as Cynochaestes by Megirin, 1880, p. 120 ; as Cynorhoestes by Trouessart, 1891,
p. 290.)

Crotonus Dumeril, 1822, p. 56. — 1829, p. 401 (ft 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 <% . 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 Eschatocephalus by various authors, see List of condemned species.

Note: Eschatocephalus was retained as a genus by Neumann (1902, p. 116)
until 1904, p. 445, when he made it a subgenus of Ixodes, because he considered
that the generic characters were insufficient to maintain the generic rank, the
differences "being only specific." He states that "the <J, 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 Eschatocephalus as a subgenus.

Dermany8SU8 Kolenati, 1857, p. 20 = Ixodes (ride List of condemned species).

Sarconyssus Kolenati, 1857, p. 21 (various "species " — Ixodes vespertilionis. See
List of condemned species).

Hyalomma Pickard-Cambridge, 1878, p. 222, Hyalomma puta = Ixodes pulus 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 <J as in Ixodes, but absent in the $ ; anal and adanal shields
in the $. Neumann, 1904, p. 444, subsequently recognized that the ? (/.
putus q.v.) differed so slightly from other "J Ixodes, that the genus could not
be retained ; the 0 was found to possess an anal groove and the $ only differs
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 Ceratixodes and Eschatocephcdus (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 always surrounded by a prominent marginal body-fold, and the male
venter presents seven non-salient chitinous plates, namely, a pregenital,
a median, an anal, two adanal and two 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

\ „ without such hairs 1

(Legs inordinately long. ..... vespertilionis 272

( „ moderately , .2

(Basis capituli with dorsal punctate excavation . coxaefurcatus 270
1 „ M without „ „ .3

136
3.

4.

5.
6.

9.

10.

11.

12.

13.

14.

15.

16.

17.

18.

Genus Ixodes

[Dorsum showing two lateral folds . . .4

| „ „ one „ fold . . .5

(Anal grooves divergent ..... loricatus
( „ „ convergent holocyclus

(Anal grooves convergent . . . . .6
( „ „ parallel or divergent . . .10

(Anal grooves approaching but not uniting behind 7
{ „ „ uniting behind. . . . .8

| Anal plate oval, basis capituli with median point

ventrally pilosus

Anal plate racket-shaped, basis capituli tricuspid
( ventrally schillingsi

(Coxa IV with long internal spur .... cordifer
I „ „ without long internal spur . . .9

(Tarsus 4 humped some distance from end . . rasus

\ „ „ tapering ugandanus

(A single distinct spur on coxa I . . . .11
(Two spurs or none 12

(Median ventral plate about as broad as long . hexagonus
( H ,, „ much longer than broad . ricinus

Coxa I with two spurs ..... 13

„ „ without spurs ; at most slightly pointed . 16

(Both spurs very long and close together . . gigas
1 „ „ otherwise ...... 14

(Both spurs short, subequal 15

(Inner spur long and strong, outer short. . . boliviensis

(Strong cornua ; anal grooves short, divergent . minor
(No cornua ; anal grooves long, parallel . . angustus

jTarsi humped canisuga

\ „ tapering 17

(Coxae trenchant tenuirostris

( „ not trenchant ...... 18

Punctations very fine, equal .... cavipalpus

„ unequal, nearly obsolete . . . rubicundus

266
235

221

238
233

228
230

178
147

203

166

164
195

210

246

193
204

Key to Species

137

Females.

1.
2.

3.

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
( „ „ sub-parallel or divergent

(Anal grooves horseshoe shaped .

( „ „ 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)

[Anal grooves a circle (Africa)

| „ „ 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, norma
\ (Australia) ......

(Coxae practically unarmed ....
(Some of the coxae with distinct spurs .

rScutum broadest quite in front (on marine birds)
1 „ ,, near middle or posteriorly

jScutum broadest behind the middle

I „ „ in middle or a little anteriorly

jScutum broader than long ....
| „ longer than broad ....

1 Palps with article 1 greatly enlarged on ventral
aspect
Palps with article 1 small ....

vespertilionis 274
1

2

8

3

5

lunatus
4

226

schillingsi 239

pilosus 222

7-asus 229
6

ugandanus 232

australiensis 250

holocyclm 236

9
20

putus 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.

I 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
| „ elongate ; no cornua (on bats)

(Sharp spurs on trochanters (Africa)

(No trochantal spurs

Coxae trenchant; scutum smooth, with fine punc-
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 .....
„ „ „ short; a slight external spur

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 posterior (Mexico) .
1 » „ long (Brazil)

tenuirostris
fecialis

247
248

vestitus
15

252

16
17

japonensis
simplex

208

207

cavipalpus
18

194

rubicundus

206

19

canisuga
texanus

212
214

21
23

percavatus
22

220

unicavatu8

264

signatus
24

261

33

25

27

acutitarsus
26

202

bicornis
fuscipes

186
169

Key to Species

139

27.

28.

29.

30.

31.

32.

33.
34.

35.
36.

37.

38.
39.
40.

41.

Cornua well marked, sharp

„ faint or absent .....

(Porose areas large, near together (N. America) .
1 „ „ small, far apart (America)

(Hypostome very long and pointed ; anus very
posterior (Italy)
Hypostome normal

IPunctations on scutum large and few ; coxal spine
very long (Sumatra)
Punctations small and numerous
(Anal grooves short ; auriculae strong
I „ „ long ; no auriculae .

(Porose areas transversely elongate (Madeira)
(possibly identical with ricinus)
Porose areas less elongate .
(Scutum nearly as broad as long .
( ,, elongate, distinctly longer than broad
(Anal grooves pointed in front ; tarsi humped
\ „ „ rounded in front ....

I Scutum oval; auriculae, but no cornua (America)
„ cordiform ; cornua, but no auriculae
(America)

(Article 1 of palp with anteriorly directed spur (on
birds, Brazil) .......
No such spur

(Basis capituli winged, pointed laterally (S.
America)
Basis capituli normal ......

(Tarsus 4 humped, or narrowing abruptly

( „ „ tapering more or le.ss gently

(Anal grooves rounded in front ....

\ „ „ 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)

28

29

sculptus

171

diversifossus

163

acuminatus

161

30

spinicoxalis

171

31

boliviensis

167

32

nigricans

159

ricinus

150

34

36

hexagonus

179

35

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 loDg and sharp ; basis capituli 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 ;

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) .

rubidus

175

marxi

173

■minor

165

44

brunneus

190

45

caledonicus

198

40.

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
arefuscipes, p. 170 ; brunneus, p. 190 ; simplex, p. 208.

With legs inordinately long
„ „ otherwise

1.

(Scutum without cervical grooves
1 » with

(Scutum with lateral grooves

I „ without lateral grooves .

(Coxae unarmed ....
( „ armed ....

(Palp with article 1 large and pointing forward
1 » „ „ „ otherwise

J With anal grooves fused to a point behind .
( >, h ., otherwise . , ,

vespertilionis
1

275

tenuirostris

248

2

3

9

vestitus

254

4

angustus
5

197

holocyclus
6

237

Key to Species

141

6.

7.

8.

9.

10.

11.

12.

13.

14.
15.
16.
17.

[With anal grooves in the form of a horseshoe
( „ „ „ parallel or divergent

jWith long palps, concave externally. .
„ palps otherwise . . ...
jWith tarsus 4 tapering
{ „ „ „ humped
(With coxae unarmed .
| „ „ armed
(With tarsus 4 humped
( „ „ ,, otherwise
(With very long scutum
[With short, very broad scutum

! 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

7

boliviensis
8

169

ricinus
rubidus

151
176

10

12

canisuga
11

213

pvi-us 259

ornithorhynchi 242

hexagonus

13

unicavatus

14

15

neumanni

schillingsi

16

signatus

caledonicus

pilosus

loricatus

181
265

219
241

263
199

224
268

Larvae.

N.B. There are 14 species whose larvae have been recorded, three
of which (fuscipes, p. 170; brunneus, 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
1 „ „ „ n present

vespertilionis 275
1

tenuirostris
2

248

142 Genus Ixodes

2.

3.
4.
5.
6.

(Scutum with lateral grooves

| „ without lateral grooves .

(Coxae unarmed

( „ with spurs

(Palp with article 1 large and pointing forward
( „ otherwise

(Capitulum with dorsal ridge pointing out laterally pilosus
\ „ otherwise

(Basis capituli with latero-ventral protuberances
| „ „ „ central ridge

7 (Basis capituli without auriculae ....
\ „ „ with auriculae ....

I Scutum with cervical grooves diverging widely
behind
Scutum with cervical grooves otherwise

9.

(Coxa IV with two spurs
( „ „ without two spurs

vestitus
3

255

4

7

angustus
5

197

pilosus
6

224

canisuga
hexagonus

213
181

unicavatus
8

265,

9
signatus

263

caledonicus

200

ricinus

152

143

SPECIFIC DESCRIPTIONS

OF VALID SPECIES OF IXODES

AND OF THEIR

VARIETIES AND SUBSPECIES

1. IXODES RICINUS1 (Linnaeus, 1746 and 1758).

Pis. IV and VII, Text-figs. 115, 139-148.

Synonymy: not Reduvius Charleton, 1668, p. 49.

? Ricinus caninus Ray, 1710, p. 10 (Synon. in Walckenaer and Gervais, 1844,

p. 236). ?.
not Acarus reduvius Linnaeus, 1746, p. 479. Syst. natur. 1758, 10th edit.,

p. 616; 1788, 13th edit., p. 2925.
Acarus ricinus Linnaeus, 1746, p. 480 ; 1758, p. 615 ; 1788, p. 2925.

Regarding 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 Reduvius Charleton {Onomasticon zooicum,
1668) and Pedicidus 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), retaining 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 Pediculus 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, vn, p. 98 ; p. 101 {reduvius).
Ixodes ricinus Latreille, 1804, I, p. 156 (9).
Ixodes reduvius Latreille, 1804, p. 51.

1 According to Dubreuilh, 1900, p. 456, the word Ricinus, used to signify a tick by
Pliny, gave its name to the castor-oil plant, as stated by Dioscoridea, iv, 161, because the
seeds of the latter resemble a tick {i.e. the gorged females).

144 Genus Ixodes

Cynorhaestes reduvius Hermann, 1804, p. 65.
Cynorhaestes ricinus Hermann, 1804, p. 66.
Ixodes megathyreus Leach, 1815, XI, p. 398.
Ixodes bipunctatus 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 trabeatus Audouin, 1832, xxv, p. 420 ( $ ).
Ixodes plumbeus Duges, 1834, i (2), p. 20.

not Ixodes plumbeus Duges, 1834 c, PI. VII, Figs. 7-12 ( = Rhipicephalus sp.).
Ixodes reduvius Hahn, 1834, n, p. 62.
Ixodes fiiscus Koch, 1835, Heft xxxvu.

Ixodes lacertae Koch, 1835, Heft xxxix ; 1844, p. 234, larvae and nymphs, fide
Canestrini, 1890, p. 497.

Ixodes rufus Koch f Koch' 1835> Heft v> No" H> Heft xxxvn- No- 8 '> 1844>
Ixodes mlcalus Koch J P- 232 i 1847' P- *li The ^^ examined bv Neumann
Ixodes sciuri Koch I (1901> PP- 281-289) proved to be I. ricinus nymphs, all

^ collected in Germany.
Ixodes reduvius Linn., in Koch, 1835, Heft v, No. 11.
Ixodes pustularum Lucas, 1866, vi ; Bull., p. lvii.
Ixodes fodieiis 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

ricinvs. — Donitz, 1905, p. 132.
Ixodes ovatus Neumann, 1899, as modified by Neumann, 1904, p. 4b2 = Ixodes

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, PL 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 5 in copula (recognized as
coitus), <£ dorsum, capitulum. Excellent, considering the date ; better than
some figures printed a hundred years later. — Audouin, 1832, PL XIV, Figs.
3a-n (trabeatus).— Duges, 1834, PL VII, Figs. 7-12 (plumbeus).— Hahn, 1834,

TICKS

PLATE IV

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 6. Fig. 7.

Figs. 1-7. Ixodes ricinus.
Fig. 1. i dorsum, x 12. (Wheler, 1906, Fig. 16.)
Fig. 2. i 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 t coupled and a second t "waiting his turn." (Wheler, 1906, Fig. 36.)
Reprinted (Fig. 2 excepted) from the original blocks illustrating the paper by
E. Q. Wheler, Journ. Agric. (Cambridge), Vol. i.

/. ricinus 145

II, PL LXVI, Fig. 152 ((J).— Koch, 1835-44, Heft 5, Fig. 11 {reduvius ?);
Heft 37, Figs. 5-7 (ricinus <J, 9 s), Fig. 8 (sciuri) ; Heft 39, Fig. 11
(lacertae), Figs. 3, 4 (fuscus <J, 9), Fig. 7 (>m/w, 9 ).— Pagenstecher, 1861, II,
PL I, Figs. 1-10 ; PL II, Figs. 1-22. Really excellent figures of anatomy.—
Megnin, 1867, I, PL IV.— Murray, 1877, Fig. 2 (fodiens; figure useless).—
Haller, 1882, PL V, Fig. 4 (ricinus).— Braun, 1883, Figs. 55, 56 (ricinus).—
Railliet, 1886, p. 348 (ricinus).— Aurivillius, 1886, Figs, a and 6 of 9 ; very
poor. — Berlese, 1888, fasc. xlix, PL 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, PL XLIII, Figs. 1-1 g
(I. reduvius) : <$ venter, hypostome ; 9 dorsum (fed and unfed), part of
capitulum, digit, etc. (very poor). Figs. 3-3 6 " Phaulixodes ru/us (Koch)"
dorsum, digit, hypostome ; scarcely recognizable. — Blanchard, 1890, Figs. 592-
595 (ricinus). — 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 ; reproduced by Salmon and Stiles, 1901, Fig. 222, and by other authors ;
<J venter, capitulum, tarsus 4 and foot. Somewhat schematic (reduvius). —
Gene, in Railliet, 1895, Fig. 480 ( $ ovipositing, original figure by Gen£). —
Braun, 1895, Figs. 127, 128 (ricinus). — Megnin, 1895, figs, worthless, of different
stages.— Osbom, 1896, Fig. 155; poor.— Niles1, 1898, PL IV, Figs. 3-6 (ricinus).
—Morgan1, 1899, Pis. VII, VIII, Figs. 1-5 (ricinus).— 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, £ , J, 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, $ (gorged),
much enlarged, dorsum and venter ; PL XCIV, Figs. 225, 226, 9 capitulum in
ventral aspect, capitulum and scutum, much enlarged (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, PL I, Figs. 2, 2 a, 2 6 (9 sketchy, coloured).— Nordenskiold, 1905, text-
figures, illustrating anatomy and histology. — Wheler, 1906, Pis. VII, VIII,
reproductions of photographs of (a) mounted specimens ( $ and $ ), poor ; and
(6) of unmounted ( <$ , 9 dorsum and venter, o and larva ; here reproduced) ;
PL X, Fig. 38, shows an abnormality : a 9 minus capitulum ; Fig. 36 ( £ and
9 in copula) ; other figures illustrate gorged 9 1 and 9 ovipositing. — Donitz,
19Q7, PL VI, Figs. 34-36, 38, <$ and 9 capitula, $ venter ; good.— Banks,

1908, PL II, Figs. 9, 10, 13 ; PL IX, Fig. 5, brief description of 9 only ; figures
relate to 9 and are poor.— Bonnet, 1908, p. 254, Figs. 15, 16 ; poor. —
Nordenskiold, 1908, Pis. XXVI-XXVIII, Text-tigs, 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, PL IX,

1 Inaccessible, cited by Salmon and Stiles, 1901.

146 Genus Ixodes

7 text-figs, (spermatogenesis). — Braun and Liihe, 1910, Figs. 89, 90. Original,
(J venter and $ capitulum in ventral aspect. Good representations after
drawings by A. Dampf.
Literature : 1710. Ray, p. 10 (Ricinus caninus). — 1746. Linnaeus, p. 480 (Acarus
ricinus). — 1758. Linnaeus, p. 615 (Acarus ricinus, not Ac. reduvius ; latter
confused with Melophagus ovinus). — 1778. de Geer, pp. 98, 101 (Acarus
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)1. — 1805. Fabricius, p. 351 (/. ricinus). —
1807. Chabrier, p. 366 ; Illiger 2, p. 370.— 1815. Leach, p. 398 (/. megathyreus
on hedgehogs and dogs) ; p. 397 (/. ricinus on dogs). — 1817. Mtiller2, p. 278
(/. ricinus). — 1826. von Heyden, p. 610 ; Risso, p. 183 (/. bipunctatus,
Cyn. hermanni, Cyn. megathyreus (Leach)). — 1829. Dumeril, p. 401 (Crotonus
ricinus) ; von Heyden, p. 288. — 1831. Treviranus, p. 191. — 1832. Audouin,
p. 420 (/. trabeatus).— 1834. Hahn, p. 62, O3; Duges, p. 20 (/. plumbeus).—
1835-44. Koch, Heft 37, No. 5 (<J) ; H. 5, No. 11 ; H. 37, No. 8 (/. sciuri);
No. 3 (/. fuscus on deer); No. 7 (/. rufus, $, on deer); No. 11 (/. lacertae,
o on Lacerta agilis, Munich). — 1837. Burmcister, p. 579 (/. marginatum). —
1844. Walckenaer and Gervais, p. 236 ; Gend, pp. 1 et sea., 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 (/.
ricinus) ; 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. Kiichenmeister, p. 422 (mere cited description
of $ /. ricinus).— 1857. Kolenati, p. 24.— 1858. Milner2, 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. Gerstaecker2, p. 344.— 1864. Cox, p. 82 (on dogs
and ferrets, but may have been hexagonus). — 1866. Lucas, p. lvii (/.
pustularum).— 1867. Megnin, p. 107.-1869. Packard2, p. 613.— 1870. Verrill2,
p. 118.— 1876. Wagner, p. 129, O.— 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. Conil2, p. 100.— 1879. Cobbold, pp. 267, 350.— 1880. Haller, p. 38 ;
Megnin, p. 127 ; Oudemans, p. xvii ; Taschenberg, p. 151, O. — 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, ? ricinus, penetrating
beneath the skin in man ; thinks ticks do so normally ! — 1885. Johannessen,
p. 401 (effects of bite on man).— 1886. Railliet, p. 497. Aurivillius, (a), p. 105,
3 figures of $ (poor) ; (b), 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 O denotes that the publication referred to contains nothing worth noting.

