SYlVIPOSIUi.

\1
ON
PHYSIOLOGY IN RELATION TO BEHAVIOUR
(Sat~wday, zznd. july, r967)
Chairman : T. E. HUGHES (U.K.)

THE BEHAVIOUR AND PHYSIOLOGY OF TICKS

BY

A. D. LEES

Agricultural Research Councü Unit of Insect Physiology,

Zoological Department, Cambridge.

An ideal method of approaching the study of behaviour has been outlined by

by MARKL and LINDAUER (rg65). Observation and description will come first.
This is followed by an experimental analysis which may, with good fortune, reveal
the conditions - both environmental and within the animal itself - which permit
the behaviour patterns to be expressed. Lastly, the behaviour is interpreted in
terms of the known properties of the nervous system.
It must be admitted that the subject of acarine behaviour has not yet advanced
far towards this goal. Experimental studies, which are not numerous, have
usually been concerned ·with the discovery of the sensory perceptions and with
the contribution of different environmental and other factors to the whole beha-
viour pattern as observed in the natural habitat. The neurophysiological basis
of behaviour is a virgin field, and even the gross morphology of the nervous system
is incompletely known in many groups.
The sensory physiology of free-living mites has recently received sorne atten-
tion. For example, the reactions of Bry obia praetiosa to gradients of humidity
have b een analysed in depth by Winston (rg63a, b). The response to light, humi-
dity and temperature have been described in severa! other Tetranychidae by
Susrn and NAEGELE (rg63) and l\-fom (rg6r, rg6za and b). The sensory percep-
tions of certain oribatids have also been examined by MADGE (rg64, rg66) and
WooDRING (rg66). Hovvever, it is the parasitic mites with their strongly differen-
tiated behaviour patterns that seem to offer particular advantages as experimental
material. The most striking behavioural adaptations occur in ticks in which the
A caro/ogia, t. XI, fasc. 3, rg6g.
 -- 398 -
clear separation of the free-living and the parasitic phases of the life cycle leads
to considerable elaboration in the behaviour mechanisms associated •vith host-
finding.
CAMIN (r963) distinguishes three main types of behaviour pattern in parasitic
mites. Species which live continuously on the host, such as Sarcoptes scabiei,
feed more or less continuously and their behaviour tends to " become so reduced
and sim pli fied as to be almost uninteresting ". CAMIN's second group includes
mites that are associated with nest or burrow-dwelling vertebrates. Many Der-
manyssid and Laelaptid mites take sizeable meals of blood or lymph. vVhen
feeding is completecl they leave the host but do not move far away, usually remain-
ing in the nest material. The location of the host does not therefore present great
difficulties. Nevertheless, these mites exhibit a much wiàer array of sensory per-
ceptions. Carnin notes from his own work on the snake mite Ophionyssus natn:cis
and the spiny rat mite Echinolaelaps echidnimts that these species react negatively
to light and gravity and positively to moist air up to 95 % R.H. Y et the response
patterns are highly inflexible, seeming to be little influenced by the nutritional
state or by other physiological modalities. The specificity of the host-fincling
reactions is conferred by the response to odours, temperature and tactile stimula-
tion. vVhen close to the host, the snake mite makes use of temperature and olfac-
tory elues and finally responds to the tactile sensations imparted by the over-
lapping scales. The olfactory sense is particularly acute in Echinolaelaps and the
temperature sense in the chicken mite, Dermanyss1,ts gallinae.
CAMIN's third group includes the ticks and chiggers which spend nearly all
their life cycle away from the host and which are therefore less protected from
environmental hazards. Their sensory physiology is relatively elaborate ; they
are sensitive to a wide variety of environmental signais; and the response often
depends on the physiological state of the individual.
The host-finding behaviour of Ixodes n:cinus will be taken as an example. As
is well known from the work of MILNE (1944, 1945, 1950) and others, the typical
ground habitat of this tlu·ee-host species is the moist vegetation layer that is typi-