/. ricinus 147

beneath human skin, excision of a $ ricinus) ; Batelli, pp. 78 et seq. (physiology
of digestion). — 1892. Neumann, p. 95 ; Megnin, p. 32 ; Efisio1, p. 256 ; Janson
and Tokishige1, p. 349.— 1894. Mosler and Peiper, p. 320 ; Dolly1, p. 1000.—
1895. Braun, p. 257 ; Neumann, 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; Schneidemuhl, p. 339; Dubreuilh and
Beille, p. 69, O.— 1897. Wood and Fitz1, p. 346.— 1898. Niles1, pp. 29, 45 ;
Weichselbaum, p. 325. — 1899. Morgan, p. 129, O ; Neumann, p. 116 (/. ovatus
in part), p. 112 (/. reduvius) ; Wheler, pp. 38, 49 (biology and structure) ;
Mosso1, p. 20; Nuttall, 1899, p. 42, cites Dubreuilh, 1838, Despres, 1867,
and others re effects of bite. — 1900. Kossel and Weber, relation to redwater
in Finland. Ward, {a), p. 203 ; (b), 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, pp. 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., O.
Mohler, 1905, p. 14, O.— 1906. Wheler, p. 411 ; Vassal, p. 285, also Martin,
p. 105 (experiments with trypanosome infection : negative) ; Braun, p. 364. —
1907. DSnitz, p. 90 ; Pocock, p. 201 (cites Wheler, 1902).— 1908. Nuttall
(figures illustrating a lecture, reproduced here) ; Banks, p. 24, O ; Bonnet,
p. 254, O; Galli-Valerio, p. 611, biology; Nordenskiold, p. 637, histology,
important ; Stockman (reprint, relation to redwater). — 1909. Samson, (a),
pp. 185-236 ; (b), 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 Ltihe, p. 178, O ; Samson, pp. 1 et seq., dissertation, anatomy and biology.

Male (PI. IV, Text-figs. 115, 139-142) : average L. 245, W. 1-33 mm.2.
Body dark red-brown, narrow in front, broadly rounded behind, very

1 Cited in bibliography by Salmon and Stiles, 1901, unimportant.

2 Note regarding measurements : The size of different specimens may vary, thus 6 S s,
comprised in two lots collected in England (N. 933, 934), measured in mm. —

Scutum

Length, including Body ,. * s

capitulum width length width

f 2-6 13 1-85 1-1

Lot 933 \ 2-5 1-4 20 1-2

(2-3 1-35 1-75 1-1

Lot 934

2-5 1-4 1-9 1-1

2-4 1-3 1-8 10

22 1-2 1-6 0-95

Wheler (xn. 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 ? at

148

Genus Ixodes

convex when gorged. Marginal fold slight, generally of lighter colour
than the scutum. Scutum (average 18 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. Emargination 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, anal plate with lateral borders divergent, adanal plates
narrowing posteriorly. Spiracles large, oval, with long axis directed

Fig. 139. I. ricinus & : 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 6-5 mm. respectively. Nymphs, unfed, measure l-5 x 1-68 mm.,
when gorged they attain 3-3 mm. in length (Wheler, 1899, p. 49).

/. ricinus

149

forward and macula anterior. Gapitulum 500 fi long ; base trapezoid,
broader in front, slightly convex behind, without comua. 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 s ■ 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 cf and ? respectively. (Neumann, 1899, Figs. 2, 3,
illustrating the structures in 2\ ovatus. We find them identical in I. ricinus.)

150

Genus 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 (VS x 1*15 or 1*4 x l-05 mm.) dark brown, sub-pentagonal,
with rounded angles ; emargination 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. ricinus ? : capitulum in dorsal and ventral aspects, x 70. Specimen from
Coilessan, Scotland. (N. 933. Original, G. H. F. N. del.)

I. ricinus

151

Venter: vulva between coxae IV; spiracles circular, genital grooves
only slightly divergent, anal grooves rounded in front, parallel behind
anus. Capiiulum long (800 ft). 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 dorsally, 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. 145, 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. ricinus o : 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.1
400, 401, 417), j/Vs $s, from dog, cattle and sheep (R. Bramwell) ; from
Shelfhill Farm, Hawick, v. 1908 (N. 405), /a, ?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), </" from sheep, v. 1910 (Captain Leschallus); from
Thornhill, 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, Baukfort, VIII. 1904; from deer, near Crawford,
X. 1908 ; from wild cat, no locality 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 grouse1. [Mr W. Evans (i. 1907, p. 35) records this tick
on cows, near Callander, mostly attached to udders and adjacent parts
of legs (sexes in copula, 21 September) ; o s found on head of red-deer

Fig. 147. /. 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.).

1 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); J near Killin (Det. by Neumann)]. England and Wales:
From Northumberland (N. 1039) Js, $s, os, on sheep, v. 1905
(J. Hedley); from Huntingdonshire (N. 1037) ?s, on cat, in. 1906
(E. C. Burleigh); from Grantchester, Cambs. (N. 123) o, 1906 (N. D. F.
Pearce); from Longner Hall, Shrewsbury (N. 1036) ?,on young rat,
iv. 1906 (R. F. L. Burton); from Lyndhurst, Hants (N. 656), larva,
on Mustela erminea, vn. 1900 (G. Tate ; ex N. C. Rothschild coll.) ;
from Tavistock, Devonshire (N. 963) £ , %$,, 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 Gowanburu
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, vn. 1905 (J. W. Steen); (N. 1038) °.s, on cattle affected
with piroplasmosis, v. 1905 (A. E. Mettam); (N. 1185) </", 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, vm. 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 ; Crookbaven, 1895 ; Roundstone, Ardagh, Castletown, vi.
1893; Kenmore, vn. 1898; Lough Corrib, 1905. Prance: [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 fiedgehog and deer, Chantilly, ix. and XI. 1909 ; and from
the roe-deer, Halles (Paris), x. 1908. Belgium: (N. 631) ? from
Erinaceus europaeus (E. A. Boulenger, ex N. C. Rothschild coll.).
Holland : [larvae found on Mus decumanus (Oudemans)]. Germany :
From Kittendorf and Schwandt, Mecklenburg-Schwerin, on various
hosts, as follows : (N. 57) f, on man, IX. 1905 ; (N. 769, 770) ^s, ? s,
on Cervus elaphus, vm. 1909; (N. 771) £ , %, o, on Capreolus capraeu,

/. riciniis 155

vm. 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, vn. 1910, and (N. 1184)
? , O, from a dog, 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,
Schiitz, 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 Regulus ignicapillus and on Strix brachyotus, 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 Rhinolophus 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 avellanarius (Bureau of Animal Industry coll.,
Washington, D.C.)J. Albania and Rhodes: [Neumann, 1901, p. 282,
states that </s, $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 also records, 1899, p. 116, a o
from Caucasia in the Megnin 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) J1, ?, on dog, Azazea, in. 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, n. 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, vm. 1903,
collected by Dr E. Dschunkowsky, 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 Felts concolor, xil. 1894
(Gilroy), and (N. 291) on mouse, Palo Alto, in. 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 /. ricinus). 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 wild 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 afiinis 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.

I. ricinus var. scapularis

157

Ixodes scapularis Say, in Banks, 1904, p. 144 (occurrence in Florida); 1908,
pp. 25, 26, PL IX, Figs. 1, 2 ; PL II, Fig. 15 ( $ unfed, inaccurate ; sexes
in copula ( <J attached too far posteriorly), $ venter and capitulum, details
fairly accurate (Fig. 15 very sketchy). Hooker, 1908 a, observed 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 sub-parallel 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 aflinis was found by
C. Curtice on Felis pardalis in Costa Rica.

Fig. 149. I. ricinus var. scapularis 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 Genus Ixodes

We possess specimens (N. 626), </\ $, 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 described contained typical ricinus. The figs, of the <J

and $ hypostomes and digits as in ricinus.
Ixodes ovatus Neumann, 1904, p. 452, species maintained (description quoted

below).
Ixodes ovatus Neumann, in Donitz, 1905, p. 132. Donitz considers the species

probably but a variety of ricinus.

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 Yamagnchi. The differences noted only appear to ns to be varietal.

nigricans

159

Ixodes ricinus var. californicus (Banks, 1904).

Lit. etc. : Ixodes californicus Banks, 1904, p. 369, PI. XLI, Fig. 57 (scutum and
coxae)1; 1908, p. 24, PI. II, Fig. 12 ($ capitulum and scutum).

We have not seen this tick, which appears, from its description, to
be specifically indistinguishable from /. ricinus. Banks states that
the capitulum 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 : Colour : Scutum yellowish-brown, legs darker brown, abdomen
yellowish-grey. Scutum l-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, vn, 1908, p. 75. Name given for the reason that
obscurus was preoccupied (/. obscurus 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
posteroexternal border, but no (sic) basal spine ; coxae II and III with prominent spine, and
coxa IV with small protuberance at the postero-extemal border. Found on Toxostoma
r.rissalis, at Claremont, California (Baker). This description does not agree with the later
one.

160

Genus Ixodes

Female (Fig. 150) : Body short oval, 7x5 mm. Dark brown all over. Scutum
oval, borders rounded, l-3 x l'l 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. Capitulum 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. /. nigricans $ : capitulum and scutum ; parts of venter, spiracle and tarsus 4.
Drawn from the type. (Hamburg Mus. 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 apparently
differs from /. ricintis only in (a) the scutum, which is oval (without
lateral angularity) and faintly grooved, and (b) 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 n long ; dorsal process longitudinally crescentic ; external article with five
successive teeth, the last stout.

T. acuminatus

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 I. 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. 151 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 (1'3 x 0"9 mm.), lozenge-
shaped oval, antero-lateral borders subrectilinear, divergent, the portion
bounded by the posterior rounded border forming f 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. acuminatus ? : venter. (Neumann, 1901, Fig. 7.)

Fig. 152. I. acuminatum ? : H, hypostome, C, digit, 120 /j. 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 | 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 ( <? 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. 35 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
■ — -ofone-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 4. Palps with article 2 slightly longer than 3. Legs short,
maroon-brown. Coxae (Fig. 154) close together, not occupying the

7. 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.). \ tf#/t

16. &A

■<t us**.

Fig. 153.

Fig. 154.

Figs. 153, 154. I. dentatus ? : capitulum 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. 136, 137 (no figure).

Banks, 1908, p. 27, PI. Ill, Figs. 12, 14 (? capitulum, scutum, spiracle, coxae 1
and II, tarsus 1); not reproduced, as we do not trust their accuracy.

Male : unknown.

Female : Body oval, reddish yellow, 5 x 25 mm. [? partly gorged].
Scutum oval, 1'3 x l'l 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. Venter : 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. Capitulum 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 j 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— I V. Tarsi long, tapering gradually.
Description, by Neumann, based on 2 $s (mutilated) collected by
Hassall from Procyon lotor, 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. T45, W. OS 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 i : venter. (Neumann, 1902, Fig. 1.)

Fig. 156. I. minor J : venter (L. 1'73 mm. ), from the type mounted in balsam.
(Neumann coll. Original, G. H. F. N. del.)

I. minor

165

hairs fairly abundant over the whole ventral surface. Spiracles large,
slightly oval. Capitulum (Fig. 157) long (0'45 mm.), 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. /. minor <? : base of capitulum, showing cornua. Drawn from the same
specimen as Fig. 156. (Scale = 025 mm. Original, G. H. F. N. del.)

Fig. 158. I. minor <? : hypostome, drawn from the same specimen as Fig. 156. (Scaler
0-2 mm. Original, G. H. F. N. del. )

Female (gorged, Fig. 159) : Body elongated oval, L. 5, W. 2*5 mm.,
broadest at the posterior third, yellowish brown. Scutum oval, much
longer than broad (l*3l 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. Capitulum long (0"8 mm.) ; dorsal base subtrian-
gular, with very prominent cornua ; porose areas large, circular, separated
by a space equal to their diameter; auricula forming a prominent
horn. (Digit1.) Hypostome (absent^ but probably sharply lanceolate as
in /. acuminatum). 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 {/1,
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. l-6, W. 1"1 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 /i long ; dorsal process crescentic, longitudinal and with retrograde
points, external article with five teeth, larger and more separated proceeding toward the
base (as in J. acuminatum).

I. boliviensis

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. Capitulum 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. /. boliviensis i (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. 19 mm.
[partly fed], yellowish brown, the capitulum, scutum and legs maroon-
brown. Scutum oval, sides convex, longer than broad (To x 1*2 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 cornua ;

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. 1. boliviemii ? (type): capitulum and scutum; part of venter; spiracle.
(Original, G. H.F.N, del.)

Fig. 162. /. bolivierwis o (type, unfed) : dorsum, part of venter, tarsus i. Same
origin as <f and ? in Figs. 160, 161. (Original, G. H. F. N. del.)

I. fuscipes

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 £, 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. : Ixodes 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. I. fuscipes ? : capitulum and scutum ; capitulum in ventral aspect, coxae,

spiracle and tarsus 4. Found on Danyprocta aguti. (Neumann coll. 760. Original,

G. H.F.N, del.)
Fig. 164. I. fuscipes ? : capitulum (mutilated) and scutum, coxae and part of capitulum,

showing auricula. (Sketch from specimen in British Mus. Original, C. 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 margins 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 (G"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.1 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 1 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 07 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 of 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.

/. sculptus and spinicoxalis 171

9. IXODES SCULPTUS Neumann, 1904.

Lit.: Neumann, 1904, p. 462 (no figure).

Banks, 1908, p. 30, PL IV, Figs. 7 and 9 ( ? capitulum, scutum, coxae) ; not
reproduced.

Male : unknown.

Female : Body oval, narrower anteriorly, brownish, L. 2, W. 1*8 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 punctations, 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 /m), 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
tapering abruptly.

Description (from Neumann) from a specimen found with a ?
J. 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.,
glossy, 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

Genus Ixodes

a line with the posterior border of coxae IV. Sexual grooves uniting
in front of the vulva, straight 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. (Digit1.) Hypostome (Fig. 166)
narrow, lanceolate, long, 4 1 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. O. Forbes (British Museum).

Fig. 165. Fig- 166.

Fig. 165. J. spinicoxalis J : capitulum (mutilated) and scutum, coxae and part of basis

capituli. (Sketch from the type in the British Mus. Original, C. W. del.)
Fig. 166. /. spinicoxalis ? : hypostome x 115. (Neumann, 1899, Fig. 6.)

1 Digit 160 n 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.

I, niarxi

173

11. IXODES MARXI Banks, 1908.

Fig. 167 (original).

Lit. etc. : Banks, 1908, p. 32, PI. IX, Fig. 8 ( ? dorsum, G. Marx del.), PI. Ill,
Fig. 10 ($ capitulum, scutum and coxae; Banks del).

Male : unknown.

Female (Fig. 167): Scutum (1-2x0-9 mm.), longer than broad,
broadest rather in front of the middle (but not so anteriorly as in
/. 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. Capitulum : 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 j 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.

^jrV

Fig. 167. I. marxi s : capitulum and scutum, ventral aspect of capitulum with coxae,
tarsus 4, spiracle, genital and anal grooves. From specimen kindly loaned by Di-
li. 0. Howard, Government Entomologist, Washington, D.C., U.S.A. (Original,
G.H. F.N. del.)

174

Genus Ixodes

Described from a % 1 found on 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 (=1. canisuga). It is somewhat allied to /. canisuga,
but can easily be distinguished from it by the different 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) " L. 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.)

1 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 marxi Banks.

/. pratti and rubidus 175

two humps behind, and III with one hump; venter punctate ; stigmal
plate twice as long as broad, finely granulate."

Female (Fig. 168): Scutum 12 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 beween 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 auriculae1. Palps rather short and
broad, article 1 strongly prominent ventrally. Hypostome strong, well
covered with 2 [ 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 J1 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 14 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 s in our key, Banks' description being insufficient.

176

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. Capitulum:
(08 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. Pig. 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. J. 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 (06 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.).

/. hexagonus 177

14. IXODES HEXAGONUS Leach, 1815.

Figs. 171-177 (original).

Synonymy: 1. Ixodes hexagonus Leach, 1815, xi, p. 397, on hedgehog.

2. Ixodes autumnalis Leach, 1815, xi, p. 398, on pointer dog1.

3. Ixodes erinacei Audouin, 1832, xxv, p. 415 (?)1,2.

4. Ixodes reduvius Audouin, 1832, xxv, p. 422; nee Linnd {$)1.

5. Ixodes crenulatus Koch, 1835-44, Heft 39 L

6. Ixodes vulpis Pagenstecher, 1861, II, p. 40 1>2.

7. Ixodes erinaceus Murray, 1877, p. 190 '.

8. Ixodes ricinus 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. (I. canisuga Johnston,

1849 ; I. cookei Packard, 1869 ; /. cruciarius Fitch, 1872, are listed by
Blanchard, 1909, p. 86, as synonyms ; see our List of condemned species.
We recognize canisuga.)

Iconography : Audouin, 1832, xxv, PI. XIV, Figs. 2, 4.— Koch, 1835-44, Heft xxxix,
Figs. 5, 6 ( $ /. sexpunctatus) ; Figs. 8, 9 ( (J § /. crenulatus). — Pagenstecher,
1861, ii, PL I, Figs. 12, 13.— Berlese, 1889, fasc. 55, No. 4 (I. crenulatus
v. Synon.) ; 1892, fasc. 61, No. 10 ; $ dorsum, venter, capitulum and scutum,
spiracle and tarsus 4.— Canestrini, 1890, iv, PL XLIII, 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,. 1893, 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 (v. 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, PL II, Figs. 4, 4a, 4 6 (sketchy, coloured). — Wheler,
1906, PL VIII, Fig. 22, $, rough sketch after Neumann, 1899, Fig. 10;
PL VIII, Fig. 20: $ dorsum, good photograph of opaque object (reproduced). —
Banks, 1908, PL III, 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 /. hexagonus is
undoubtedly described, and this is rendered certain by reference to the Icono-

1 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 in 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
/. 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. 234; 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. Railliot, 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.

Male1 (Fig. 171): L. 35 to 4 mm. (capitulum inch), W. 2 to
25 mm. (Nn.). Body oval, nearly as broad in front as behind. Scutum
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 s : 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 ForSt
de la Londe, Seine Interieure. We have recently (7. m. 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.

I. hexagonus

179

almost a regular hexagon, preceded by two small plaques behind
coxae I; median plate very broad; anal plate ogival ; adaual 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 (12 x 1*4, or 15 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. . /. lie.r<itjtmu» ? : dorsum, venter, spiracle and tarsus 4. Specimen off
MmteUa rultjaris, Tring j F. J. Cox coll., 1908. (N. 668. 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 Myopotamus coypu, S. America.
(N. 826. Original, F. M. H. del.)

Fig. 174. J. hexagonus ? \ capitulum in dorsal, ventral and lateral aspects. (Original,

G. H. F. N. del.)

Fig. 175. I. hexagonut ? : tarsi 1 and 4, highly magnified, from mounted specimen.
(British origin. Original, G. H. F. N. del.)

/. hexagomis

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 | 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 j 2, with fewer teeth per file.

Fig. 176. I. hexagonm 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.)

TFSv

Fig. 177. I. hexagomis 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 Genus 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, Os and larvae from the following places
and hosts [observations by other authors are added in square brackets] :

EUROPE : Scotland : Mr W. Evans' collection iu Edinburgh con-
tains os from the polecat, Ross-shire, xn. 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 europaeus,
both in Ross-shire (Neumann det.).] England: (N. 404) °.s, os, from
Mustela putorius, Holme, Peterborough, in. 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, vn. 1908 (W. A. Harding); ?,
Grantchester, near Cambridge, vm. 1906 (N. D. F. Pearce) ; o s,
Longnor Hall, Leebotwood, Salop, ix. 1910 (R. F. L. Burton).
(N. 1160) Larvae from young Canis vulpes, Boxworth, Cambs., in. 1900
(N. C. Rothschild). (N. 262) $ , from 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, Hauts., 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, in. 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 woman, Caxton, Cambs., vi. 1905 (Dr Giles, presented by
Dr Wherry); (N. 604) Mustela vulgaris, Tring, Herts., in. 1902
(N. C. Rothschild); (N. 668) ?s, os, 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 putorius,

I. hexagonus var. cookei 183

Aberystwyth: (N. 746) larvae, vm. 1900; (N. 606) ?s, os, larvae,
xi. 1902; (N. 609) os, xn. 1902. Ireland: (N. 612) ?s, os, from
Mustela erminea, Inismore, in. 1902 (N. C. Rothschild). Belgium:
(N. 597) from Erinaceus europaeus (E. A. Boulenger). Germany :
(N. 1168) ?s, os, from ferrets, Kittendorf, Mecklenburg-Schwerin,
vi. 1910 (G. v. Oertzen's gamekeeper) ; (N. 608) from Erinaceus
europaeus, Dernburg, ix. 1898 (N. C. Rothschild); (N. 1186b) %,
from dog, Godesberg, vn. 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 <$ 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 " crenulatus " 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 ou Myopotamus coypu (ex Rothschild coll., no particulars).

Ixodes hexagonus var. cookei (Packard, 1869).

Syn. : Ixodes cookei Packard, 1869 a, p. 67.

Ixodes hexagonus var. lonyispinosus Neumann, 1901, \i. 283.
Ixodes cookei Packard, in Banks, 1908, pp. 28, 29, PI. IX, Figs. 1-5. (Larva,
dorsum ; palp of o (surely wrong !) ; <J 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 capitulum, scutum and spiracle.
Ixodes hexagonus 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 Genus Ixodes

Neumann examined 43 Js and 127 ?s from various mammals in
different 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 inchoatus 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 J. cookei with Neumann's types of /.
hexagonus var. longispinosus, and finds them identical. Dr L. O. 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, correbt 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 Js and 113 ? s, found on Lutra, Mustela
vison, sheep, Texas; Spermophilus; Felis domestica, Maine; fox, 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).

I. 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. 25 to 3"5 mm., W. 15 to 19 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, 750 fj, 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 1 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, thosf
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 macleari 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

± rfwtM^WK**

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, 4 x 21 mm. (partly fed), broadest
toward the posterior third. Scutum oval, 1*7 x To 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 1 4 followed by 3 1 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

^ -«a\

St

i jtt ^

• ^j

$W v dk*

(A '

Fig. 179. I. bicornis ? : part of venter ; hypostome. (Neumann, 1906, Fig. 1.)

/. auritulus 187

coxa II ; the external spur pointed, slightly longer than broad, almost
parallel to the internal ; a short spur at the postero-external 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 " Conchuda " at Atoyac, is considered to inflict " a bite fatal to
children "(A. Duges coll., now in Neumann coll., Toulouse). /. bicomis
is very closely allied to /. fuscipes Koch (/. spinosus Neumann). We
possess (N. 637) 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 /. bicomis Neumann as
synonymous with I. diversifossus Neumann, 1899. In the absence of
a figure of I. diversifossus, we would point out that the descriptions
differ as follows :

I. diversifossus. I. bicomis.

Scutum: 1*3 x 1 mm. 17 x To mm.

Palps: Article 2 scarcely longer Article 2 one and one-half times as

than 3. l°ng as 3.

Hypostome : 2 | 2 or 3 | 3 4 | 4, followed by 3 1 3.

<-^ O*.

C ecJi

17. IXODES AURITULUS Neumann, 1904.

Figs. 180 and 181.

Lit., Icon, and Synon. : Ixodes thoracivus Neumann, 1899 ; not Ixodes

t/wracieiis C. L. Koch, 1844, as described in Neumann, 1899, p. 149.
Ixodes thoracicus Neumann, 1899, pp. 149-151, Figs. 22, 24 (reproduced).
Ixodes auritulus Neumann, 1904, p. 450.

Male : unknown.

Female : Body short oval, swollen, 7 x 5'5 mm., dark reddish brown,
legs reddish yellow. Scutum oval, with lateral borders almost straight
along their anterior half, 13 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. Gapit-
ulum 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. nuritulus ? : capitulum and Bcutum ; V ventral and D dorsal aspects.

(Neumann, 1899, Fig. 22.)
Fig. 181. /. auritulus ? : hypostome, x 65. (Neumann, 1899, Fig. 23.)

/. brumieus 189

Description based on 4 Js 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 Neumaun's figure of
the type, and correspond more to our figure of /. brunneus (Fig. 182).

18. IXODES BRUNNEUS Koch, 1844.

Figs. 182, 183.

Lit. etc. : Ixodes brunneus C. L. Koch, 1844, p. 232 ; 1847, p. 99, PI. 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 aoisugus Berlese, in Oanestrini, 1890, p. 505.
not Ixodes bifurcatxis Neumann, 1899, p. 122, as stated by Neumann, 1901, p. 347,

in relation to a $? specimen he subsequently 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).
V Acarus frontalis Panzer, 1795, fasc. 59, Fig. 23.
? Ixodes pari Leach, 1815, p. 399, found on Parus major.
1 1xodes pall ipes Koch, 1835-1844, Heft 39, Fig. 10, found on Sitta enropaea

(Larva).
? 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" 1 '. 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) : L. 7 to 8, W. 5#2 mm. Body brown, with
numerous white hairs. Scutum (Fig. 182) very long, 1*4 x 11 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. Capitulum (Figs. 182, 183) of medium length (08 mm.) ; base
triangular, comua 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 j 2 l. 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 /. 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
albicollis Gmelin, from North America. The larva described by Koch
(1844) as /. pallipes was found on Sitta coesia by Panzer (1795), and
Leach (1815) 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 arboretis, A. pratensis, Erythacus rubicula, Merula
nigra. Canestrini (1890) possessed two specimens found on Praticola
rubetra and Emberiza schoeniclus.

The description of frontalis by 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 rubicola

1 In Neumann's figure (Fig. 183 of frontalis, here reproduced) the dentition ranges
from 4 1 4 to 3 j 3, 2 | 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.

/. brunneus

191

(Paris Mus.), Turdus aonalaschkae Pallasi, from Baltimore (Bureau of
Animal Industry) ; 1 o from a goose in Paris (Trouessart coll.) ; 1 larva
from Turdus merula from Saintes.

Fig. 182. J. brunneus ? : A, capitulum and scutum, x 20 ; B, 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. brunneus ?: A, digit, x220; II, hypostome, x80; C, ventral aspect of
basis capituli and coxae I, x40. (Neumann, 1899, Fig. 11, of J. frontalis (Panzer).
See footnote to p. 190. )

192 Genus Ixodes

We possess, or have examined, females from France : (N. 523) from
Passer montanus, St Gdnis de Malgoire (Gard), xi. 1908 (A. Hugues
coll.; presented by Hon. N. C. Rothschild); England: (N. 224) from owl,
Cambridge, 1904 (G. S. Graham-Smith) ; California : (N. 2791) from
Cyancitta stelleri frontalis, XII. 1895 (R. C. Snodgrass) ; (N. 2801) from
thrush, Palo Alto, XI. 1895 ; (N. 282) from Piranga ludoviciana Sunol,
V. 1896 ; (N. 283) from Habia melanocephala Sunol, v. 1896 ; from " a
bird1," 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 (Haplopelia 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 (16 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 bearing a few white hairs. Spiracles whitish, in front of half
the body-length. Capitulum 1 mm. long. (Digit 145 ft long.) The rest
as in the type.

19. IXODES CAVIPALPUS Nutt. and Warb. 1907.

Figs. 184-180.
Lit. and Icon. : Nuttoll and Warburton, 1907, p. 396, Figs. 1-5 (reproduced).
1 Described by us as /. kellogyi.

/. cavipalpKx

193

Male (Figs. 184, 185) : L. 19, W. 12 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 grooves1 and punctations very faint. Venter
brown with long hairs ; sexual 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 A, 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 i : 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 ;t and 270 /t£.
orientated so that the tick's capitulum was to the left and the dorsum above ; the ?
spiracle shows a slight angularity postero-dorsally. (B) i and ? digits, respectively,
138 ix and 140 /x. I., viewed from the dorsal surface. (Nuttall and Warburton, 1908,
Figs. 4 and 5.)

1 Accentuated in Fig. 184.

194

Genus Ixodes

Female (Figs. 185, 186), more or less fed : L. 29 to 5, W. 18 to
33 mm. Body brown, with a few scattered hairs. Scutum (I. 13 to 1*4,
w. 1 to 1*8 mm.) dark brown, long oval with slightly sinuous posterior
border ; cervical grooves shallow, almost reaching the margin ; no lateral
grooves ; numerous fine shallow punctations. Venter 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 | 2. (Digit ; see Fig. 185 B.) Legs as in J, 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. I. cavipalpus ? : capitulum in (A) dorsal and (B) ventral aspects. (Nuttall
and Warburton, 1908, Fig. 3.)

Description based on (N. 245) 2 fs and 7 ? s from a baboon (Cyno-
cephalus 6a6wTO),!Kansanshi, N .W. Rhodesia, 1. 1907 (Dr A. Yale Massey) ;
(N. 353) 1 £ from a native baby, Benguella Hinterland, Angola, XII.
1907 (Dr F. Creighton Wellman) ; and (N. 394) ^s, $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.

/. angustus

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 ? ). In this paper
Neumann re-described what he took to be angustus from 2 $ s found on
Didelpkys azarai, Buenos Ayres (C. Berg coll.). Banks (1908, v. infra) has
pointed out that 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 angustus,
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
( 9 capitulum, scutum, coxae and spiracle ; sketchy, but not very inaccurate).

Male1 (Fig. 187) : oval, rather narrower in front, broadest in the
middle. Scutum (l-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
border somewhat pointed. Capitulum : base trapezoid, broader in

Fig. 187. J. angustus i : 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.)

1 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 [ 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, tarsus 4 tapering obliquely ; pad long.

Female (Fig. 188) : Body elongate, with numerous fine punctations
and short white hairs. Scutum (15 x l'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. Venter: 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. angustus ? : 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.) %

T. angusttis

197

Nymph1 (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. Capitulum :
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.

Larva1 (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. I. angustws 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. I. anijustus 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 J1 (N. 1065) from Sciurus hudsonius 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, xn. 1909 and ix. 1910, N. 506, 991, etc.).
Mr S. Hadwen has sent us (N. 1195) a </ and ? which copulated in
confinement, the £ 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 angttstus on the
strength of their structure.

198 Genus Ixodes

squirrel aforementioned, vin. 1910. We have also received a %
(N. 678) from Tamias townsendi, Chilliwack, British Columbia
(ill. 1900, N. C. Rothschild coll.), a ? (N. 284) from Sciurus sp., La Honda,
California (n. 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, xn. 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*8 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. Capitidum : 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. Venter: 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

I, caledonici is

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.

\^ZT^

Fig. 191. I. caledonicus ? : capitulum and scutum, capitulum in ventral aspect with
coxae, anal groove, tarsus 4 and spiracle. (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 0-7 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 hies. Venter : spiracle bluntly oval. Otherwise resembling
the ?.

200

Genus Ixodes

Fig. 192. /. caledonicut o : capitulum 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 08 mm. long, resembles the °.
and o in its chief characters (coxae, trochanters, tarsi). Scutum more
rounded (0*4 x 04 mm.) than in the o, with deep cervical grooves.
Capitulum : hypostome 2 j 2, with six teeth in the external files.

v%

<5

Fig. 193. I. caledonicus larva: capitulum and scutum, ventral aspect of capitulum with
coxae, tarsus 3, anal groove. (Nuttall, 1910, Fig. 3, F. M. H. del.)

Described from 1 !j! (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. vm. 1910, communicated
by Mr William Evans (Edinburgh).

/. fossalatxis 201

22. IXODES FOSSULATUS Neumann, 1899.

Fig. 194 (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 sbagreened 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
conical 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 ? : capitulum (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 Mus.), 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. : Haemalastor acutitarsus Karsch, 1880, p. 142 (Latin

description in five lines ; quite insufficient).
Ixodes laevis Neumann, 1899, p. 148, Fig. 21 (reproduced); 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 (25 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. I. acutitarsus J : (1) dorsum; (2) digit, 265/i'-> and (3) coxae. (Neumann,
1899, Fig. 21 of I. laevis Nn.)

/. gigas 203

in front, widely divergent behind ; short hairs, mostly behind the coxae.
Spiracles large, subcircular, with macula eccentric (anterior). Gapitulum
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 long or
slightly longer than the palps; dentition 4 | 4, then 2|2 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.
Gapitulum 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 oi'ifice 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 & : dorsum and venter. (C. Warburton, 1910, Figs. 1 and 2.)
The anal grooves are somewhat too divergent in the figure.

Female : unknown.

Described from two specimens in the India Museum, Calcutta
(No. £fp 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 £ 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).

Jiit. and Icon. : Neumann, 1904, pp. 460-462, Fig. 2 (reproduced).

Howard, C. W., vm. 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,

/. rvhicundus

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 /j, 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. /. rubicundus S : venter and hypostome. (Neumann, 1904, Fig. 2.)

206

Genus Ixodes

Female (gorged, Fig. 198): Body elongate oval, may attain L. 10 mm.,
W. 65 mm. toward the posterior third, brick-red in specimens of small
size, maroon-red in others. Scutum a short oval, scarcely longer than
broad (16 x 14 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 IV1; anus fairly anterior; anal grooves
long, parallel, united in a flat curve in front of the anus. Spiracles
small, circular, whitish. Capitulum 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 140/4 long ; dorsal process long, parallel to the shaft ; external
article with 5 teeth growing progressively larger antero-posteriorly.)

Fig. 198. I. rubicundus ? : 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 j 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 tbe specimen examined by us the vulva is placed just behind coxae III.

/. simplex 207

Description based on two Js and seven ? s found on sheep 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 (13 x 115 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 (073 mm. I.); 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, l'2x 09 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 type 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*6 mm. Scutum subcircular, as broad as long
(11 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 hairs ; 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 055 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 denned, oval, longer than broad,
the space between them equal to their breadth ; ventral surface

/. japonensis and canisuga

209

uniform, slight auriculae. Hypostome long, narrow, 4 1 4 near the tip
(corona), then 2 1 2 with a median unarmed ridge. Palps medium,
article 2 scarcely longer than 3. Leys: coxae unarmed; coxa I with
internal angle sharp, all, especially coxa II, trenchant on their posterior
border. Tarsi long, slender, tarsus 4 tapering 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 i. The

same specimen as the one to which the preceding figure relates. The scutum is

injured, there being a triradiate crack in the centre, 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. Harmand, 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 plumbeus, in Wheler, 1899 ; see Wheler, 1906, p. 413.
Ixodes hexagonus var. inckoatus Neumann, 1901, p. 283; Wheler, 1906, 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 hexagonus 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 11 to 23 x
l-5 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 nearly 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 g : 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.)

canisuga

211

fairly distinct lateral teeth on either side and corresponding ventral
crenulations (varies somewhat in structure, see Fig. 203). Legs : coxae
practically unarmed, coxa I only bluntly prominent at its internal angle,
slight protuberances at the postero-lateral angles of coxae II-IV.
Tarsus 4 fairly long, prominent dorsally at some distance from its
extremity.

Fig. 203. /. canisuga 3 : oapitulum 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 4 : hypostome. Drawn from specimen (N. 1123) from Gowanburn,

N. Tyne, 1901. Presented by Mr E. G. Wheler. (Original, G. H. F. N. del.)
Fig. 205. I. canisuga l : legs 1-4 in profile, x 20. (Original, G. H. F. N. del.)

212

Genus Ixodes

Female (Figs. 206, 207) : unfed, 21 mm. I., may attain 8x5 mm.
Scutum : 1 x 09 to 1*1 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. 20G. Fig. 207.

Fig. 206. I. canituga ? : capitulum and scutum ; capitulum in ventral aspect. Found

on dog, Tynron, Scotland, by J. McMonnies, vn. 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
coruua; 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,

/. canisuga

213

about 7-8 teeth per file, 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. canisuga o : capitulum and scutum ; coxae and ventral aspect of capitulum ;
tarsus 4 ; anal groove and spiracle. Specimen from dog, Tynron, Scotland,
J. McMonnies coll., vn. 1908. (N. 447. Original, G. H. F. N. del.)

I

Fig. 209. I. canisuga, 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 dog, Thornhill, Dumfriesshire,
v. 1908 (J. K. Wallace); (N. 422, 447, °.s, os) from dogs, Auchenbrack,
Tynron, v. and vn. 1908 (J. McMonnies); (N. 402) °.s from sheep,
Blackadie, Sanquhar, V. 1908 (R. Bramwell). [Evans (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 </ found.
(Det. by Neumann)]. England: (N. 1123, J), Gowanburn, N. Tyne,
v. 1901, and (N. 1046, ?) from dog, same locality, v. 1901 (E. G.
Wheler); (N. 1079, J, ?) 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 horse, Cambridge,
V. 1908 (G. A. Banham); (N. 1068, j"s, ?s) from nest of Cotile
riparia, Lyndhurst, Hants, vil. 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,
Lougner Hall, Shrewsbury, IV. 1906 (R. F. L. Burton.) 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 ,/s, !j!s, Os) from nests of three sand-martins
(C. 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,
vin. 1908 (G. H. F. Nuttall). Prance: [Neumann, 1901, records (?)
"inchoatus" from the fox (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 red squirrel or 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 broad1, 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

1 The scuta of our three specimens measure respectively l'lxl-2, 1-2x1*1, and
lxl mm. Our most replete specimen measures 4-9 x 3'3 mm. Banks gives the body-length
at 6-5 mm., but docs not state if this is the maximum length which may be attained in
gorged females.