cal of the rough moorland grazings in northern England and Scotland. The hun-
gry lm·vae, nymphs and adults of both sexes all climb the coarse grass and rush
stems and it is there that the host is usually encountered. The latter is most
likely to be a hill sheep but small mammals and birds of the wild fatma play a
minor role (MILNE, 1949a, b). After engorgement the replete tick drops to the
grouncl and regains the moist microclimate of the vegetation mat.
After the moult the tick is most difficult to arouse, and shows none of the beha-
viour characteristic of the hungry tick. No activity occurs before the digestion
of the previous meal has been completed - a process which may last for sorne
months. Activation may be assisted by the spring temperature rise, but is not
closely bound to this factor since populations of ticks which are placed in the natu-
ral habitat at different times of year also show activity peaks which are out of
phase with the local tick population. This means, of course, that seasonal activity
 - 399-
maxima (which vary with the locality) are determined by the availability of ticks
in the right physiological condition rather than by extrinsic ractors.
The phase of locomotor activity which succeeds the quiescent phase is meta-
bolically exhausting. Captive female ticks which are allowed to remain undisturb-
ed in humid air survive for nearly r8 months, even at high temperatures (25° C).
But the duration of survival is drastically reduced if they are provoked to prema-
ture activity by removing them periodically from their storage tubes (LEES, 1964).
In nature the active phase does not last for more than 30 days on average; and
the actual survival period is mainly determined by the length of the quiescent
p eriod prior to the onset of activity.
The 'appetetitive' behaviour displayed by the active ticks is characterised also
by a lowered threshold to sensory stimulation. The information received from
the environment is used by the tick in a sequence of manœuvres which culminate
in attachment to the host. The tick is first guided to a position near the vegeta-
tion tips 'vhere a host is most likely to be encountered . If a host approaches, the
tick is immediately alerted . Orientation to the host follows and the tick climbs
on. Finally, the host is either accepted or rejected. If the former is the case,
attachment ensues and with engorgement proceeding, the appetetitive behaviour
is inhibited.
Several kinds of orientation are involved during the first activity phase. A par-
ticularly important one is displayed vvhen the object climbed is a short vertical
plant stem (or in the laboratory a glass rod). The tick climbs steadily to the tip
but rarely cornes to rest immediately. Before this occurs the direction of move-
ment is usually reversed several times, although on each occasion the tick departs
only a short distance from the terminal point. This yo-yo-like movement appears
to be a response to the arrivai at a 'free end' of the substratum, since it is not seen
if the tick is confined in a glass tube. It may be partially tactile in nature since
the ticks can be seen to touch the top of the rod with the fore legs. But there
seems also to be an additional mechanism for limiting the height of climb . This
cornes into operation tick is made to climb an unusually high obj ect. In such
circumstances the tick often fails to reach the apex but turns round and retreats
to the base, without showing the usual repetetive movements. vVrLKINSON (1953)
has observed that when larvae of the cattle tick Boophihts micropl~ts were offered
a 6 ft. lath for climbing, larval groups tended to form no more than 2-4 ft. from
the ground (that is, within the range of a bovine host) ; although a few isolated
larvae climbed to the top none of them stayed there. In sorne Norwegian habitats
of 1. 1'icinus, where alders and other trees are quite common, there is evidence of
zonation within the vegetation cover : the adults are found at a height where they
could only encounter birds, cattle or horses; nymphs occur lower in the vegetation
and larvae lowest of all (TAMBS-LYCHE, 1943). It appears possible that the ticks
possess a mechanism which registers the distance climbed and which initiates a
turning movement if this has not already been induced by the tactile perception
of a 'tip'.
 -400-