1. texanus

215

rather sharply prominent ventrally. Hypostome thick and strong, with
2 | 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. 7. texanus ? : eapitulum aud 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.1)

Fig. 211. I. texanus ? : eapitulum and scutum, eapitulum in ventral aspect, coxae,
anal groove, spiracle and tarsus 4. Drawn from specimen found on grey squirrel,
King's river, California, vm. 1896. (N. 270. Received from V. L. Kellogg and identified
by N. Banks. Original, F. M. Howlett del.1)

1 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.

216 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 grey squirrel,
King's river, California, vn. 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 ; see 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, 1*5-1 '65 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 posterior
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 ft I. (see Fig. 212). Palps 1 mm. I., narrow, cultriform ;
article 1 with ventral 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-extcrnal
spur on coxae II-IV ; tarsi slightly humped distally ; pads large, almost
as long as claws.

/. stilesi

217

Fig. 212. J. stilesi j : 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. 213, 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 Neumann, 1899, p. 121; 1902, pp. 119-121; 1904, p. 461 (no

figures).
Ixodes intermedins Neumann, 1899, p. 132; 1904, p. 451 (no figure),
not Ixodes eudyptidis 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 /. praecoxalis and
/. intermedins to the rank of synonyms of his /. eudyptidis Maskell.
We recognize /. signatus Birula ( = /. parvirostris Neumann, 1901,
and /. 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 (1'7 x 12 mm.1), hexagonal2, 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

CP r^v.

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 1-5 x 1 mm., that of
J. praecoxalis Neumann, 1899, p. 121 ; 1902, p. 119, measures M x 1mm.

5 The postero-lateral borders are more rectilinear than is shown in the figure.

/. neumanni

219

white hairs ; punctations inconspicuous. Venter : very finely striate,
without punctations, but with numerous 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:
(08 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 8. 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 1 2 behind, the outermost
teeth (10 per file) much the strongest1 ; 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. neumanni 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 | 4, that of 7.
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 os, collected in New Zealand by Suter and presented to Neumann by
Trouessart in 1894.

Neumann's specimen of /. intermedins (1 %) was collected by
Dr Schauinsland from Phalacrocorax sp., and i". praecoxalis (2 ? s, 1 o)
from jEstrelata 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, 1-6x1-3 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. Gapitulum short (shorter than in the figure);
base trapezoid, broader behind, with slight cornua ; porose areas very
large1, 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.

I. percavatus var. rothschildi 221

auricula forming a stout retrograde process. Digit? Hypostome
narrow, rounded distally, 4 1 4, teeth equal. Palps short, article 2 barely
longer than 3, 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 retrograde 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.
Lounsbury, 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. Gapitulum : 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 | 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. /. pilosus s : dorsum and venter, spiracle and tarsus 4. The <f 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. Pillers.
(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

/. pilosus

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 numerous,
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 j 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. I. pilosus ? (unfed) : dorsal aspect of body, venter, spiracle, tarsus 4.
(N. 927. Original, G. H. F. N. del.)

224

Genus Ixodes

Fig. 219. I. pilosus ? : oapitulum in ventral and dorsal aspects. From the same ? as in
Fig. 218. (Original, G. H. F. N. del.)

Nymph (Fig. 220): resembles the $, especially in 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).

I. pilosus

22f>

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 </" 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, horses 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, xn.
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. howardi.

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. howardi was a microscopic preparation of
the ? , which appears to us identical with the specimens of /. pilosus
from Lounsbury'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 vin. 1908, p. 95, PI. IV, Figs, a-k ;
describes (J , ? , larvae, eggs ; figures <J and $ and details thereof somewhat
sketchily.

Male: resembles the type, but is smaller, 2-.3xl-2 mm. (type measures
3*15 x 1*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 smaller, shallower,
wider apart. Legs much feebler, especially the coxae. General colour, unfed, orange-
red.

Description based on 2 $ s and 12 9 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. 27, W. T8 mm., body oval, at least twice as broad
behind as in front, brownish, broadest towards the posterior third.
Scutum 1*57 x 139 mm., narrow, elliptical, maroon-brown, glossy,

I. hmatus

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 084 mm. long,
narrow, cultriform, article 2 slightly longer than 3. Legs slender, long
(pair 4 about 2-o 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.

Pig. 222. Fig. 223.

Figs. 222, 223. J. lunatus ? : dorsum and venter. (Neumann, 1907, Figs. 1 and 2.)

Description based on 1 $ from Hallomys audeberti Jent. (Muridae),
Madagascar (Leyden Natural Hist. Museum).

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 (27 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 (055 mm.), slightly longer than broad, broadest just
behind the palps. Hypostome (Fig. 224 (7) short and broad, sometimes
slightly indented, dentition 4 1 4 to 3 1 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. roam s: .4 venter; B digit, x210; C hypostome, x 135.
(Neumann, 1899, Fig. 12.)

/. rasus

220

Female (Figs. 225-226) : Body rather short-oval, broadest at the
posterior third. Scutum (1*9 x 1*5 mm., Nn.), reddish brown, oval,
glossy, dark brown, with very numerous fine, equal punctatious ; 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 (l-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. 225). Hypostome lanceolate ;
teeth 4 1 4, then 3 | 3, finally 2 j 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 f, but the spurs tend to become obsolete in large ?s.

Fig. 225. /. ratus ? 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 man and dog, Kumasi, Ashanti, by W. Oraham,
x. 1907. (N. 928. Original, G. H.F.N, del.)

230

Genus Ixodes

Fig. 226. I. rasus ? : 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., g % 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 22 x 12 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

/. Uganda mix

231

behind ; cervical grooves narrow, superficial ; punctations many, distant,
very fine (inconspicuous). 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 (05 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. /. ugandanus l : 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*8 xl'l mm.), broadest
slightly behind the anterior third, dark brown, glossy, slightly emar-
ginate1 in front, very fine punctations, slight lateral grooves1, 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, 0'92 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-brown.
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.)

i Neumann states that the ? scutum is not emarginate and does not possess lateral
grooves ; that the spiracle is small, and the porose areas oval.

/. cordifer 233

Neumann's original description was based on 1 £ and 1 % in coitus,
found on Aulacodus sp., from Uganda, collected by E. Degan (British
Mus.). Neumann, 1907 a, p. 18, has since reported 1 </, 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. n. 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 l-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 068 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, vn. 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 (17 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 front1, finely punctate and fused
behind the point of the anal plate ; spiracles large, oval, much longer
than broad. Capitulum 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. I. cordifer <? : venter and tarsus 4. (Neumann, 1908, Fig. 1.)
1 Neumann's figure, reproduced here, appears to be inaccurate in this respect.

/. holoci/clus

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 / from Sekroe, New
Guinea, collected by S. Schadler (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 pu notations 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. /. holocyclus s : 1, ventral aspect ; 2, hypostome, x 75.
(Neumann, 1899, Fig. 24.)

16-2

236 Genus Ixodes

as long as broad, tapering much behind, whitish. Capitulum short
(0-6 mm.), its base quadrangular, scarcely broader than long. (Digit, see
footnote1.) Hypostome (Fig. 230) short and broad, with some blunt
denticles in front, then 2 1 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 17
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 (2'3 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 footnote2.) Hypostome (Fig. 231) long,
narrow, lanceolate, numerous small sharp teeth, followed by 3 | 3 files of
12-13 stout teeth, increasing3 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 /t long; dorsal process (?) ; external article 4-toothed, the two anterior teeth
very small, the others strong.

a Digit 210 /i 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.

3 Neumann states they diminish outwardly, but this does not agree with the figures
nor with our specimens.

I. holocyclus

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 /. ornithorhynchi (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. /. holocyclus ? (partially gorged): 1, capitulum and scutum in dorsal aspect;
2, venter ; 3, hypostome, 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 07 mm.
Capitulum 0'88 mm. long (dorsally); base more conical. (Digit, see
footnote1.) Hypostome narrower, sharper, with 2 j 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 n long ; dorsal process a subterminal hook at the end of a narrow shaft.

l»:js

Genus Ixodes

Fig. 232. I. holocyclus o : dorsal aspect of capitulum, x 88 /t I.
(Neumann, 1899, Fig. 26.)

India (R. Blanchard 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, vn. 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 (</*, $ ) 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 (<J, reproduced).

Male (Fig. 233) : Body oval, 2"3 x T38 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

/. schillingsi

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 (05 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 /a I. (see Fig. 233).
Hypostome short, broad, with six transverse rows of 6-8 rounded teeth
and a large basal tooth 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 ; II 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. schillingsi <f : 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. 43 mm., W. 35 mm., yellowish white. Scutum reddish
brown, much longer than broad (T5 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 1 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. schillingsi ? : capitulum 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.)

/. schillingsi

241

indication of a spur at the postero-external 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. schillingsi 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 $s and
1 o, found on Colobus caudatus, Kilimandjaro, Kibonoto, 8. vn. 1905,
and 1 ^ found in the same locality on grass, 12. vn. 1905, collected
by the Sjiistedt Zoological Expedition, German East Africa. Neumann
(1910, p. 171) has recently described the £ from a specimen found in
copula, collected by G. Vasse in 1907, in Mozambique, Province of
Gorongoza (Paris Mus.) ; the description we give of the <$ is condensed
from Neumann, who states that the 2 % s collected with the £ were
somewhat larger than the types (7 mm. 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, PI. 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*2x1*9 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.
Capitulum 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 1 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.

Nymph {Ixodes ornithorhynchi Pagenstecher) : resembles $ . Body
relatively broader (3 x 1*5 mm.), oval, the colour varying from brownish
yellow to earthy yellow. Capitulum with base relatively shorter and
broader, without porose areas. Digit 85 //. long, as in $. Hypostome
with 2 | 2 files of 5 teeth.

Neumann was the first to describe the species adequately from
(a) six $8, from the Paris Museum, of which five were from the
Ornithorhynchus(Luca& coll.),and one from the Marianne Islands, collected

/. omithorhynchi

243

Fig. 236. I. omithorhynchi ? : 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.)

Vif. 237. Fig. 238.

Fig. 237. /. omithorhynchi ? : tarsi 1 and 4, x 45. (Neumann, 1899, Fig. 19.)
Fig. 238. I. omithorhynchi ? : C, left digit viewed dorsally to one side, 150^ I. ; H, hypo-
stome, x 75. (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 Ornithorhynchus ;
he took them for females); (c) three %s, from the Ornithorhynchus
(Me'gnin 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 M^gnin 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.
Scutum1 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 /t), 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 I. ornithorhynchi
(Fig. 237). Claws short ; pad almost reaching their free extremities.

1 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).

1. tasmani

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.

Fig. 239. I. tasmani ?: A, capitulum and scutum ; B, hypostome, xl05;
C, tarsus 1, x 70. (Neumann, 1899, Fig. 20.)

Fig. 240. I. tasmani y (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.

PL 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 ( 9 , reproduced).
Ixodes tenuirostris Neumann, 1902, p. 119, Fig. 5 (jj, reproduced).
Ixodes tenuirostris in Wheler, 1890, p. 40 ; 1906, p. 414, Figs. 23, 24 ( 9 capitulum,

<$ 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 : To x 08 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; punctatioDS 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 divergent 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 (033 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.

Kg. 241. Pig. 242.

Fig. 241. I. tenuirostris 3 : capitulum in dorsal aspect. From the type at the British

Museum. (Original, sketch by G. H. F. N.)
Fig. 242. J. tenuirostris 3 : posterior part of venter. (Neumann, 1902, Fig. 5.)

TICKS

PLATE V

Fig. 1.

Fig. 2.

m

^^m

^^i^tf

m

19

Fig. 3.

Fig. 4.

Figs. 1, 2. Ixodes tenuirostru.

Fig. 1. <f venter. (Original, E. G. Wheler, phot.)

Fig. 2. ? venter, mounted specimen, x 11. (Original, E. G. Wheler, phot.)

Figs. 3, 4. Ixodes putus.
Fig. 3. i dorsum, x 7. (Wheler, 1906, Fig. 26.)
Fig. 4. ? dorsum, x 7. (Wheler, 1906, Fig. 25.)

(All specimeus of British origin.)

/. tenuirostris

247

Female (PL V, Fig. 2; Figs. 243, 244): Body lead-coloured,
elongate-oval, may attain 8x5 mm. in fully gorged specimens. Scutum:
08 X 07 mm., broadest near the posterior end ; antero-lateral borders
subrectilinear, divergent, united by a broadly rounded posterior border ;
emargination slight ; surface 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. Gapitulum 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 J1 ; tarsi slender, tapering gradually ;
pad almost as long as claws.

Fig. 243.

Pig. 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 %&, two os) from Evotomys glareolus,
Island of Rugen (Lemm coll., Berlin Mus.) and (one % ) from " Arvicola
pratensis," Painswick, Gloucestershire, England (E. G. Wheler's coll.,
1893). The </\ 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.1 (N. 1165, $ s), from field vole, Longner Hall, Shrewsbury,
VIII. 1910 (R. F. L. Burton). (N. 622, %), from Microtus amphibius
(water vole), England, IV. 18921. (N. 620, %) from Mus minutus
(harvest mouse) ; no particulars1. (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. 18991.

Switzerland: (N. 624, 625, 1069, a J captured with mouthparts
buried in $'s body close to anus, $s and Os) from Microtus arvalis,
Campfer, St Moritz, Engadine, vn. 1904 (K. Jordan)1. (N. 619, f and
$ ), from Evotomys glareolus (bank vole), Campfer, St Moritz, Engadine,
vn. 1904 (K. Jordan)1. (N. 1161, %), from Tarasp, summer of 1901 \

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 13 mm.), broadest near the posterior end; uniformly punctate
except along the cervical grooves, which are otherwise obsolete; the

1 Ex N. C. Bothsehild coll.

2 Recalling a conventional form of heraldic shield, hence fecialis.

/. 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-
lateral^ 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. Legs 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, in. 1900, presented by the Hon. N. C.
Rothschild.

Fig. 245,1.

Fig. 245 B.

Fig. 246.

Fig. 245. I. fecialis ? ; 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. Capitulum : 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. vm. 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 /. 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. /. tenuirostris 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
27 mm. Scutum hexagonal, with rounded angles, shorter than broad

/. australiensis

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. Anal grooves closed posteriorly ; oval, tangential to the

Fig. 247. J. australiensis ? : capitulum 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 = l mm. Original, G. H. F. N. del.)
Fig. 249. I. australiensis ?: 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. Cajritulum very short (540 fi), 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 1 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 Ganis 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 Ganis 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
cordiform1, with rounded sides, broadest (l-35 mm.) toward the middle,
1'25 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 Bcutum in the type are very ill-defined. They
appeared to us almost rectilinear (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.

7. 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. Capitulum short (07 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 and broad, article 1 very broad,

Fig. 250. J. vestitus ?: 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 very small, terminal. Legs1 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. vettitut ? : capitulum in 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. 7. 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 Myrmecobius fasciatus Waterhouse
(Dasyuridae), West Australia (Neumann's type, British Museum). Our
collection contains (N. 642) $ , O and larvae from Diemenia superciliosa
(a snake), Herdman's Lake, near Perth, W. Australia (B. H. Woodward,
ex N. C7 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, PL XIII, Fig. 4 (o).
We are indebted to the author for sending us one of his types to examine
in vn. 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 putus by Neumann.

Ixodes borealis Kramer and C. J. Neuman, 1883, p. 518, PL

Ixodes fimbriatus 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 fimbriatus— putus <$ . (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 putus (Cambridge) in Neumann, 1902, p. 117, Fig. 4 (capitulum of <J,
dorsal and ventral aspects, palp in profile, somewhat inaccurate, consequently
not reproduced. Lahille, 1905, p. 26, reprints description of <J, $ and
o given by Neumann ; on pp. 138-148, more detailed description (original),
gives many measurements, etc., PL XIII, coloured Figs. 1-13 of ° and o
with details.

Ceratixodes putus Cambridge, in Wheler, 1906, pp. 415, 416, Figs. 25 ( $> ) and
26 ( (J ) photographs (reproduced).

Ixodes borealis Kramer and Neuman, in Evans, 1906, pp. 85, 86, Fig. of °
(reproduced).

? Ixodes uriae White, 1852, p. ccx, found on Uria troile, Baffin's Bay (P. C.
Sutherland).

? Ixodes eudyptidis Maskell, 1885 (see Notes on Doubtful Species, p. 291).

Male (PI. V, Fig. 3; Text-fig. 254): very large (37 mm. I,
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 punctatious, 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 dorsally 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. Capitulum very small ; base twice as broad as
long, broader in front, the posterior border straight, 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 Bhosilly, Gower, Glamorganshire, 1909, by
Dr J. W. W. Stephens. (Original, G. H. F. N. del.)

2f>8

Genus Ixodes

Female (PL V, Fig. 4 ; Text-figs. 255-259) : L. 33 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 (03 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 1 2 strong equal teeth, about
8 per file. Legs : long and more slender than in the J, 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. /. putus ? : dorsum, venter, right palp in profile, spiracle and tarsus 4. Same
origin as the J 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 J. borealis
Kramer and Neuman, 1883.)

7. putus

259

Fig. 257. X. putus ? : capitulum in dorsal and ventral aspects. (Neumann, 1899,
Fig. 8, G. Marx del.)

Fig. 258.

Fig. 259.

Fig. 258. I. putus ? : 1, scutum; 2, left digit; 3, hypostome; 4, tarsus 4. (Neumann,

1899, Fig. 9.)
Fig. 259. I. putus ? : spiracle, G. Marx del. (Neumann, 1899, Fig. 7.)

Nymph (Fig. 260) : bears an unmistakable resemblance to the $ .
L. 125 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 i and ? above figured. (N. 968.
Original, G. H. F. N. del.)

260 Genus Ixodes

Our description of /. putus is based upon numerous specimens
(</". ? i 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) : o a on guillemots ; % s on puffins, Morthoe,
N. Devon (Pocock); fa, $s, on cliffs frequented during the nesting
season by guillemots and other sea-birds, at Bempton and Buckton on
the Yorkshire coast (Mr Howett, of York, occasionally found sexes in
copula beneath stones) ; on Pinnacle Rocks, Fame Islands (found by
Mr Wheler, who has- kindly presented us with some of these specimens :
N. 935, 936, 937). We have adults (N. 454, J, ?) found on ledges
frequented by guillemots, Bempton Cliffs, Yorkshire, vi. 1908, by
Mr O. Grabham, who states that the jackdaws feed on the ticks with
avidity. Wales: (N. 968, </" ? o) found on cliffs, Rhosilly, Gower,
Glamorganshire, "guillemots, razorbills and puffins breeding there,"
vi. 1909 (sent us by Dr J. W. W. Stephens). Scotland: (N. 459, %)
from seagull, Handa Island, Sutherland, vm. 1908 (presented by
Dr W. L. H. Duckworth) ; Mr W. Evans, of Edinburgh, has kindly sent
us specimens he collected: (o) from a gannet, Bass, vi. 1910, and ($)
from a puffin, Firth of Forth, vn. 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 Js. Asia

/. signatus 261

and Australasia : The specimens (</" and $) described respectively as
jimbriatus and borealis by Kramer and Neuman were found on BERING
Island, Bering Sea, by the 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.) aud 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 signatus Birula, 1895, pp. 357, 358, PI. I, Figs. 10-13
( ? capitulum in dorsal aspect, tarsus 1, scutum, coxa I, drawn from a
mounted specimen aud 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 eudyptidis var. signata Neumann, 1904, p. 451.
Cerativodes 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. Dept. of Agriculture), 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 35 mm. when replete), elongate,
yellow (in young specimens). Scutum: 17 x 1"2 mm., 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

l>i;l>

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.

Fig. 261.

Fig. 262.

Figg. 261, 262. I. tignatus ? : 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.)

7. signatw

263

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. J. signatus o : capitulum and scutum, ventral 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.

<3

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, Pai-ific 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
Phalacrocoraw 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 !j! 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-o mm. (capitulum
included), broadest (3"2 mm.) in front of the middle of the length.
Scutum oval lozenge-shaped, 1'5 mm. I., 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 (065 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 | 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,

/. unicavatus

265

relatively short and broad, uniformly broad along their length ; article 2
half the length of 3 ; article 4 terminal, clearly visible. Legs 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. 266.

Fig. 265. I. unicavatus ? : capitulum and scutum. (Neumann, 1908, Fig. 1.)
Fig. 266. /. 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. I. 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.)

„ r 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, os, from Phalacrocorax
graculus, Cromarty, m. 1898 (W. Roy); both lots presented by
Hon. N. 0. 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, XL 1906.

Note: In a tube labelled "I. eudyptidis 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 I. 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 I. 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. angustus Neumann, 1899, on
p. 195 of this work) ; Neumann, 1910, p. 30.

Male (Fig. 268) : Body oval, narrow, broader and rounded behind,
4'6 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 l-9 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 grooves
slightly marked ; lateral grooves distinct, straight ; punctations many,
equal, very fine. Dorsum with a deep marginal groove, 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. I. loricatus s : dorsal and ventral aspects. (Neumann, 1899, Figs. 15, 16.)
Fig. 269. 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 /j, long, resembling that of /. 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 leugth, 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. Venter 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 08 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.).

/. loricatus var. spinosus

289

Neumann (1901, p. 283) examined a gorged $ from Tierra del
Fuego (C. 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, XL 1901 ; (N. 638) f, $, from
Didelphys sp., Sapucay ; W. F. Cooper's collection contains a </" from
Estancia Cooper, Paraguay, VI. 1906 (Foster).

Ixodes loricatus var. spinosus Nuttall, 1910.

Fig. 270.
Lit. and Icon.: Nuttall, xn. 1910, p. 411, Fig. 5 (reproduced).

Male: unknown.

Female : differs from the type as follows :

It is larger and less hairy generally. The scutum (16 x 125 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 (0'7 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. /. loricatus var. spinosus ? : 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 poster o- 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, in the month of May (ex Hon. N. C. Rothschild's
collection).

50. IXODES COXAEFURCATUS Neumann, 189a.

Fig. 271 (original).

Lit. : Neumann, 1899, p. 127 (no figure).

Male : Body elongate, narrow, sides parallel, 3"7 x 17 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 j : dorsum, parts of venter, spiracle and tarsus 4. Sketched
from the type in the Berlin Museum. (Original, G. H. F. N. del.)

I. coxaefurcatus 271

adanal plates with their sides curved and parallel. Spiracles large,
oval, with long axis directed forward and macula eccentric. Capitulum
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, Brazil1 (Hensel coll., Berlin Mus.).

51. IXODES VESPERTILIONIS Koch, 1844.

Figs. 272-282 (Figs. 272, 273, 277, 280, 281 original). See also
PI. VI, Figs. 1 and 2 (?s).

Synonymy : Ixodes vespertilionis Koch, 1844, H. 37, Fig. 9 ( $ ) ; 1844, p. 2.32, listed ;

1847, p. 21.
Ixodes flavipes Koch, 1844, H. 39, Fig. 2 (? ?); 1844, p. 232, listed; 1847, p. 21.
Ixodes troglodytes Schmidt, 1853 {vide Frauenfekl, 1853, p. 57) ((J).
Eschatocephalus gracilipes Frauenfeld, 1853, p. 57, PI. ( <J : dorsum, venter, capit-
ulum ; $ : dorsum and capitulum. Sketchy, but resembling I. vespertilionis) ;

1857, PI. II ( ? ).
Haemalastor gracilipes Frauenfeld, 1854, p. 23, PL Figs. 4, 6 (?).
Sarconissus flavipes Kolenati, 1856, p. 21, 1857, p. 21, PI. I, " <} ," immature

stages. See Neumann, vm. 1910, p. 191.
Sarconissus hispidulus Kolenati, 1856, p. 21 ( $ ).
Sarconissus brevipes Kolenati, 1856, p. 21 ( $ ).

Sarconissus kochi Kolenati, 1856, p. 21 (?) ; 1860, p. 573, PI. II, Fig. 4 ( <$ ).
Sarconissus flavidus Kolenati, 1856, p. 21 ( ? ).
Sarconissus exaratus Kolenati, 1856, p. 22; 1860, p. 757, PI. I, Fig. 2 (Neumann

det. vm. 1910, p. 191).
Haemalastor gracilipes (Frauenfeld) Kolenati, 1860, p. 573. Neumann, det. vm.

1910, p. 191.
Eschatocephalus frauenfeldi'L. Koch, 1872, p. 26, PI. II, Figs. 29-32 (<J); 1877,

p. 150.

1 Neumann erroneously states that the specimen came from Siwa, Egypt.

272 Genus Ixodes

Eschatocephalus seidlitzi L. Koch, 1872, p. 26 ((J); 1877, p. 151.

Ixodes longipes Lucas, 1872, p. lxxiv. (cJ).

Ixodes siculifer Megnin, 1880, p. 132 ($).

Haemalastor vespertilionis (C. L. Koch) Neumann, 1899, p. 169; see further
under Literature.

Eschatocephalus vespertilionis (Koch) Neumann, 1901, p. 290; 1902, p. 116.

1 Sarconissus nodulipes Kolenati, 1860, p. 576, PI. I, Fig. 3.

1 Eschatocephalus fiavipes (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).— 1890. 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 ( (J : Fig. 27, venter ; Fig. 28, ventral aspect of capitulum (these two
figures not reproduced), hypostome and digit, leg 1, foot ; 2 '■ 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. 19a (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 Eschatocephalus and Haemalastor, pp. 134, 281.

Male (Figs. 272-276) : Body long-oval, narrower in front, yellow.
Scutum (3'5 x 21 mm.), very long and narrow, indistinctly denned, 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. 273.

Fig. 272. I. vespertilionis 3 : venter. Specimen from Grotto of Alsaaua, E. Simon

coll., 1896. (Neumann coll. 760. Original, G. H. F. N. del.)
Fig. 273. I. vespertilionis j : capitulum. Same specimen as Fig. 272. (Original,

G. H.F.N, del.)

Fig. 274.

Fig. 276.

Fig. 274. /. vespertilionis s : A, hypostome, x84; B, B', right digit in dorsal and

ventral aspects, x238. (Neumann, 1899, Fig. 29.)
Fig. 276. I. vespertilionis i ; ambulacrum, x 110. (Neumann, 1899, Fig. 31.)

Fig. 275. J. vespertilionis 3 : 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*9x1*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. veepertilionis ? : eapitulum (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. 278. J. vespertilionie ? : A, bypostome, x85; B, left digit in dorsal aspect, x 225.

(Neumann, 1899, Fig. 33.)
Fig. 279. /. veepertilionis ? : capitulum in dorsal and ventral aspects, x 40. Drawn from

mounted specimen. (Neumann, 1899, Fig. 32.)

/. vespertilionis

275

sides ; spiracle as in £. Gapitulum : base shorter and broader than in
the </", with salient lateral points ; porose areas large and near together ;
palps normal ; hypostome tapering to a point, 4 j 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 (PI. 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. /. vespertilionis larva: capitulum and scutum; ventral aspect of capitulum,

with coxae, tarsus 3. Specimen found on Rhinolophus sp. by Trouessart, 1896.

(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 J1, I J, lo and 1 larva kindly lent by Professor
Neumann.

This species appears to be widely distributed geographically. It has
hitherto only been found on Cheiroptera 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,
beiDg 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, vn. 1902 (Dublin Nat. Hist. Mus.,
specimen determined by us.) Prance : One of us (G. H. F. N.) examined
a number of mounted specimens in the Me'gnin collection through the
kindness of Professor R. Blanchard (Paris). They had mostly been sent
to Me'gnin by H. Gadeau de Kerville, of Rouen, during n-v. 1883 ;
$ s and O from Plecotus auritus, grotte de la Briqueterie, 10. III. ;
os and larvae from Rhin. ferrurn equinum, grotte Cuvette, 10. III.;
larvae from Rhin. hipposideros, grotte de Dieppedalle, Seine inferieure,
10. in.; larvae, no particulars, 25. n. ; larvae from Rhin. hippo-
sideros, Carrian de la Londe, 14. v. ; all of these were labelled
/. longipes by Megnin, 1 $ , labelled " /. aculeifer 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 (/"s from caves and grottoes in the following places : Abeille, Ariege ;
Aspradels and Espezel, Pyrenees ; les Baux, near Aries ; Penne, Tame ;
Serres, Hautes Alpes ; 3 $s respectively from Bize and Minerve,
He'rault, 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 Ordufia,
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 54 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 Rosenmiiller cave,
near Muggendorf, IX. 1869, and in caves near Almas, Frankischer

TICKS

PLATE VI

Fig. l.

Fig. 2.

Figs. 1, 2. Ixodes venpertilionis.

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, vn. 1902. (Dublin Mus.,

Original, L. E. Robinson, phot.)

/. vespertilionis 277

Jura. Austro-Hungary : os and larvae have been found on Rhin.
ferrum equinum 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 had 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. ferrum 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
Institut 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.
hexagonus : ,, „

Special Hosta.

canisuga: ,, ,,
brunneus : ,, ,,

Birds.

vespertilionis : ,, „
tenuirostris : Great Britain, Switzerland,

Island of

Biigen.

Bats.

Small mammals,

acuminatus : Italy.
putus: Great Britain.
unicavatvs : Great Britain.
caledonicus : Scotland.

Marine birds.
Marine birds.
Birds.

AFBICA.

North Africa :

ricinus : Algeria, Tunisia.

hexagonus: Algeria.

vespertilionis : Algeria. Bats.

nigricans: Madeira (perhaps = ricinus, see text).
Central Africa:

rasus : E. and W.

ugandanus : E. and W.

ugandanus var. djaronensis ; Abyssinia.

schillings^: E.

rubicundus var. limbatus : Congo Free State.

cavipalpus: Angola, N.W. Bhodesia.
South Africa :

rubicundus : Cape Colony.

pilosus : Cape Colony.

percavatw : Tristan d'Acunha Islands (W. of Cape Colony).
Madagascar :

lunatus.

ASIA.

ricinus : Transcaucasia, Arabia, Japan.

ricinus var. ovatus : Japan.

acutitarsus : India, S. Formosa, Japan.

signatus: Japan, Unalaska (Aleutian Islands). Marine birds.

japonemis : 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.
cordifer: New Guinea.

australiensis : Australia.

vestitus : ,,

holocyclns : ,,

fecialis : „

fecialis var. aegrifossus : Australia.

vespertilionis : Australia. Bats.

tamtam : Tasmania, Island of St Pierre.

ornithorhynchi : Tasmania and Marianne Islands. Ornithorhynchus.

putus : Campbell and King's Islands. Marine birds.

neumanni : New Zealand. Birds.

AMEBICA.

North America :

putus : Alaska, Newfoundland. Marine birds.

angustus : Alaska, Canada, United States.

ricinus: Canada, United States.

ricinus var. scapularis : Canada, United States.

marxi : Canada and United States.

texanus : Canada and United States.

hexagonu8 var. cookei : Canada and United States.

hexagonus : United States.

canisuga :

brunneus :

signatus: „ Marine birds.

pratti :

sculptus :

diversifossus :

dentatus :

ricinus var. calif ornicus : United States.

rubidus : Mexico.

bicornis : , ,

loricatits var. spinosus : Mexico.

minor : Guatemala.

fuscipes: Panama.
Sooth America :

fossulatus : Ecuador.

elegans : Chili.

boliviensis : Bolivia.

fuscipes : Brazil.

hexagonus : Locality not stated.

coxaefurcatus : Brazil.

loricatus : Brazil, Paraguay, Argentina, Tierra del Fuego.

auritulus : 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.

americanui Linn., 1758, p. 615 ; Fabricius, 1794, p. 428 (given as synonym of
Ixodes americanus by Fabricius, 1805, p. 355, and many subsequent
writers) = Amblyomma arnericanum (Linn.) Koch, 1844.
frontalis Panzer, 1795, H. 59, n. 23 = ? I. 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. lipsiensis (Fabr.) condemned, see p. 286.
pcdlipes „ 1794, p. 426=? I. brunneus Koch, 1844.
plambeus 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 = /. ricinus (Linn.) $.
ricinus Linnaeus, 1746, p. 480=7. ricinus (Linn). $.
tristrialus Panzer, 1795, H. 59, n. 24 = 7. tristriatus (Panzer), condemned,

see p. 290.
undatus Fabricius, 1794, p. 427 (undeterminable).
Ceratixodes Neumann, 1902.

putus (Cambridge), 1878, in Neumann, 1902, p. 117=7. pulus (Pickard-

Cambridge).
putus Cambridge, in Banks, 1908, p. 21 = Ixodes putus (Pickard-Cambridge).
signatus Birula, in Banks, 1908, p. 21 =7. signatus Birula, 1895.
Crotonus Dumeril, 1822.

ricinus Dumeril, 1822, p. 56 = 7. ricinus (Linn.).
Cynorhaestes Hermann, 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. ricinus by Neumann.)
ricinus Hermann, 1804, p. 65, possibly = 7. ricinus (Linn.).
hermanni Risso, 1826, p. 183 = 7. ricinus (Linn.).
Dermanissus Kolenati, 1866.

rubiginosus Kolenati, 1857, p. 20, PI. 1 = 7. rnbiginosus (Kolenati), in Neumann,
1899. Condemned, see p. 288.

Condemned Species, etc. 281

Eschatocephalus Frauenfeld, 1853.

acutitarsus (Karsch), in Neumann, 1901=7. acutitarsus (Karsch).

brevipes Neumann, 1899, p. 179 (crassipes Joseph, 1882, renamed)=7. brevipei

(Neumann). See Notes on Doubtful Species, p. 291.
crassipes Joseph, 1882, p. 16. See brevipes above.

flavipes (Koch) in Bonnet, x. 1908, p. 325 = probably I. vespertilionis Koch.
frauenfeldi L. Koch, 1872, p. 26, PL II, Figs. 29-32 {$); 1877, p. 150=/.

vespertilionis Koch, 1844.
gracilipes Frauenfeld, 1853, p. 55. PL— Z vespertilionis Koch, 1844
seidlitzi L. Koch, 1872, p. 26 (<J); 1877, p. 151 = 7. vespertilionis Koch, 1844.
vespertilionis (Koch) Neumann = 7 vespertilionis Koch, 1844.
(See Sarconyssus.)
Haemalastor Koch, 1844.

acutitarsus Karsch, 1880, p. 142 = 7. acutitarsus (Karsch), from Japan.
brevipes Neumann, 1899, p. 179 = 7 brevipes (Neumann), see p. 291, Notes on

Doubtful Species.
crassipes Kolenati, 1857, p. 437 ; 1860, p. 578, PL II, 5 ; Neumann, 1899, p. 178,

and 1901, p. 290. Found on Rhinolophus clivosus, in Egypt. Description

insufficient.
crassitarsus Karsch, 1880, p. \A\= Amblyomma longirostre (Koch), Neumann,

1905, p. 231, not Amblyomma crassitarsus (Karsch) Neumann, 1901, p. 315;

not Hyalomma crassitarsus (Karsch), as stated in Neumann, 1899, p. 293.

Original description insufficient ; from Caracas.
longirostris Koch, 1844, p. 233 ; 1847, p. 49 = Amblyomma longirostre (Koch)

Neumann, 1905, p. 231, not Hyalomma longirostre (Koch) as stated in

Neumann, 1901, p. 315.
Hyalomma

puta Pickard-Cambridge, 1878, p. 222 = 7. putus (Pickard-Cambridge).
Ixodes Latreille, 1796.

acanthoglossi 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 7. savignyi by Gervais,

1844, p. 245. See savignyi= Hyalomma aegyptium (Linn.).
aequalis Banks, 1909. See Notes on Doubtful Species.
affinis Neumann, 1899, p. 120= Ixodes ricinus var. scapularis (Say) Nuttall and

Warburton, 1910.
africanus Megnin, 1876, p. 831 = Hyalomma aegyptium (Linn.), fide Neumann,

1899, p. 286.
albipictus Packard, 1867, p. 65 = 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. 7 albipictus also referred to

as attacking man by Holland, 1898, p. 96.
algeriensis Megnin, 1880, p. 121, PL II. Undeterminable : see Neumann, 1901,

p. 314.

n. i. 19

282 Genus Ixodes

ameivae Pagenstecher, 1861, p. 41, PI. II, Figs. 23, 24, hypostome and part of
basis capituli with mandibles; probably an Amblyomrna or Aponomma,
from Mexico, as rightly suggested by Neumann, 1899, p. 159.

" americana" (Linn.) Koch, of Murray, 1877, p. I9i = americanus, see below.

americanus (Linn.) Fabricius, 1805, p. 355 = Amblyomrna americanum (Linn.)
fide Salmon and Stiles, 1901, p. 475.

americanus Gervais, 1844, p. 247 ' = Amblyomrna americanum (Linn.) fide
Neumann, 1897, p. 366.

angustus Neumann 1901, p. 284 = Ixodes loricatus Neumann 1899.

annulatus Say, 1821, p. 75; 1859, p. \9 = Boophilus annulatus (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
australis, 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.