The response to humidity is an important factor governing orientation within

the vegetation layer. Fully hydrated ticks tend to avoid high humidities and this
behaviour may serve to reinforce upward-turning if the tick happens to descend
into the lower parts of the vegetation cover where the humidity gradient becomes
steeper. But this taxis is very transient and disappears completely with the
slightest degree of desiccation. The strong hygropositive response which replaces
it causes the tick to come to rest on entering the zone of humid air. Since I. rici-
mts readily absorbs water from unsaturated air (dawn to 88% R.H. in the case
of the young tick) this function would be expected to have considerable adaptive
value. An examination of the activity records of individually marked ticks has
shown that the ticks do indeed disappear into the vegetation mat at irregular inter-
vals. And in these circumstances the replacement of water lost by evaportation
at the tips will undoubtedly follow. However, sorne active ticks remain conti-
nuously at the tips and are presumably able to recover sorne water at night when
the humidity usually rises to saturation point.
Ticks that are waiting at the vegetation tips either extend their forelegs stiffly
or wave them about in the manner of antennae. If no stimulus is forthcoming
the degree of excitation appears to subside and they assume an attitude of repose
with the legs folded. Nevertheless, the questing response is instantly elicited if
they are shaded or if the vegetation is disturbed in a way that might be associated
with the passage of a host - very rapid adaptation occurs in response to the nor-
mal swaying movements of the vegetation in the wind. It also seems likely that
co2 from the breath of the host may have an activating effect, as it does in other
ticks (NEVILL, rg64). 1 have noticed that most investigators seems to check the
vitality of their tick stocks by breathing on them !
Directional orientation is only seen when the host is close by. The gradients
of temperature and odour are important in these circumstances, particularly if
the host is upvvind. It is of interest that both stimuli must be presented together
in order to elicit a vigorous response. Sheep wool at room temperature is not
attractive to ticks; and a warm odourless object is actually repellent, except for
the less discriminating juvenile stages. When enumerating the many factors
involved in the attainment of a host one must not omit the remarkable facility
with which ticks cling to moving abjects. A blanket dragged over the vegetation
'sweeps np' ticks most effectively even though the temperature and odour compo-
nents are missing from the stimulus situation.
During the final phase of the food-locating process the tick continues to expe-
rience olfactory and temperature stimulation while moving about actively on the
host. In addition, contact chemoreceptors on the palps are brought into play.
Attachment then ensues if the host is acceptable. After engorgement is com-
plete, the tick drops to the ground and, actuated particularly by the strong pho-
tonegative behaviour, immediately burrows dawn into the deeper layers of the
vegetation mat.
Experimental studies have shown that most of the organs responsible for the
 -- 401 -

sensory perceptions are borne on the forelegs. Haller's organ, which is a duplex
structure, contains olfactory and probably humidity receptors. Two other types
of sensillum are also found, short thick-walled temperature receptor bristles on
the dorsal and lateral surfaces of the foreleg and tactile mechanoreceptors which
rest lightly against the substratum when the legs are folded and which are there-
fore ideally placecl for perceiving vibrations.
Since host-finding and host selection involves a relatively complex sequence
of behaviour patterns it is worth enquiring whether the degree of host specificity
observecl in ticks is associatecl with differences in the sensory physiology or whe-
ther other attributes are involvecl. An experimental approach has been macle to
this problem, using Ixodes ricin2ts and Ixodes canisuga Johnson as material. These
two species occur in the same localities in the north of England - indeed they
are often found on the same farm - yet they are ecologically isolated.
I. canis2tga has been recovered from foxes, but the sheep clog is an important
secondary host. On hill farms in northern Englancl the dogs are often kennelled
in disused stables or stone-walled onthouses where very heavy infestations are
sometimes built up. As a habitat the kennel serves as a excellent substitute for
the fox's earth. The dogs spencl only part of the day gathering sheep on the fells
but during this time they pick up sheep ticks from the vegetation. Ixodes ricimts
engorges normally on this host and is certainly droppecl, along with the engor-
ged Ixodes canisztga, on the floor of the kennels. If I. canis2tga shows a
drop-off rhythm (which is not known) one would expect that most ticks
would fall to the ground during the daylight hours, as the fox is mainly nocturnal.
There can be little doubt therefore that the dogs woulcl also drop engorged I. cani-
suga on the sheep grazings. Y et in spi te of this continuons invasion and conn ter-
invasion, I. ricin2ts cloes not become established in the kennel environment, nor
does I. canis2tga become established on sheep pastures. Incleed this species -vvas
never recorded from sheep by MILNE (1949a, b) clespite intensive sampling.
The reason why I. ricimts does not survive in the kennel is simply that the
environment is slightly too dry. Evaporation records kept in one particular
kennel in Northumberland, showecl that the average humidity was consi-
derably less than the near-saturated atmosphere of the vegetation mat on the sheep
grazings. Moreover, engorgecl females which were placecl in the kennel habitat
became clesiccated during the summer months before the eggs could be laid. Indeed,
the shrivelled bodies of ticks dropped by dogs were sometimes discovered on the
floor of the kennel. Of course, MILNE (1948) has already shown that the sheep
tick is confined to areas where the ground cover is capable of maintaining a humid
microclimate throughout the year. The kennel is merely a special case of this
relationship. The explanation is a physiological one. The effectiveness of the
epicuticular wax as a waterproofing agent varies widely in different species of
ticks (LEES, 1947). I. ricin2ts proved to have an exceptionally high transpiration
rate - approximately twice the rate characteristic of I. canisuga. Incidentally,
it would be interesting to know whether I. ricimts is confined to a humid environ-
Acarologia, t. XI, fasc. 3, I969. 26
 -402-

ment because it has a relatively permeable cuticular wax, or whether the high
permeability has resulted from the relaxation of selection pressure in a species
which, for other reasons, has come to inhabit a humid environment.
The reasons for the lack of success of 1. canisuga in the ricinus environment
are more subtle and probably involve the behaviour. Observations in the kennel
showed that engorged 1. canis~tga climb the walls and, since the bloated opistho-
soma hangs clown like a plumb-line, they proceed vertically upwards, finally disap-
pearing into a crevice where moulting or oviposition takes place. The unfed ticks
walk over the surface of the walls, questing freely. The largest aggregations
occur at 'dog height' where the ticks are attracted by the adour left by the dogs
as they rub against the wall. Although there is no apparent reason why engorged
canis1-tga ticks should not moult successfully in the moist environment of the vege-
tation layer, the behaviour of the unfed ticks is totally unsuited to host-finding
in this habitat. This can readily be appreciated if hungry females of bath species
are placed in a pot-grown clump of Molinia grass and junc~ts. 1. ricimts behaves
in the normal fashion, climbing the stems and finally coming to rest near the tips.
Since their movements are almost entirely vertical, few ever attempt to escape.
1. canisuga, on the other hand, usually fails to climb the tangle of stems and spends
much time walking on the sides of the pot. If an ascent is made, the tick imme-
diately returns to the base without exhibiting any upward-turning movements
when near the tip. The preference of 1. canisuga for plane surfaces rather than
the highly convex surfaces of plant stems, together with the absence of the turning
response, may well be sufficient to exclude the tick from that part of the environ-
ment where it could encounter a sheep.
There are, of course, other possibilities. The sheep might not be attractive
as a host for 1. canisuga. However, tests showecl that hungry ticks responcl stron-
gly both to clog haïr and sheep wool. The response occurs in the absence of a tem-
perature difference, although it is intensified if such a gradient is present. It
seems therefore that non-host specifie odours play a more important role in host-
finding thau in 1. ricimts. The sensory stimuli associated with the attachment
process also lack specificity. Hungry females of 1. canisuga attach themselves
to the shaven flank of a sheep as readily as do 1. ricimts and, after feecling to reple-
tion, lay viable egg batches of normal size.
The sex ratio and mating behaviour may also be significant factors in limiting
the distribution of 1. canisuga. Parthenogenetic egg development is sometimes
possible in 1. canis~tga, as it is in 1. ricimts, but the viability of the eggs is extre-
mely law. The sex ratio appears to be close to equality in 1. ricimts. On the
other hand, a collection of 596 nymphs of 1. canisuga, reared on hedgehogs, pro-
duced 400 female and 140 male ticks, a ratio of nearly 3 : I. The high proportion
of males in 1. ricimts is associated with the fact that mating takes place on the
host. Bath sexes therefore have to undergo the hazardous process of finding a
host. Not unnaturally, the male has the same sensory responses as the female
tick and even occasionally takes a 'vestigial' blood meal after climbing on to the
 -403-
sheep. Males of I. canisuga meet and copulate with engorged females in the more
confined environment of the burrow or kennel. The male is not required to find
a host and is in practice never found on kenneled sheep dogs. The reason for
this is not completely clear since the males seem to possess many of the reactions
necessary for host finding. Whatever the explanation, it is quite evident that
opportunities for mating would never ocur in the spacious ricinus environment.
It will be seen then that the two hasts, sheep and dog, are in many ways equally
suitable for bath species of tick. I. ricinus is excluded from the canisuga habitat
because of its greater cuticle permeability. Other characteristics, including malad-
justments of the host-finding behaviour, prevent I. canisuga from colonising the
ricinus habitat.
CAMIN (r963) and his associates, who have made a detailed study of the beha-
viour of Haemaphysalis leporispalustris, have found that whereas the immature
stages occur on a variety of birds and mammals, the adult ticks are virtually con-
fined to one host, the cottontail rab bit. The behaviour pattern is adapted accordin-
gly. The larvae and nymphs show much the same kind of orientations as I . rici-
mts in climbing to the top of low growing vegetation. Haemaphysalis executes
the same turning movements and shows an initial negative hygrctaxis. But all
the stimuli are not identical, since Haemophysalis is strongly photopositive whe-
reas I. ricimts is indifferent to light. Desiccation causes the light response to
disappear, with the result that the ticks leave their position of vantage and return
to the base of the vegetation layer where they absorb water. In contrast, the
adult ticks are uniformly photonegative and hygropositive, a response pattern
usually associated with the engorged stages. This behaviour virtually restricts
the adult to the temporary 'forms' constructed by the rabbit.
The sensory physiology of argasid ticks has received less attention. A feature
of their host-finding behaviour is clearly the token response to carbon dioxide.
The activating effect of co2 was reported in trombiculids by SASA, TANAKA, UENO
and MruRA (r957) and later by GARCIA (r962) in ticks. The mode of action of
co2 was described in greater detail by NEVILL (rg64) in the African tampan Orni-
thodoros savignyi. These ticks live in sand at a depth of several inches but readily
come to the surface if an artificial source of carbon dioxide is presented. The
ticks have considerable powers of locating the source of the gas. Moreover, C0 2
activation lowers the thresholds to other 'host stimuli'. The ticks are then most
strongly attracted to warm, moist objects. They are less attracted to warm, dry
abjects and are indifferent to cold, dry ones.
In a well lmown footnote NuTTALL (rgrr) reported an experience in which a
cow heavily infested with fully engorged females of Boophilus decoloratus was led
from a warm stail into the cooler air outdoors. The ticks immediately began
dropping off and " rattled like peas" on the ground. Nuttall believed that this
response to temperature would tend to confine the ticks to cattle paths which
would usually be traversed in the cool of the evening. This question of tick 'drop-
off' is of great ecological significance. Any mechanism which will prevent slow-
 -404-