"aquilae" (in Hamburg Museum) = Amblyomrna 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.

aureolatus (Fabr.) Fabricius, 1805, p. 355 (Acarus aureolatus Fabricius, 1794,
p. 428) = Amblyomrna aureolatus (Fabr.).

auricularius 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 = Amblyomrna tenttcdinarium Koch, 1835-
1844, Heft xi, 1 ; fide Neumann, 1901, p. 303.

australasiae Fabricius, 1805, p. 355 = ? Amblyomrna.

autumnalis Leach, 1815, p. 398 = hexagonus Leach, 1815.

avuugus Berlese, 1890, fasc. lv, No. h — brunneus Koch, 1844.

bengalensis Supino, 1897, p. 15, PL II, Figs. 6-10 ; 1897, p. 268, PI. XII, Fig. 4
(tarsus). Description insufficient, from Burma.

berlem Birula, 1895, p. 359, PI. II, Figs. 1-6. Description and figures
insufficient ; a $ Ixodes from E. Africa.

bibroni Gervais, 1842, p. xlviii ; Walckenaer and Gervais, 1844, p. 248;
Packard, 1867, p. 68 ; also Neumann, 1899, p. 157. Probably an Amblyomrna ;
from S. America.

Condemned Species, etc. 283

bifurcatus Neumann, 1899 ( <? ), not /. brunneus Koch, as stated in Neumann,

1901, p. 347 = Dermacentor bifurcatus Neumann, 1904, p. 453.
bimaculatus Denny, 1843, p. 312, PL XVII, Fig. 1 (?) = Amblyomma

hippopotamense (Denny).
bipunctatus Risso, 1826, p. 183. Nominal species, see Neumann, 1899, p. 158.

From S. France.
birmanensis Supino, 1897, p. 12, PL I, Figs. 6-10 ; 1897, p. 247, PL XII, Fig. 2
(tarsus 1) ; see also Neumann, 1897, p. 164. Description insufficient ; from
Burma.
boarum Stoll, 1886-1893, p. 18, PL XIII, Fig. 4= Amblyomma dissimile Koch.

See Neumann, 1899, p. 227 (o).
borealis Kramer and Neuman, 1883, p. 518 = 7. putus (Pickard-Cambridge).
bovis Riley, 1869, p. 168 = Boophilus annularis (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.
cajennensis (Fabricius) Fabricius, 1805, p. 354. {Acarus cajennensis Fabr.,

1794, iv, p. 427), undeterminable.
calcaratus Birula, (?), p. 137 = Boophilus annulatus (Say) Curtice, 1891.
californicus Banks, 1904, p. 369, PL XLI, Fig. 57 ; 1908, p. 24, PL II, Fig. 12 =

I. ricinus var. californicus (Banks) Nuttall and Warburton, 1911.
camelinus Fischer von Waldheim, 1823, p. 13, PL, Figs. 1 and 2 = Hyalomma

aegyptium (Linn.).
cenereolus Lucas, 1840, pp. 47, 48, PL VII, Fig. 10 (after Gervais, 1844, p. 244).
Insufficiently described ; see Neumann, 1899, p. 165; from Canary Islands.
chelifer Megnin, 1880, p. 132 = Haemaphysalis concinna Koch, 1844.
chordeilis Packard, 1867, p. 67 = Haemaphysalis leporis palustris (Packard), fide
Neumann, 1897, p. 343, but stated to be a distinct species of Haemaphysalis
by Banks, 1908, p. 34.
cinctus Fabricius, 1805, p. 356 ; undeterminable.
ductus Lucas, 1840, pp. 47, 48, PL VII, Fig. 12 (after Gervais, 1844, p. 243).

Insufficiently described ; from Canary Islands.
columbae Fabricius, 1805, p. 356= Argas reflexus.
"communis" Marx (label: no date) = 7. ricinus var. scapularis (Say) Nuttall

and Warburton, 1911.
cookei Packard, 1869 a, p. 67 = hexagonus var. cookei (Packard) Nuttall and
Warburton, 1911 ; not Haemaphysalis concinna Koch, as stated in Neu-
mann, 1897, p. 338.
cornuger Kolenati, 1857, p. 431, PL VI, Fig. 52 (very poor) = Hyalomma aegyptium
(Linn.) ((J) 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.
crenatus Say, 1821, p. 76; 1859, p. 20. Original description useless.
crenulalus Koch, 1835-1844, H. 39, Figs. 8, 9 ; 1844, p. 232 (listed) ; 1847, p. 22
= 7. hexagonus Leach, 1815, from Germany. "I. crenulatus Koch," as
described by Canestrini and Fanzago, 1877, p. 116, is undeterminable.

19—2

284 Genus Ixodes

cruciarius Fitch, 1872, p. 366 ( = /. cookei fide Banks, 1908, p. 53) = /. hexa-

gonus var. cookei Nuttall and Warburton, 1911.
decorosus L. Koch, 1867, p. 241 = 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), /. plumbeus Duges

renamed = ? Rhipicephalus sanguineus (Latreille), as suggested in Neumann,

1897, p. 385, but undeterminable from description. On dogs in France.
dugesii Gervais, in Megnin, 1880, p. 126, Fig. 44 = Boophilus sp.
elegans Guerin, 1829-1843, p. 16, PI. VI, Fig. 1 = Amblyomma variegaium

(Fabricius), 1794 ; in Neumann, 1899, p. 268.
elegant Neumann, 1910, p. 191, Fig. 1=1. stilesi Neumann, 1911 (original

name preoccupied ; species renamed.)
elephantinus (Linn.) Fabricius, 1805, p. 351 (Ac. elephantinus Linn.) = ?

Amblyomma elephantinus (Linn.).
erinacei Audouin, 1832, p. 415, PI. XIV, Fig. 32 ( ? ) = / hexagonus Leach, 1815,

in Neumann, 1899, p. 129.
erinaceus Murray, 1877, p. 190 = 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) = Hyalomma aegyptium

(Linn.).
fimbriatus Kramer and Neuman, 1883, p. 518 = Ixodes putus (Pickard-

Cambridge, 1878).
flawdus 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 Janeiro = Amblyomma

dissimile Koch.
fiavipes Koch, 1835-1844, H. 39, Fig. 2 ; 1847, p. 21 (Germany) = /. vespertilionis

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. 19l=ricinus (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 Amblyomma. 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.

Condemned Species, etc. 285

gervaisi Lucas, 1847, p. x.civ = Aponomma gervaisi (Lucas), 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.
globulus Lucas, 1860, p. 538. Insufficiently described, may be an Aponomma.
gracilentus Lucas, 1845, p. 58, PI. I, Fig. 2 (o) = Hyalomma aegyptium (Linn.),

in Neumann, 1899, p. 286.
granulatus Supino, 1897. See Notes on Doubtful Species, p. 291.
herrerae Duges, 1887? p. 487; 1891, PI. VIII, Fig. 5 ($) = Amblyomma

cajennense Koch, 1844.
hexagonus Leach, in Salmon and Stiles, 1901, p. 467. See synonymy of I.

hexagonus var. cookei.
hexagonus var. inchoatus Neumann, 1901, p. 283 = canisuga Johnston, 1849,

p. 371.
hexagonus var. longispinosus Neumann, 1901, p. 283=7. 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, PL 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 (?)?= 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.
humanus Koch, 1844, p. 233 ; 1847, p. 104, PL XXI, Fig. 78 d)- Amblyomma

dissimile Koch ; see Neumann, 1899, p. 227 ; a o, from Brazil.
hydrosauri Denny, 1843, p. 314, PL XVII, Fig. 4 = Amblyomma hydrosauri

(Denny), in Neumann, 1899, p. 197.
iguanae (Fabr.) Fabricius, 1805, p. 354 (Acarus iguanae Fabr., 1794, iv,

p. 427) = Amblyomma iguanae.
imperfectus Neumann, 1899, p. 118. See Notes on Doubtful Species, p. 292.
indentatus Gamgee, 1869 — Booph'dus annulatus (Say), in Neumann, 1897,

p. 408.
indus (Linn.), Syst. nat., II, 1022 ; Fabricius, 1794, p. 428 (I. indus) ; 1805,

p. 355. Undeterminable.
inermis Neumann, 1901, p. 283. See Notes on Doubtful Species, p. 292.
intermedins Neumann, 1899, p. 132 ; 1904, p. 451=7. neumanni Nuttall and

Warburton, 1911.

286 Genus Ixodes

juvenis Neumann, 1899, p. 124. See Notes on Doubtful Species, p. 292.
kelloggi Nuttall and Warburton, 1907, p. 396 = 7. brunneus Koch, 1844.
lacertae Koch, 1835-1844, H. 39, Fig. 11 ; 1847, p. 22 (Germany) = ricinut

(Linn.), in Neumann, 1899, p. 112. " Ix. lacertae" in Pagenstecher, 1861.

p. 41, Fig. 22 ; may be the same.
laevis Neumann, 1899, p. 148 = acutitarsus (Karsch), 1880.
lagotis Gervais, 1849, p. 49. Insufficiently described ; may be a Haemaphysalis.
leachii Audouin, 1827, p. 428. (Savigny, PL IX, Fig. 9) = Haemaphysalis leachi

(Audouin).
leporis-palustris Packard, 1867, p. 67 = Haemaphysalis leporis-palustris (Packard).
lineatus (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 = Rhipicephalus sp. ? Undeterminable.
lipsiensis (Fabricius) Fabr., 1805, p. 354. {Acarus lipsiensis Fabricius, 1794,

pp. 427, 428) = young of ricinus or hexagonus ; see Neumann, 1899, p. 157.
lividus Leach, 1824, n, in Koch, 1844, p. 234 (undeterminable) = young of

ricinus or hexagonus in Neumann, 1899, p. 157.
Hindus van Beneden, 1873, XL. A purely nominal species from Vesperugo.

No description ; see Neumann, 1899, p. 160.
longipes Lucas, 1872, p. \xxiv = vespertilionis 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 9 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,

undetermi nable.
marginalis in Gervais, 1844, p. 242 = ? (Salmon and Stiles, 1901. Synonymy of

I. 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. i¥l) = Argas reflexus (Fabr.) Latreille, 1796.
marginatus Burmeister, 1837, p. 579 = 7. ricinus, fide Oudemans, 1896, p. 191

in Salmon and Stiles, 1901, p. 466.
marmoratus Risso, 1826, p. 183 = Dermacentor reticulatus (Fabricius) 1794, in

Neumann, 1897, p. 360.
maskellii Kirk, 1887, p. 66. Undeterminable, description useless. Found on

Albatross (Diomedia exulans), New Zealand.
megathyreus Leach, 1815, p. 398 = 7. ricinus (Linn.).
mixtus Moniez, 1896, p. 497 = A mblyomma mixtum Koch = Amblyomma cajen-

nense (Fabr.), in Neumann, 1901, p. 348 (Index).
moreliae L. Koch, 1867, p. 241 = Amblyomma moreliae (L. Koch), according to

Neumann, 1899, p. 258. Description insufficient to make it a good species,

however, in our opinion. Found at Brisbane on Morelia argus var.
fasciolata Jan.

Condemned Species, etc. 287

naponensis Packard, 1867, p. 65 = Dermacentor electus Koch in Neumann, 1897,

p. 366 ; but electus=D. variabilis (Say, 1821) fide Stiles, 1910, p. 29.
nigrolineatus Packard, 1867, pp. 66, 67 = Dermacentor nigrolineatus (Packard)

Banks, 1907, fide 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 Qeer) Latreille, 1804, p. 52 = Amblyomma americanum (Linn.) Koch,

1844 ;fide Salmon and Stiles, 1901, p. 475.
nigua Guerin = Amblyomma americanum (Linn.) Koch, 1844, in Neumann,

1901, p. 342 (Index).
obliquus Koch, 1844, p. 232 ; 1847, p. 99, PI. XX, Fig. 73 ( ? and its scutum).

Insufficiently described.
obscurus 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 = ? Aponomnia 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 part n'cmws (Linn.), says Neumann,

1904, p. 452.
ovatus Neumann, 1899, p. 112, 1904, p. 452 = 7. ricinus var. ovatus (Neumann).

Nuttall and Warburton 1911.
pollens Fabricius, 1805, p. 356. Undeterminable.
pallipes {Acarus pallipes Fabr., 1794, p. 426) referred to in Koch, 1835, H. 39,

10 ; 1847, p. 22 (Germany) = ? I. brunneus 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 " Parus major" = ? /. 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. 239 = Dermacentor reticulatus (Fabricius), in Neumann,

1901, p. 265.
pilosus var. howardi Neumann, 1908, p. 125 = 7. pilosus Koch.
plumbeus (Panzer) in Fabricius, 1805, p. 353 {Acarus plumbeus Panzer).

Undeterminable.
plumbeus Leach, 1815, p. 397 ; 1824, p. 11. Undeterminable=7. lividus Koch,

in Neumann, 1901, p. 348 ; 1899, p. 157 ; see lividus in this list. On

Hirundo riparia.
plumbeus Duges, 1834 c, PI. VII, Figs. 7-12 = Rltipicephalus sp., as clearly

indicated by the figures ( = 7 ricinus, according to Neumann's synonymy).

Also figured by Wagner, 1841, Figs. XII, XIII (unrecognizable).
plumbeus in Wheler 1899 = 7soc/es canisuga Johnston, 1849.

288 Genus Ixodes

poortmani Lucas, 1850, p. xli ; Belval, 1861, p. 97, PL = Amblyomma hebraeum

Koch, in Neumann, 1899, p. 266.
praecoxalis Neumann, 1899, p. 121=7. neumanni Nuttall and Warburton,

1911.
pulchellus Lucas, 1845, p. 61, PL I, Fig. 4 ; 1849, p. lxxx ; 1849, p. 582 ((J, good

figure of ca.pitn\\m>) = Amblyomma dissimile Koch, in Neumann, 1899,

p. 227.
punctulatus Say, 1821, p. 78 ; 1859, p. 21. Original description useless, said to

resemble "7 variabilis" = Dermacentor variabilis (Say, 1821).
punctulatus Canestrini and Fanzago, 1877-1878, pp. 115, 183 ; Canestrini, 1890,

p. 502. Doubtful if it is an Ixodes ; see Neumann, 1899, p. 160. From

Europe.
pustularum Lucas, 1866, p. lvii = Ixodes ricinus (Linn.), in Neumann, 1899, p. 113.
pygmaeus Koch, 1844, p. 233 ; 1847, p. 107, PL XXII, Fig. 81 ( ? ), from Brazil

and Mexico. Types examined by Neumann (1901, p. 289), and found to

be Amblyomma nymphs, of undetermined species.
quinquestriatus Fitch, 1870, p. 366 = Dermacentor variegatus Marx and Neumann,

in Neumann, 1901, p. 266.
reduvius Audouin, 1832, nee Linnaeus ((J), p. 422, PL XIV, Fig. 4 = 1. hexa-

gonus Leach, 1815.
reduvius Hahn, 1834, p. 62, Fig. 152 = 7. ricinus (Linn.), in Neumann, 1899,

p. 112.
reduvius de Geer, in Megnin, 1880, p. 126 = 7 ricinus (Linn.).
reduvius (Linn.) = ricinus (Linn.), in Neumann, 1901, p. 348, and numerous

other authors, this incorrect, see p. 143.
reflexus (Latreille) Fabricius, 1805, p. 352 = Argas reflexus (Fabr., 1794, p. 426)

Latreille, 1796. (See our Part 1, p. 22.)
reticulatus Latreille, 1804, p. 157 ' = 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 ocelala, in Spain.
rhinocerinus Denny, 1843, p. 313, PL XVII, Fig. 3 = Dermacentor rhinocerotis (de

Geer), in Neumann, 1897, p. 370.
rhinocerotis Fabricius, 1805, p. 351 = Dermacentor rhinocerotis (de Geer), 1778.
rhinocerotis Gervais, 1844, p. 246 (Walckenaer and Gervais) = Dermacentor

rhinocerotis (do Geer) 1778, in Neumann, 1897, p. 370.
ricinus Megnin, 1880, p. 129 nee Latreille = 7 hexagonus Leach, 1815; see

Neumann, 1899, p. 129.
rostralis Moniez, 1896, p. 49Q=Oonixodes rostralis Duges, 1888, p. 129 =

Haemaphysalis leporis-palustris (Packard) in Neumann, 1901, p. 348 (Index).
rubiginosus (Kolenati), Neumann, 1899 (Dermanyssus rubiginosus Kolenati,

1857, p. 20, PL 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 = 7. ricinus (Linn.).
sanguineus Latreille, 1804, p. 157 = 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. Atlas, PI. XXXII, Fig. 1; XXIII, Fig. 2 =

Hyalomma 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 = 7. ricinus var. scapularis (Say)

Nuttall and Warburton, 1911. Original description useless.
scapulatus Megnin, 1880, p. 132. Undeterminable.
sciuri Koch, 1835-1844, H. 35, 8 ; 1844, p. 232 (listed); 1847, p. 21 (Germany)

= 7. ricinus (Linn.) os, types examined by Neumann, 1901, p. 282.

I. sexpunctatus Koch, 1835-1844, H. 39, 5, 6 ; 1844, p. 232 (listed) ; 1847,

p. 22 = /. hexagonus Leach; see Neumann, 1901, p. 283, who examined

the types (os). Canestrini and Fanzago, 1877, p. 114, described a ° (?)

as sexpunctatus, but their description is useless.
siculifer Megnin, 1880, p. 132 = 7. vespertilionis (Koch) 1844.
spinosus Neumann, 1899, p. 146= fuscipes Koch, 1844.
b-slriatus Fitch ; see quinquestriatus.

sturni Pagenstecher, 1861, p. 40 = ? 7. brunneus Koch, 1844 (larva).
sulcatus Koch, 1844, p. 233; 1847, pp. 22, 108, PI. XXII, Fig. 82 (°) = 7.

ricinus (Linn.) os. Types examined by Neumann, 1901, p. 282.
sylvaticus Gervais, 1778, p. 162, PI. 38, Fig. 7 = Amblyomma sylvaticum (de

Geer), in Neumann, 1899, p. 274. On tortoise, Cape of Good Hope. May

attack man.
testudinis Conil, 1877, p. 25 = Amblyomma testudinis (Conil), see Neumann,

1905, p. 234.
testudinis Supino, 1897, p. 11= Amblyomma supinoi Neumann, 1905, p. 234.
testudinis Leydig, 1855, p. 382, PI. XV, Fig. 51, also p. 395 ; 1857, p. 111 =

Hyalomma syriacum, no doubt.
tkorackus 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 = auritulus Neumann, 1904, p. 450.
trabeatus Audouin, 1832, p. 420, PI. XIV, Fig. 3 (?) = /. ricinus (Linn.), in

Neumann, 1899, p. 112.
trachysauri Lucas, 1861, p. 12b = Aponomma trachysauri (Lucas).
transversalis Lucas, 1844, and in Murray, 1877, p. 195 = " Neumanniella

transversalis (Neumann)" Lahille, 1904 = 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. lxxvii = 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 = 7. vespertilionis Koch.
undatus (Fabricius) Fabr., 1805, p. 353 (Acarus undatus Fabricius, 1794,

p. 427). Undeterminable.
"unipictus Packard," of Verrill, 1870, p. 118 (lapsus for unipunctata (q.v.) fide

Salmon and Stiles, 1901, p. 476).
unipunctata Packard, 1867, p. Q& = Amblyomma americanum Koch, in Neumann,

1899, p. 209.
uriae White, 1852. Nominal species, found on Uria troile at Baffin's Bay.