feeding ticks from falling to the ground in areas which are infrequently visisted
by the host would clearly have great selective value. When the host is a wander-
ing animal such as a sheep the advantages may be less marked or even absent;
but one would expect such mechanisms to be fully developed if the host inhabits
a burrow or nest.
Working with Haemaphysalis leporispalustris, George (rg64) has shown that
drop-off from the host follovvs a definite circadian periodicity which is adapted
to the behaviour pattern of the host. The cottontail rabbit is nocturnal and
forages at random during the night. During the day the rabbit remains concealed
in grass forms or in brushwood thickets. It was shown experimentally that tick
drop-off is confined almost exclusively to the daylight hours, the peak occurring
4-6 hr. before the onset of darkness. This behaviour proved to be a true circadian
rhythm and not merely a direct response to light. Thus the entrained rhythm
persists in constant darkness, the phase being maintained for several days with
only slight drifting. In order to decide whether the drop-off response was con-
trolled by the tick itself or by sorne nutritional or sensory clue emanating from
the host, the rabbits were fed on a reversed schedule with the feeding time adjusted
so asto fall at the end of the dark period rather than at the end of the clay. In these
conditions drop-off remainecl synchronous with the light period. Nevertheless,
the phase of the rhythm coulcl be shifted by this procedure if the primary rhythm
was weakenecl by exposing both host and ticks to constant illumination. The
results of this experiment suggestecl that periodic stimuli from the host have a
definite significance but that these are not sufficiently decisive to overcome the
strong endogenous rhythm in the tick.
A similar adaptation has been found in the kangaroo tick Ornithodoros gurneyi
which lives in dusty wallows macle by the red kangaroo on the shacly side of bushes
in the ariel regions of Australia (BROWNING, rg6z). During the hot months the
kangaroo spends most of the clay resting in the wallows but leaves them at night
to forage on the open plains. BROWNING has shown that the frequency with which
the wallows are visitecl is relatecl to the density of suitable bushes. The nymphs
and adults of O. gt,trneyi are rapicl feeclers and are able to complete their engorge-
ment and drop from the kangaroo before it leaves the wallow. The larvae, on the
other hancl, require several clays to engorge. If dropped outside a wallow, their
chances of moulting and then encountering a host would be negligable.
Using rats as experimental hosts BROWNING (persona! communication) has
shown that in conditions of alternating light and clark, the drop-off pattern is far
from random and coïncides with the claylight hours. In this instance the rhythm
seems to be mainly, if not wholly, endogenous since there is, as yet, no evidence
for an indirect effect exerted through the host. Thus the ticks drop off at ranclom
when they are fed in clarkness on rat hosts with an establishecl and persistent
rhythm of activity (as displayecl in a running wheel). Clearly, the mechanism of
the drop-off rhythm, as well as many other facets of acarine behaviour, offers a
fruitful field for future research.
 -405-

REFERENCES

BROWNING (T. 0.), 1962. - Distribution of the Kangaroo-tick Ornithod01'os gurneyi in

arid South Australia. - Nature, Lond., 194 : 162-164.
CAJVIIN (]. H .), 1963. - Relations between host-fmding behaviour and life histories in
ectoparasitic acarina. - Advances in Acarology, 1 : 4rr-424.
GARCIA (R.), 1962. -Carbon dioxide as an attractant for certain ticks (Acarina : Arga-
sidae and Ixodidae). - Ann. ent. Soc. Am., 55 : 605-6.
GEORGE (J. E.), 1964. - The circadian rhythm of "drop-off" of engorged Haemaphy-
salis lep01·ispalttstris from rabbits. - Proc. lst int. Congr. Acarology. Acarologia,
6 (suppl. part) : 315-323.
LEES (A. D.), 1947. - Transpiration and the structure of the epicuticle in ticks. -
J. exp. Biol., 23 : 379-410.
LEES (A. D.), 1948. - The sensory physiology of the sheep tick (Ixodes ricinus L.).-
J. exp. Biol., 25 : 145-207.
LEES (A. D.), 1964. - The effect of ageing and locomotor activity on the water trans-
port mechanism of ticks. - Proc. lst int. Congr. Acarology. Acarologia, 6 (suppl.
part) : 315-323.
LEES (A. D.) & MILNE (A.), 1951. -The seasonal and diurnal activities of individual
sheep ticks (Ixodes ricin-us L.). - Parasitology, 41 : 189-208.
MADGE (D. S.), 1964. - The humidity reactions of oribatid mites. - Acarologia, 6 :
566-591.
lVIADGE (D. S.), 1966. - The significance of the sensory physiology of oribatid mites
in their natural environment. - Acarologia, 8 : 155-160.
lVIARKL (H.) & LINDAUER (lVI.), 1965. - The physiology of lnsecta, 2:3-122. Ed. lVI. Rock-
stein. New York & London.
MILNE (A.), 1944. - The ecology of the sheep tick, Ixodes ricimts L. Distribution of
the tick in relation to geology, soil and vegetation in northern England. - Para-
sitology, 35 : 186-196.
MILNE (A.), 1945. - The ecology of the sheep tick, Ixodes ricin-us L. The seasonal
activity in Britain with particular reference to northern England. - Parasito-
logy, 36 : 142-152.
MILNE (A.), 1949a. - The ecology of the sheep tick, Ixodes ricimts L. Host relation-
ships of the tick. Part r. Parasitology, 39 : 167-172.
lVIILNE (A.), 1949b. - The ecology of the sheep tick, Ixodes 1•icimts L. Host relation-
ships of the tick. Part 2. - Parasitology, 39 : 173-197.
lVIILNE (A.), 1950. - The ecology of the sheep tick, Ixodes ricimts L. lVIicrohabitat
economy of the adult tick. - Parasitology, 40 : 14-34.
MaRI (H.), 1961. -Comparative studies of the thermal reaction in four species of spider
mites (Acarina : Tetranychidae). - ]. Fac. Agric. Hokkaido Univ. Sapporo, 51 :
574-59!.
MaRI (H.), 1962a. - A seasonal difference of phototactic response in three species of
spider mites (Acarina: Tetranychidae).-]. Fac. Agric. Hokkaido Univ. Sapporo,
52 : 1-9.
 -406-
MoRI (H.), 1962b. - The effects of photo-stimulus on the thermal reaction in four species
of spider mites (Acarina : Tetranychidae). - J. Fac. Agric. Hokkaido Univ. Sap-
poro, 52 : 10-19.
NEVILL (E. M.), 1964. - The role of carbon dioxide as stimulant and attractant to the
sand tampan Ornithodoros savignyi (Audouin). - Onderstepoort J. Vet. Res.,
31 : 59-68.
NuTTALL (G. H.F.) & WARBURTON (C.), 1911.- Ticks, a monograph of the Ixodoidea:
Part. II. Cambridge Univ. Press, 348 pp.
SASA (M.), TANAKA (Y.), UENO (Y.) & MruRA (A.), 1957. -Notes on the bionomics of
Trombicula scutellaris and Trombicula akamushi, with special reference to the
mechanism of cluster formation and reaction to carbon dioxide expired by the
host. - Jap. J. exp. Med., 27 : 31-43.
SusKI (Z. W.) & NAEGELE (J. A.), 1963. - Advances in Acarology, 1 : 435-453. Ed.
J. A. Naegele, etc.
TAMBS-LYCHE (H.), 1943. - Ixodes riciwus og piroplasmosen i Norge. - Norsk Vet-
Tidsskr. nos. 9-12.
WILKINSON (P. R.), 1953. - Observations on the sensory physiology and behaviour of
larvae of the cattle tick, Boophil~ts microphts (Can.) (Ixodidae). - Aust. J. Zool.,
1 : 345-356.
WINSTON (P. W.), 1963a. - Humidity relations in the claver mite, Bryobia praetiosa
Koch. - Ecology, 44 : 669-678.
WINSTON (P. W.), 1963b. - Possible humidity receptor mechanisms in the claver mite,
Bryobia praetiosa Koch. - J. Insect Physiol., 9 : 89-103.
WooDRING (J. P.), 1966. - Color phototactic responses of an eyeless oribatid mite. -
Acarologia, 8 : 382-388.