Description and figure inadequate; may = ? I. putus.
varanensis Supino, 1897, p. 13, PI. II, Figs. 1-5 = Aponomma gervaisi (Lucas),

in Neumann, 1899, p. 182.
varani L. Koch, 1867, p. 241 = Aponomma decorosum (L. Koch), in Neumann,

1899, p. 194.
varani Lewis 1892, p. 10, PL I = Aponomma exomatum (Koch) 1844.
variabilis Say, 1821, p. 77 ; 1859, p. 21. Original description useless, said to

resemble erraticus and punctulatus. Listed, however, by Banks, 1895, p. 433,

as occurring at Fort Collins = Dermacenlor variabilis (Say) Banks, 1907, fide

Stiles, 1910, p. 29.
variegatus Lucas, 1852, p. lxxxviii = Ambly 'omnia 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.
vibrant (Fabr.) Fabricius, 1805, p. 356, No. 28 (1794, p. 428, 20: Acarus

vibrans). Undeterminable.
viperarum Koch, 1844, p. 234 ; 1847, p. 102, PI. XX, Fig. 76 (larvae). From

Greece, undeterminable.
vulgaris Fabricius, 1805, p. 352 = 7. ricinus or hexagonus.
vulpis Pageustecher, 1861, p. 40, PI. I, Figs. 12, \Z = hexagonus Leach, 1815.
walctenaeri 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 Eschatoee-

phalus by Neumann).
brevipes Kolenati, 1856, p. 21=1. vespertilionis Koch.
exaratus Kolenati, 1856, p. 22 ; 1860, p. 757, PL I, Fig. 2=7. vespertilionis Koch,

vide Neumann, vnr. 1910, p. 191, who examined type, a nymph.
flavidus Kolenati, 1856, p. 21=7 vespertilionis Koch.
flavipes Kolenati, 1856, p. 21=7. vespertilionis Koch. (Neumann 1910, p. 192,

states the types were not adults.)
hispidulus Kolenati, 1856, p. 21=7 vespertilionis Koch.
kochi „ „ p. 21 ; 1860, p. 573, PL II, Fig. 4 = 7 vespertilionis

Koch, (J.
nodulipes Kolenati, 1860, p. 576, PL I, Fig. 3 = probably 7 vespertilionis Koch.

Notes on Doubtful Species 291

NOTES ON DOUBTFUL SPECIES OF IXODES.

Ixodes aequalis Banks, xi. 1909, p. 276 ; 1910, p. 6, PI. Ill, Fig. 23 ( $? capitulum
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. I., nearly diamond shaped, antero- and postero-
lateral borders subequal, many uniform punctations, lateral carinae distinct.
Spiracle circular. Capitulum 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, Qoba dol, etc., Krain, Austria), the hosts probably being
bats. Insufficiently described ; said by Joseph to have shorter legs than /.
vespertilionis.

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 I. 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. l-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 (J to
be either a ? 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 subtriangular, porose areas rounded, divergent.
Hypostome long, narrow, lanceolate, 3 [3, 10-11 sharp teeth per file; digit

292 Genus Ixodes

slender (110 /x I.), 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 Sciurus gordoni, S. rufigenis, S. striatus and Felis tigris, at
Bharuo, 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 2-1x1-2 mm. Scutum oval, glossy (0-56x0-51 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.
Capitulum (033 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 : LHdelphys 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 <Js identical with Ixodes
ricinus were found in the same lot, but differed 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 1-1 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 Species 293

superficial, almost forming the lateral border. Venter glabrous, or almost so ;
sexual grooves straight, divergent, attaining the posterior border ; anal grooves
parallel, united in an arc in front. Spiracle large, circular. Capitulum
0-35 mm. I., base rectangular, broader than long ; hypostome elongate, lanceo-
late, 2 1 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 apj>ears 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 trianguliceps 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.

Introductory 294

Outline of the life-history in Ixodes 295

Ixodes ricinus 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 .... 301

The influence of season and local conditions . . . 303

Copulation 303

Oviposition 308

Belation 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 anguslus .......... 315

Ixodes pilosus 316

Ixodes canisuga .......... 316

Ixodes pntiis .......... 317

Appendix I. On the process of copulation in Ornithodorus

moubata 318

Appendix II. On the adaptation of Ticks to the habits of

their hosts 324

Introductory.

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

hexagonus 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 pilosus experimentally; and Hadwen, in British
Columbia, has made partial raising experiments on angmtus. Of the
remaining species whose various stages are known {tenuirostris, ves-
pertilionis and loricatus), 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 </", $, 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.

J, % and o „ „ „ 4

</" and % „ „ „ 6

o is >> » " »>

+ >. » » «" »

$ and O, are „ „ 4 „

$, o and L, „ „ „ 6

It follows that these species are known by 21 Js, 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 feeding1. 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 ground2. I have frequently observed females

1 Although I. ricinus <r may attacli 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."

Biology of I. ricinus 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 (canisuga, angustus, putus).
Having fallen upon the grouud, 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, they 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 egg 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 days1, 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.

R. 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

Experiment I. 10. x. 1905. Many larvae were placed on a hedgehog in a

tick-proof cage.

13. „ 68 larvae dropped off' gorged.

14. „ 365 „ „ „ „
J6- „ 60 „ „ „ „

18. i. 1906. Nymphs began to emerge from the larval skins.

3. vi. „ Thirty of these nymphs were placed on a hedgehog.

7. „ 10 nymphs dropped off gorged.

"- 1) H !! >] h »

Experiment II. 18. x. 1905. Many larvae were placed on a hedgehog.

23. „ 250 larvae dropped off gorged.

24. „ 26 „ „ „ „
18. I. 1906. 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 nymphs for 4-5 days, when raised on hedgehogs in the laboratory
at a temperature of about 15° C.

These results are in agreement with the statement by Kossel,
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. 145) reports that he once placed a
hungry </" and % upon his arm, with the result that the £ remained
attached for only eight hours whilst the $ remained upon him for eight
days. Kossel and his colleagues once saw hungry Js readily attack a
guinea-pig upon which they were placed ; they state that the $ renmins
attached for a week. According to Sainson (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 frequently found in 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. vin. (Bertkau, 1881, p. 148).
„ 8 „ „ 25. v. to 19. vii. at 65° F. (Wheler, xn. 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.

Nymph* 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. vn. (Wheler, xn. 1899).

(in winter) 7 „ (Kossel, Schiitz, Weber and Miessner, 1903, p. 43).

(kept cool) 12 „ ; 14, vin. 1899 to 4. vm. 1900 (Wheler, 1902),

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 greatly
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. vin. 1898 were alive and active on 7. vi. 1899 :
10J 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 cherries, 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 l£ 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.

Allowing 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 ...
Nymph
Adult ?

13 weeks (l£ free, \ on host, 11 gorged).
13 „ (1J „ i „ 11 „ ).
H m (1* „ 2 „ 4 „ ).

Days required

Shortest period

Longest period

8

27

42

49-252

10

300-570

3

6

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 five 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 :

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 ...

Nymphs harden and wait to attack host
No. 2 after

Nymphs remain upon the host for

(Nj mphs, having abandoned the host, under-
go metamorphosis.)

Adults issue from the nymphal skin after

Adults harden and wait to attack host No. 3
after

Adults ( ? s) remain upon the host for ...

Fecundated ? abandons the host

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

28

84-140

10

200-540

3

5

56

210-360

10

450-810

8

14

Total = 178 days.

1345-2724

Biology of 1. ricinm 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, Schiitz, 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 tnay be captured in a free
state at the end of April, proving that all stages may hibernate. They
are readily captured by dragging a woollen cloth 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 coupling1) 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. Miiller (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 <f mouthparts into the vulva, hereinafter described, con-
stitutes only a stage in the process of copulation ; see pp. 318 et teq.

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
scienza1." Gen6 (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
(1835-1844, 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. Megnin (1878, p. xcii)
affirms that the males possess a penis (!) which is directed toward the
vulva by the male mouthparts. Claus (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 unfed adults and
between replete females and males either upon the host or otherwise.
He considered that the female became impregnated through the male

1 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. ricinus

305

mouthparts. Wheler sent specimens to Lewis (1900, p. 382) 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 £ 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 & in ventral and lateral aspects, showing at
A, A1, 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, as 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 ^s 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 </ 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 Js may

Biology of I. ricinus 307

copulate and that $s may copulate several times; moreover that $s
will only attach themselves to a host when fertilized, i.e. during or after
copulation. The last statement requires confirmation. This author
(p. 217) denies that the penetration of the </ mouthparts into the ?
genital orifice (what we term coupling) represents coitus. On examining
a J which had been coupled for an hour with a £, her gonoduct was
found to be empty and no spermatophore was found in the £ excretory
ducts. That coitus should take place by means of the mouthparts of
the </" is difficult to understand because (a) the <$ mouthparts cannot be
brought in contact with the </" genital orifice so as to receive the sperm
therefrom ; (b) the structure of the £ mouthparts shows no adaptation
for this function ; and (c) the £ mouthparts are immediately introduced
into the vulva when the sexes meet. On one occasion what appeared to
be 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 bis 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 Gene' 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 sperm atophores 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 facilitated 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 one 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 Gene" (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 bringen 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 americanus " 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.
Muller (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

Biology of I. ricimis

309

the vagina is everted and protruded forward and the egg 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 Genu's organ, in honour of its discoverer, completely envelops the egg
as it emerges from the everted vagina. According to Gene- it takes

f"^^^C

Fig. 284.

Fig. 285.

.^^

Fig. 286.

Figs. 284-287. Oviposition in Ixodes ricinus (Wheler, 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 Gene'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 ricinus.

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 Genu'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 egg 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 covering the eggs with the sticky
secretion, which may serve to harden the chitinous envelope of the egg,
that the organ may convey spermatozoa in some way from the female's
uterus to the egg as she is laying. He did not, however, pretend to explain
how fertilization actually takes place1. The ovipositor and Genu'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

1 Railliet (1895, p. 706, Fig. 480) reproduces a figure of an ovipositing ricinus from
Gene (1848), and briefly refers to the process and to the everted vagina or oviduct serving as
an ovipositor or " oviscapte"; he refers to Geno's organ as the "bourse seminale de Gene,"
or "vessie bilobee." Wheler (1899) calls it the " dorsal gland." Samson (1909, p. 223)
calls it the " Subscutalblase." Dutton and Todd (1905 b, p. 17) describe and figure the same
structure in Ornilliodorus moubata as a " pulmonary sac" and Brumpt (1910, p. 523, Fig.
363) calls it the " cephalic gland," following Christophers (1906, p. 95), who gave this name
to the structure in O. savignyi. None of these names adequately describes Genu's organ.
We give pre-eminence to Gene's observations because they have been generally ignored by
writers on the subject.

TICKS

PLATE VII

Ixodes ricinus. 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. x4. (Original, L. E. Robinson, phot.)

Biology of I. ricinus 311

when Gene's organ was not pricked but simply made to retract itself by
being touched with a small rod. The eggs which then appeared had dried
up in 6-8 hours, whilst eggs laid normally remained fresh for weeks1.
Wheler (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 are 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
egg, 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 Genu'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 ricinus, we glean the following notes on the subject from the
literature :

Gene" (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 (xn. 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, Schlitz, 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-Valerio (1908, p. 611, with figure) illustrates, by means of a

1 Schleohtendal (1891, p. 11) givea a very imperfect description of oviposition observed
in the case 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 Gene'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 Haemaphy sails, 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 ir 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. ricinus 313

Ireland. Kossel found ricinus in N. Wales in redwater 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 of 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 /. 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
Eaymondaud, 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

n. i. ' 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 albus 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.
Megnin (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,
Ric. vitirin., cited by Megnin, 1892) observed a cyst at the end of a
dog's ear which contained a tick supposed to be ricinus. Vau 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 ricinus 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.

Biology of I. 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 M£gnin (1867, R4c. veterin., redescribed
1892, p. 56 ; in the earlier paper he referred the ticks to a uew 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 Js taken from
the same dog, was shortly after found coupling with 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 ^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 £ beneath a % upon a host
but the sexes were not coupled. He has only captured 8 or 9 Js
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 J1 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 Js 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 Larvae 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 Gowanburn, on the North Tyne, and all the males we
otherwise possess have come from the nests of the sand-rnartiu or bank-

Biology of I. putus 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 7. plumbeus ? in 1899.

Through the courtesy of Mr. R. F. L. Burton, of Longner Hall,
Shrewsbury, who kindly supplied me with 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 ricinus. 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 (xn. 1899) kept the unfed larvae of this species alive for
ten months (9. x. 1898-vin. 1899), after they had hatched out from
eggs laid 4. vm. 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
known 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 Ornithodorus mignini 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 (1906, 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 Bucktou, 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 moubata. 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 moubata, that is, impregnation takes place by
means of spermatophores.

We have repeatedly observed the process in moubata in Cambridge,
and the description which follows is based on the study of several pairs
in copulation. When a male and female moubata, 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] Copulation 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]. Ornithodorw 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 Appendix 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 be 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 35 mm.
in length and about l-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 1"6 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 mouthparts 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. rnoubata 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 " (prospermiums) 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. rnoubata 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
Haemaphy salis, Dermacentor, Rhipicentor, Rhipicephalus, Boophilus,
Margaropus, Hyalomma, Amblyomma 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 HOSTS1.

By GEORGE H. P. 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 recording 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 habits2.

General considerations regarding 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

1 Read before the Cambridge Philosophical Society Meeting, 20 February, 1911.

2 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 reflexus and Argas vespertilionis, 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 nymphal stages in Argasidae (one only in Ixodidae),
and the nymphs, with one exception, Omithodorus megnini, are 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 !j! Ixodidae, though possibly in Js).

The nymphs1 and adults of Argasidae are rarely 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 Amblyomma,
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 the 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
nymphal1 and adult stages; the females lay their eggs in and about
1 Omithodorus megnini excepted.

:!!'<;

Appendix 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. reflexut

A. vapertilion.it

Hosts and habitat etc. of Tick

Birds : fowls, clucks, 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 Tickt, Part I, p. 39, where the bats are listed. A variety infests penguins at
Queenstown, Cape Colony.

[49]

Adaptation of Ticks

327

Argasidae
A. brumpti

0. savigmji
O. moubata

0. turicata

0. talaje

O. talaje var. ca-
pensis

O. pavimentosus
0. tholozani ...
O. lahorensis ...

0. megnini

Hosts and habitat etc. of Tick

Porcupine : found infesting dusty burrows of
Hystrix 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
chiefly in the ears.

Geographical

distribution

(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 egg, 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 Appendix II [50]

The larvae of remain attached to the host for

A. persicus 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)1.

0. megnini 5 days (but they stay on as nymphs upon the host).

The first stage nymphs of

A. persicus 1J-2 hours (Nuttall).

0. moubata 10 minutes-1 hour (Nuttall).

O. 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).

O. moubata 20 minutes-2 hours (Nuttall).

O. coriaceus 45 minutes-1^ hours (Nuttall).

O. turicata Feed more rapidly than the last (Nuttall).

O. tholozani 30 minutes (Me'gnin).

O. lahorensis 25 minutes-2J hours (Nuttall).

O. savignyi Feed like O. 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. moubata and savignyi.

„ 3. O. 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 record8 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 Boumania
(vide Ticks, Part I, p. 27), hut I feel warranted in doubting the statement.

[51] Adaptation 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 reflexus 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
bats1. 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. moubata 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 moubata)
as does the adult, i.e. in 10 minutes to one hour. Ornithodorus moubata
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
moubata. 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.

1 See footnote, p. 326.
N. i. • 22

330 Appendix 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 575x450/*. 5. Boophilus decoloratus 625 x 400 ix.

2. Hdemaphysalis punctata 570 x 380 /x. 6. Amblyomma hebraeum 625 x 525 /*.

3. Hyalomma aegyptium 550 x 425 p. 7. Argas persicus 625 x 575 /x.

4. Rhipicephalus evertsi 550 x 400 ix. 8. Ornithodorus moubata 950 x 750 /*.

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 which 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. brumpti, 0. turicata and 0. pavimentosus
may prove, on investigation, to have a life-history similar to that of
0. movhata 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. lahorensis will be worked out in India. We are studying 0. turicata
and 0. talaje var. capensis.

Note on the structure of the Hypostoines 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, reflexus and vespertilionis, where the larvae attach them-
selves to the host for some days, or longer, the larval hypostome is

Fig. 2901.

Fig. 291.

Fig. 292.

Fig. 293.

Fig. 294.

Fig. 290. A. persicus, larva (similar in larva of A. reflexus)

Fig. 291. A. vespertilionis, larva.

Fig. 292. A. persicus, adult.

Fig. 293. A. reflexus, adult.

Fig. 294. A. vespertilionis, nymph.

1 Figs. 290-297 are reproduced from Ticks, Part I.

[55]

Adaptation of Ticks

333

Fig. 295. Fig. 296. Fig. 297.

Fig. 295. 0. navignyi, adult. Fig. 296. 0. moubata, adult.

Fig. 297. 0. moubata, 1st stage nymph.

Fig. 298'.
Fig. 298. 0. megnini, larva.

Fig. 299'.
Fig. 299. O. megnini, nymph.

1 Figs. 298, 299 are reproduced from Salmon and Stiles, 1901, Figs. 99 and 102
(17(A Ann. Rep., V.A.I., V. S. Dept. Agricult. and from Ticks, Part I).

334 Appendix 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 hypostome. In 0. savignyi
and 0, moubata, 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
maiked 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, nigricans, fossulatus, acutitarsus, 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 eyidence 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.

II. 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 Cants 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 (b). 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 hypostome1 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 Ixodes will be discussed presently.

336

Appendix II

[58]

I. Species of Ixodes of which both J and ? are recorded as
occurring upon the host.

(a) Species in which the sexes occur in copula upon the host.