DISCUSSION.

J. BRADY (U.K.).
Is there any evidence that I. ricimts is distasteful to birds ? Questing ticks
would seem to place themselves in an ideal position for predation.
LEES.
There is no reason to believe that ticks are unpalatable to birds. Perhaps
their natural agility saves them.
HOBART (U.K.).
Did Dr. Lees find any evidence that handling ticks affected their normal beha-
viour?
LEES.
Rough handling certainly causes the sensory responses to become attenuated
or even to disappear. The ticks fail to settle on the rods and lose their foothold
easily. They also adopt a distinct and characteristic gait while walking on a hori-
zontal surface.
 -407-
J. A. WALLWORK (U.K.).
There appears to be an appreciable amount of individual variation in the acti-
vity of the tick once it arrives at the tip of the glass rod, in the experiments des-
cribed. Can Dr. Lees offer an explanation for this variation ? Could this be a
response to variations in experimental conditions ?
LEES.
This is possible. My impression is that the number of turning movements
may be related to the general activity level, and perhaps to the age, of the tick.

SoNNENSHINE (U.S.).

Marked Dermacentor variabilis adults placed on a platform containing vertical

rods (aluminium) and surrounded by a vvater trough climbed the rods in much
the same manner as yon have described, and oriented at or near the tips. But
after several days, many declined the rods and crawled to the water trough, or
oriented at the lower parts of the rods. This process continued, and very few of
the ticks climbed again to the tips. After several weeks, all ticks were far from
the tips of the rods. It may be that several populations co-exist in nature and
that these differ physiologically. The population of highly active ticks near the
vegetation tips would readily be able to find hosts, while other ticks deeper in the
vegetation mat may be less readily attracted by the host.

KEMP (U.K.).
In mid-summer Ixodes ncmus appears to be inactive at high temperatures
(6o-7o°F) but they are not dead, only immobile at the base of the vegetation.
Under these conditions they will become active if mechanically stimulated or if
the observer blows gently on them. Could there be a blockage of the influence
of temperature on activity as this reaches sorne upper threshold ?

LEES.
This is possible, but I did not observe this summer torpor in the ticks I worked
with.
A. E. TREAT (U.S.A.).
Does the shape of the vertical support influence the type of questing beha-
viour ? Is this behaviour similar, for example, on thin, fiat supports and on cylin-
drical rads ? On sharply pointed as compared with bluntly rounded rods ?

LEES.

This possibility was not investigated experimentally, but I suspect that a

fairly normal questing response would be seen if 1. ricinus was caused to climb
to the top of a narrow, fiat support. On the other hand, a vertical road gradually
tapering to a sharp point might well prove unfavourable for settling and adopting
the questing attitude.
 -408-
TREAT.

Is there any selection by the tick of the type of vegetation on which the quest-
ing is clone?
LEES.