Number of Geographical

Host's

times found distribution

Speclea

habits

Host

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

+

,, oapreolus

4

+

, , dama

1

-63

+

Capra hircus

2

+

Equus caballus ...
Homo sapiens

6
2

+

Felis concolor

1

+

Genetta sp.

1

+

Lepus europaeus...

2;

0

Erinaceus europaeus

2'

0

Mustela putorius

1

0

,, erminea...

1

0

Meles taxus

1

0

Mus decumanus ...

1

-10

0

"Mouse"

1

0

Myoxus sp.

1

0

Talpa europaea ...

1

0

Ground squirrel ...
Larvae and nymphs

1>

on

lizards (10) and birds (5)

of different species.

I. ricinus var. scapu-

+

Cattle ...

N. America, Costa

laris ( 3 ? known)

+

Deer
Dog
Man

Eica.

I. ricinus var. ovatus

+

Horse ...

Japan.

(only 2 ? s known)

±

Dog

J. ricinus var. califor-

+

Black-tail deer ...

N. America.

nicus

0

Grey fox

I. pilosus ...

+

Capra hircus

8

S. Africa.

+

Bos taurus

2

+

Equus caballus ...

3

+

Sus scrofa domestica

2

+

Ovis aries

1

+

Duikerbok

1

+

Tragelaphus sylvaticus

1

=fc

Canis familiaris ...

1

[59]

Adaptation of Ticks

337

Host's

Species

habits

Host

I. piloms . . .

+

Felis domesticus ...

+

Viverra civetta ...

+

Felis paidus
Homo sapiens

I. pilosu* var. howardi

+

Cat

( cf ? known)

0

Erinaceus frontalis

0

Khinolophus sp.

/. boliviensis

?

Icticyon (Speothos) venati-
eus (Canidae)

I. schillingsi

+

Colobus caudatus (monkey)

I. rams

+

Bos taurus

+

Felis pardus

±

Canis familiaris

0

Herpestes ichneumon

0

Hyrax sp.
Homo sapiens

I. itgandanus

+

Ovis aries

0

Aulacodus sp.

" Large rodent "(?Aulaoodus)

I. ugandanus var.

dja-

?

Genetta suahelica

ronensis ( i ? known)

0

Procavia brucei

Number of

times found

thereon

1

1

1

I

Geographical

distribution
of tick

S. Africa.

S. Africa.

Bolivia.

E. Africa.

E. & W. Africa.

E. & W. Africa.

E. Africa.

(b) Species in w

hich the sexes have been found toge

the1)

• upon the host.

I. anguslus

0
0
•>

0
0

Sciurus hudsonius douglasi
Tamias townsendi (Sciuridae)
Lepas dalli

Neotoma occidentalis (Muridae^
" Mouse "

N. America.

I. tenuirostris

0
0
0
0
0
0
0
0

Evotomys glareolus
Arvicola amphibius

,, pratensis
Microtus agrestis

,, arvalis ...
Sorex vulgaris

„ minutus ...
Mus miniatus

3
2
1
2

Europe.

I. minor

0

Hesperomys sp. (Muridae)

Guatemala.

I. cavipulpu-i

+

Cynocephalus babuin
Homo sapiens

Africa (Bhodesia
<fe Angola).

I. rubicundus

+

+

Ovis aries
Capra hircus

S. Africa.

■22 -•-)

338

Appendix II

[60]

Species

Host's
habits

Host

Number of Geographical

times found distribution

thereon of tick

I. rubicundiis var. Urn-

+

Sheep ...

1 C. Africa (Congo).

batus ( ? only known)

+

Goats

7. holoq/clus

+

Bos taurus

Australia, India.

+

Macropus sp.

±

Canis familiaris

0

Sciurus variabilis

0

Phascogale penieillata (M
supial tree shrew)

!ar

I. loricatug

+

Ateles melanochoerus

S. America.

0

Didelphys aurita

0

„ opossum

0

sp.

0 Microdidelphys sorex (o & L1.)

occurs as a parasite on small mammals which burrow ; these burrows
are, however, near the surface of the ground and liable to be 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.

1 o and h signify that nymphal 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.

Species
I. hexagonus

I. hexagonus var.
cookei ( <r ? known)

Geographical

distribution
of tick

Europe, N. Africa,

N. & S. America.

N. America.

Number of
Host's times found

habits Host thereon

0 Erinaceus europaeus ... 9'

0 Mustela erminea ... ... 8

0 ,, vulgaris... ... 7

0 ,, putorius... ... 6

0 „ furo ... ... 2

0 Lutra vulgaris ... ... 1 riO

0 Meles taxus ... ... 1

0 Lepus cuniculus ... ... 1

0 Myopotamus coypu ... 1

0 Sciurus sp. ... ... 1

0 Canis vulpes ... ... 3 '

± Canis familiaris ... ... 3

+ Ovis aries ... ... 1

+ Sus scrofa ... ... 1

Homo sapiens ... ... 1

0 Lutra ...

0 Mustela vison

0 Spermopliilus

0 Weasel

0 Porcupine

0 Marmot

0 Pocket gopher

± Dog

+ Cat

+ Sheep

± Canis familiaris ... ... 12

0 ,, vulpes ... ... 2

0 Cotile riparia (bank-swallow) 3

0 Mustela furo ... ... 1 J-21

0 Meles taxus

0 Sciurus ...

0 Talpa europaea (larva)

+ Equus caballus ...

+ Ovis aries

I.putus. On marine birds only ( $ only found in nests or beneath rocks and soil near neBts).

I. canisuga

Europe, N.
erica.

Am-

0 Phalacrocorax verrucosus

0 Pygosceles taeniatus

0 Spheniscus magellanicus

0 Fratercula arctica

0 Penguin sp.

0 Uria troile

0 Sea-gull sp.

0 Sula bassana

0 Fulmarus glacialis

0 Cormorant sp.

+

Europe, N. * S.
America, Asia,
Australasia.

Man (occasionally, see text, p. 340)

340

•

Appendix II [62]

Species

Host's
habits

Number of Geographical
times found distribution
Host thereon of tick •

I. vespertilionis.

Bats only ( t

found in oaves etc. inhabited by bats).

0
0
0
0
0
0

Plecotus auritus ... ... Europe, Africa,

Ehinolophus hipposideros Australia.

„ ferrum equinum

,, euryale

„ hippocrepia

,, blasius.

0

,, clivosus

0
0

Vesperugo pipistrellus
„ tricolor.

Ixodes putus 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 /. hexagonus1, 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 bats, 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 £ does not feed, and that it nevertheless
fertilizes the female. The same may hold for certain Ixodidae r/s.
The matter requires further investigation. The sexes of canisuga and
putus have often been observed in copula apart from the host, and it
appears probable that this is the usual occurrence in nature1.

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 known1

Species

Host's
habits

Host

Geographical

distribution

of tick

I. acuminatus

0

Mus agrarius (only 2 ? s known)

Italy.

I. dentatus

0

Rabbit (1 ? known)

N. America.

I. diversifossus

0

Procyon lotor (2 ? s recorded)

N. America.

I. fuscipes

0

Dasyprocta aguti (?Pelis pardalis)

C. & S. America.

I. sculptus

0

" Rock squirrel "

N. America.

I. spinicoxalis

0

Mu stela flavigula (4 ? s known)

Sumatra.

I. marxi

0

Red squirrel and fox (fow ? s known)...

N. America.

I. rubidus

0

Bassaris astuta (1 ? known)

Mexico.

I. nitem

0

Mus macleari (2 ? s known) ...

Christmas Island,
Pacific Ocean.

I. bicornis

+

Felis concolor, F. onca, Homo sapiens
(may = diversifossus, only 3 1 s known)

Mexico.

I. auritulus

0

Trupialis militaris and another bird.sp.?

S. America.

I. brunneus

0

Birds: many species

Europe, N. Ame-
rica.

I. caledonicus

0

Domestic pigeons ...

Scotland.

I. simplex

0

Rhinolophus ferrum equinum, Vespertilio

China.

I. texanus
I. stilesi

0

+

Grey squirrel, Procyon lotor (few ¥ s

known)
Padua humilis (Cervidae) ; (14 ? s

found once)

N. America.
Chili.

I. neumanni

0

Marine birds (10 ? s known) ...

New Zealand.

I. percavatus

0

Marine birds (presumably) ...

Tristan d'Acunlia
Island.

I. percavatus var.
rothschildi

0

Marine bird (puffin)

No locality given.

I. lunatus

0

Hallomys audeberti (1 ? known)

Madagascar.

I. ornithorhynchi

0

Ornithorhynchus (found 5 times)

Tasmania.

I. tasmani

0

Probably birds (Verreaux coll.)

■i

I. fecialis

0

Dasyurus geoff royi (1 ? known)

Australia.

I. fecialis var.
aegrifossus

0

Opossum (3 ? s); Paramelesobesula(l ? )

»»

I. anstraliensis

±

Canis familiaris (found once)

»>

0

Bettongia lusueuri (marsupial rat-kan-
garoo), (found once)

I. vestitut

0

Myrmecobius fasciatus (Dasyuridae),
(found once)

Diemenia superciliosa (snake), (found
once)

»»

I. signatus

0

Phalacrocorax

Asia, N. America.

1. unicavatus

0

Fhalacrocorax graculus and P. carbo

Great Britain.

1 Nymphal and larval stages are known in many of these species.

342 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 (&)),
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, schilling si, pilosus and boliviensis.
The hypostomes in the males of the first five species are characterised
by prominent basal teeth; boliviensis has a well-armed hypostome.

I (b). In seven species (cavipalpus, rubicundus, minor, Icricatus,
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, putus 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 («)) 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 (&)) will yet be
found in copula when they have been more carefully observed. In
Group I (a) and (b) 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]

Adaptation of Ticks

343

Figures 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

Pigs. 300, 301 represent the hypostomes of I. ricinus i and ? . That of the <? may be
taken as the type of a hypostome with large basal teeth as observed also in the males
of rasus, ugandanus, schillingsi, rubicundus, cavipalpus and pilosus, the basal teeth
being less prominent in the last two species mentioned. In boliviensis <f the basal
tooth, whilst prominent, is much less developed.

Pig. 302 represents the hypostome of I. minor s , with well-developed outer files of teeth.
(The hypostome of tenuirostris and loricatus approximate to the type of structure
found in the females.)

It is poorly but relatively better

Fig. 303 represents the hypostome of I. holocyclus <r .
armed than that of canisuga which follows.

Fig. 304 represents the hypostome of I. canisuga i , poorly armed, and much emarginated
distally. That of hexagonus is similar but not emarginated.

Fig. 305 represents the hypostome of I. putus l , 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 s 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, putus and vespertilionis males
that the capitulum is much smaller 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 hosts1.

A great many males belonging to Haemaphysalis, Dermacentor,
Rhipicephalus, Amblyomma (and Aponomma) are known, these genera
of Ixodidae being rich in species. The males of Boophilus, Hyalomma,
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 are chiefly 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 hexagonus 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.

Collections which include
the Types

Stages of each

species which

are known

Page

Toulouse, 2 ? a

?

161, 341

Berlin1, 1 ?

?

202

Washington (a), 1 ? ;
Cambridge, l o l

i ? O L

195, 315, 334 et seq.

Paris (a), 3 ? s;

Toulouse, 1 $ , co-type

?

187, 341

Cambridge, 2 ? s ;
Toulouse, 1 ? , co-type

?

250, 341

Toulouse, 3 ? s

?

186, 341

Toulouse, 1 i , 1 $ , 1 o

i ? 0

166, 334 et seq.

Berlin, 1 ? (in balsam)

$ O L

189, 341

Cambridge, 1 $ , o l
Edinburgh, 1 ? , o l

9 0 L

198, 341

Lost, but substitutes in
Cambridge : J ?ot;
Toulouse

i ? 0 L

209, 316, 334 et seq.

Cambridge, i s, ? s

<? ?

192, 334 et seq.

Leyden, 1 i

<?

233

Berlin, 1 i

t

270

Washington (6), 1 ?

?

162, 341

Washington (e), 2 ? s

?

163, 341

Cambridge, ?

?

248, 341

Cambridge, $

?

250, 341

Berlin, 1 ?

?

201

Berlin, ? ; Toulouse, o h

? O Ii

169, 341

Calcutta, 1 <f ;
Cambridge, 1 i

i

203

London, ?

cf ? O L

177, 330, 334 et seq

Harvard, ?

?

183, SSietseq.

London, Paris (o),
Toulouse, Berlin, ? s,
Hamburg

J ? O

235, 334 et seq.

Paris (a), 1 ?

?

208

Names of Species,
Authors ana Date

acuminatus Neumann, 1901

acutitarsw (Karsch, 1880)

*angustus Neumann, 1899

auritulm Neumann, 1904

*australiensis Neumann, 1904

bicornit Neumann, 1906
boliviensis Neumann, 1904
*brunneus Koch, 1844
*caledonicus Nuttall, 1910

*canisuga Johnston, 1849

*cavipalpw Nuttall and War-
burton, 1907
cortiifer Neumann, 1908
coxaefurcatus Neumann, 1899
dentatus Marx, 1899, in Neu-
mann, 1899
diversifo'ssus Neumann, 1899
*jecialis Warburton and Nuttall,

1909
*feclalis var. aegrifossus War-
burton and Nuttall, 1909

fossulatus Neumann, 1899
fuscipes Koch, 1844
*gigas Warburton, 1910

»hexagonus Leach, 1815
*hexagonut var. cookei (Packard,

1869)
•holocyclus Neumann, 1899

japonensis Neumann, 1904

1 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.

Index to Valid Species

347

Names of Species,
Authors and Date

*loricatus Neumann, 1899
*loricatus var. spinosus Nuttall,
1910

lunatus Neumann, 1907

marxi Banks, 1908

minor Neumann, 1902

neumanni Nuttall and War-
burton, 1911

nigricans Neumann, 1908
*nitens Neumann, 1904

ornithorhynchi Lucas, 1845

percavatus Neumann, 1906
*percavatus var. rothschildi Nuttall

and Warburton, 1911
*pilosus Koch, 1844

pilosus var. howardi Neumann,

1908
*pratti Banks, 1908
*putm (Pickard-Cambridge, 1876)

*rasus Neumann, 1899
*ricinus (Linnaeus) Latreille, 1804
ricinus var. californicm (Banks,

1904)
ricinus var. ovatus (Neumann,

1899)
*ricinus var. scapalaris (Say, 1821)
•rubicundus Neumann, 1904

rubicundus var. limbatus Neu-
mann, 1908
rubidus Neumann, 1901
*schillingsi Neumann, 1901

sculptus Neumann, 1904
*signatus Birula, 1895

simplex Neumann, 1906
spinicoxalis Neumann, 1899
stilesi Neumann, 1911
tasmani Neumann, 1899
*tenuirostris Neumann, 1901

Hexanus Banks, 1909
*ugandanus Neumann, 1906
ugandanus subsp. djaronensis
Neumann, 1907

Collections which include
the Types

Stages of eacli

species which

are known

Page

Toulouse, London, s 9

$ ? O L

266, 334 et seq.

Cambridge, 9

9

269

Leyden, 1 ?

?

226, 341

Washington (c), 9

$

173, 341

Toulouse, 1 i , 1 ?

t ?

164, 334 et seq.

Toulouse, 7 ? s, 12 os

? o

217, 341

Hamburg, 1 9

?

159

Cambridge, 2 ? s (o lost)

?

185, 341

Paris (a), 9 o ; (b) 9

? o

242, 341

London, 4 ? s

?

220, 341

Cambridge, 2 9 s, 3 o s

? 0

221, 341

Berlin, 1 $ ; Toulouse,

i 9 O L

221, 316, 334 et seq.

i 9 L

Toulouse, t ?

i 9

226, 334 et seq.

Washington (d), a 9

i 9

174

Paris (a), 1 9 ; Pickard-
Cambridge, o

i 9 o

256, 317, 334 et seq

Toulouse, i 9 ; Berlin

t 9

228, 334 et seq.

Lost1

i 9 O L

143, 296, 334 et seq

Washington (c)

<? 9

159, 334 et seq.

Hamburg, 1 9 ;
Toulouse, 1 9

9

158, 334 et seq.

Lost

<T 9

156, 315, 334 et seq.

Toulouse, 3 $ ;
Pretoria, <$ ?

i 9

204, 334 et seq.

Katanga, ¥

9

207, 334 et seq.

Toulouse, 1 9 , 3 o s
Paris (a), s ; Berlin, 9 s

o s ; Cambridge, 9 o ;

co-types
Washington (6), 1 9
St Petersburg, 9 ; Hamburg,

o ; Cambridge, o l
London, 9 o
London, 4 9 s
Paris (a), 14 9 s
Paris (a), 9 s ; Berlin, 1 9
London, £ ; Berlin, 9 o ;

Cambridge, 9 o l
Washington (rf), 9
London, 1 i , 1 9
Sjostedt, i 9

9 o
i 9 o

9

9 O h

9 o

9
9
9

no

175, 341

238, 334 et seq.

171, 341
261, 341

207, 341
171, 341
216, 341
244, 341
246, 334 et seq.

214, 341

230, 334 et seq.

233, 334 et seq.

1 Not in Upsala : only Hyalomma aegyptium still in the Linnean Collections, 1911.
* Species contained in the Cambridge collection ; types, and others which have been
compared with the types.

348 Genus Ixodes

Names of Species,
Authors and Date

Collections which include
the Types

Stages of each

species which

are known

Page

unicavatus Neumann, 1908

Edinburgh, ? o l;
Cambridge, h

ioi

264,

341

vespertilionis Koch, 1844

Berlin, 1 ?

i ? 0 L

271,

334 et seq

vestitus Neumann, 1908

London, 1 s ; Cam-
bridge, O L

? O L

252,

341

* 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.B.S., Quick Laboratory,

University of Cambridge.
Edinburgh ,, Collection of Mr W. Evans, F.B.S.E., Edinburgh.
Hamburg ,, Naturhistoriscb.es 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 Boad, London.
Paris (a) ,, Musec d'histoire naturelle, rue Buffon, Paris.

,, (b) ,, Laboratoire de Parasitologic, Ecole de Mfidecine, Paris.

Pickard-Cambridge signifies Collection of Eev. 0. Pickard-Cambridge, F.B.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 Sjdatedt Zoological Expedition.

Toulouse ,, Collection of Professor L. G. Neumann, Laboratoire d'Histoire

naturelle, Ecole Nationale Veterinaire, Toulouse, France.
Washington (a) signifies Bureau of Animal Industry, U. S. Department of Agriculture,
Washington, D.C.
,, (6) ,, 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.

,, ('■) ,, 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.)

< .

QL

A2T43
pt.1-2

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