I believe so. Relatively stout stems, such as those of rushes seem to be pre-
ferred to blades of grass (which are to sorne extent flattened) or to the very flexible
grass stems of small diameter. It would be interesting to examine the behaviour
of ticks on tree boles in localities such as Norway or the English Breckland, where
a tree cover is present.
FELDMAN-MUHSAM (Israel).
Y our picture shows I. ricimts standing on the grass with the capitulum upwards.
I have noticed Rhipicephahts sangttine~ts standing on the grass with their capitu-
lum downwards. Can you explain the difference of this position in the two species ?

LEES.

We11 in I. ricintts I think the direction in which they finally come to rest is ran-
dom and it does not seem to matter in the least from the point of view of host
fin ding.
FELDMAN-MUHSAM.

Is it not always upwards ?

LEES.

No, by no means.

FELDMAN-MUHSAM.

It is always downwards in Rhipicephaltts.

LEES.

It certainly is not in I. ricintts. Have you any suggestions as to the possible

function in Rhipicephaltts ?
FELDMAN-MUI-ISAM.

In R. sangttinetts the ticks always stand on the grass with their capitulum
downwards, and it seems to me that this particular position, with the first pair
of legs extended in order to detect the host, facilitates the attachment of the tick
to its host.
R. J. TATCHELL (Australia).
What effect does the presence of an observer, who will provide C0 2 , tempera-
ture and odour gradients, have on the behaviour of ticks in experimental situa-
tions ?
LEES.
If C0 2 in I. 1'icimts serves only as an alerting stimulus, it may well be that the
proximity of a human observer would have little influence on orienting reactions
to temperature and smell. But this point needs re-investigation.

lVIACFADYEN (U.K.).
Since the hungry tick must respond quickly to a heat stimulus is there any
evidence that its sense organs are stimulated by radiant energy rather thau by
temperature change ?

LEES.
No. Under experimental conditions ticks seem to respond equally well to
radiating and relatively non-radiating bodies, provided the surface temperature
is the same. In the field, the distance at vvhich ticks respond to a host is much
greater if the latter is on the windward side. In the main, this is probably a res-
panse to air temperature.

HUGHES (U.K.).
The cuticle is waxed, yet there is high transpiration and active uptake of water.
This is responsible for the up and then down movement of I. ricin-us which is meta-
bolically very wasteful. I s there any reason known for this apparently wasteful
behaviour which reduces in the end the chance of host finding ?

LEES.
I do not know of any reason. It is difficult to understand why sorne species
of ticks - especially I . ricinus - have such a poorly waterproofed cuticle, parti-
cularly as penneability is not related in any way to the ability to absorb water
from air of high humidity.

CAMIN.
As a possible partial answer to lVIr. Hughes' question, we have sorne evidence
that the critical temperature of the cuticle is lower than the host surface tempera-
ture and loss of water may serve as a feeding stimulus in the rabbit tick. There-
fore, in adapting to this they may have sacrificed the ability to retain water under
other conditions.

FRIEDHOFF (Germany).
Is there an anaesthetic in the saliva of the tick ? Persans infested with I. rici-
mts nymphs do not seem to feel the ticks.

R. J. TATCHELL (Australia).
Cattle infested with Boophiltts microphts can show intense irritation which
perhaps indicates that in this species the saliva does not contain an anaesthetic.
 -410-

FELDMAN-MUHSAM (Israel).
In our laboratory several people had been bitten by Ornithodoros ticks. They
never noticed it during feeding, only later by the local reaction which was left
at the site, did they became aware that they had been bitten.
FRIEDHOFF.
I have been bitten lots of times but have felt no pain, therefore there must
be an anaesthetic.
GREGSON (Canada).
The question of an anaesthetic is very evident in the case of D . andersoni. In
British Colombia we have sorne 300 records of paralysis in humans and rarely is
the causative tick felt or noticed before its fifth day of feeding when paralysis
first occurs.
J.H. OLIVER (U.S.).
It has been observed in my lab that there are two periods when the host seems
to be especially irritated by Dermacentors feeding : when the ticks are placed on
the host and again upon the rapid phase of feeding found in females after mating.
WINSTON (U.S.).
Initial insertion and imbedding of mouthparts was not felt when ticks attached
to me for the first 3-4 days. There must be a good anaesthetic for that part of
the attachment period.