2009. 17 (1)



Belozerov V. N.

About author:

Belozerov V. N., Biological Research Institute, St. Petersburg State University, Stary Peterhof, St. Petersburg, 198504, Russia


Concerning the problem of life history and such an important its aspect as seasonality of life cycles and their control enabled by dormant stages, the parasitiform mites reveal the obvious similarity with the acariform mites. This concerns the pres- ence of both main kinds of dormancy (diapause and quiescence). The great importance in the seasonal control of life cycles in some parasitiform mites, like in acariform mites, belongs also for combinations of diapause with non-diapause arrests, particularly with the post-diapause quiescence (PDQ). This type of quiescence evoked after termination of diapause and enabling more accu- rate time-adjustment in recommencement of active development, is characteristic of both lineages of the Parasitiformes — Ixodida and Mesostigmata (particularly Gamasida). The available data show that in ixodid ticks the PDQ may be resulted similarly after developmental and behavioral diapause. Reproductive diapause combined with the PDQ is characteristic of some gamasid mites (particularly the family Phytoseiidae), while most gamasid and uropodid mites with phoretic dispersal reveal the dormant state (apparently of diapause nature) at the deutonymphal stage. The uncertainty between diapause and non-diapause dormancy is retained in some many cases (even in ixodid ticks and phytoseiid mites), and the necessity of further thorough study of different forms of diapause and non-diapause arrests in representatives of the Acari is noted therefore.


Akimov, I.A., Grobov, O.F., Pileckaya, I.V., Baraban-
ova, V.N., Yastrebtzov, A.V., Gorgol’, V.T., et al.
1993. Pcheliny kleshch Varroa jacobsoni [Bee
mite Varroa jacobsoni]. Kiev, 256 pp. [In Rus-

Alfeev, N.I. 1948. [Diapause in ixodid ticks]. Trudy
voenno-med. akademii, Leningrad, 44: 50–60. [In

Athias-Binche, F. 1991. Ecology and evolution of
phoresy in mites. In: F. Dusbabek and V. Bukva
(Eds.). Modern Acarology. Academia, Prague,
vol. 1, pp. 27–41.

Athias-Binche, F. and Habersaat, U. 1988. An ecologi-
cal study of Janetiella pyriformis (Berl. 1920), a
phoretic Uropodina from decomposing organic
matter. Mitt. Schweiz. Entomol. Ges., 6: 377–390.

Axtell, R.C. 1969. Macrochelidae (Acarina: Mesostig-
mata) as biological control agents for synanthropic
flies. In: Proc. 2nd Intern. Congr. Acarology (1967),
pp. 401–416.

Babenko, L.V. 1956. [About seasonal events in life of
ixodid ticks Ixodes ricinus and I. persulcatus].
Med. parasitologiya i parasit. bolezni, 25:
346–352. [In Russian]

Babenko, L.V. 1967. [Variability in types and duration
of development in ticks Ixodes ricinus in depen-
dence on the weather of vegetation seasons]. In:

Voprosy parasitologii. Kiev, pp. 337–338. [In

Babenko, L.V. 1970. [Some ecological features of
nymphs Ixodes ricinus]. In: Vtoroe akarologiches-
koe soveshchanie (Abstracts.), Kiev. 1970, vol. 1,
p. 43–46. [In Russian]

Babenko, L.V. and Gal’chenko, S.S. 1976. [Peculiari-
ties of feeding and development of larvae in Ix-
odes kazakstani]. Med. parasitologiya i parasit.
bolezni, 45: 331–337 [In Russian]

Babenko, L.V. and Platonova, V.F. 1965. [On larval
diapause of Ixodes ricinus and I. persulcatus. 1.
Experimental data on the impact of photoperiod
upon unfed and engorged larvae]. Med. parasi-
tologiya i parasit. bolezni, 34: 69–73. [In Rus-

Badh, M.H. and McMurtry, J.A. 1984. Life history and
life table parameters for Phytoseiulus longipes
with comparative studies on P. persimilis and
Typhlodromus occidentalis (Phytoseiidae). Ac-
arologia, 25: 111–123.

Balashov, Yu.S. 1959. [Periodization of cycles of de-
velopment in ixodid ticks]. Med. parasitologiya i
parasit. bolezni, 28: 469–476. [In Russian]

Balashov, Yu.S. 1962. [Determination of physiological
age and the age structure of Ixodes ricinus and Ix-
odes persulcatus populations in the Leningrad Re-
gion]. Med. parasitologiya i parasit. bolezni, 31:
47–55. [In Russian]

Balashov, Yu.S. 1967. Krovososushchiye kleshchi (Ix-
odoidea) — perenoschiki bolezney cheloveka i
zhivotnykh [Bloodsucking ixodoid ticks as vectors
of human and animal deseases]. Leningrad, 319
pp. [In Russian]

Balashov, Yu.S. 1993. [Significance of continental drift
for distribution and evolution of ixodid ticks (Ac-
arina, Ixodidae)]. Entomol. obozrenie, 72:
929–936. [In Russian]

Balashov, Yu.S. 1998. Iksodovye kleshchi — parazity i
perenoschiki infektsiy [Ixodid ticks as parasites
and vectors of infections]. St. Petersburg, 287 pp.
[In Russian]

Balashov, Yu.S. 2009. Parazitism kleschey i naseko-
mych na nazemnych pozvonochnych [Acari and
Insect parasitism on terrestrial vertebrates]. St. Pe-
tersburg, 357 pp. [In Russian]

Beklemishev, V.N. 1970. Biocenologicheskie principy
sravnitel’noy parazitologii [Biocoenologic prin-
ciples of comparative parasitology]. Moscow, 502
pp. [In Russian]

Belikova, N.P. 1969. Comparative data on feeding ac-
tivity and duration of morphogenesis in ticks of
the genus Haemaphysalis. Sbornik trudov Vladi-
vostokskogo issled. instituta epidemiologii i mi-
crobioogii, 4: 248–254.

Belozerov, V.N. 1957. [About biology and anatomy of
the mite Poecilochirus necrophori Vitzt. (Parasiti-
formes, Parasitidae)]. Zool. zhurnal, 36:
1802–1813. [In Russian]

Belozerov, V.N. 1958. [Influence of atmospheric hu-
midity on the mite Ornithonyssus bacoti Hirst
(Parasitiformes, Liponyssidae)]. Entomol. obozre-
nie, 37: 47–63. [In Russian]

Belozerov, V.N.1963. [Day length as a factor determin-
ing the oviposition delay in adult females of Der-
macentor marginatus Sulz]. Med. parasitologiya i
parasit. bolezni, 30: 308–313. [In Russian]

Belozerov, V.N. 1964a. [Larval diapause in the tick Ix-
odes ricinus L. and its dependence on exogenous
conditions]. Zool. zhurnal, 43: 1626–1637. [In

Belozerov, V.N. 1964b. [Diapause and reactivation in
adult females of Dermacentor marginatus]. Vest-
nik Leningradskogo universiteta (Ser. 3, biology),
4: 5–11. [In Russian]

Belozerov, V.N. 1966. [Nymphal diapause in the tick
Ixodes ricinus L. 1. Dependence of behavior of
unfed nymphs and of development of engorged
nymphs on photoperiodic conditions]. Med. para-
sitologiya i parasit. bolezni, 6: 723–729. [In Rus-

Belozerov, V.N. 1967. [Nymphal diapause in the tick
Ixodes ricinus L. 2. Different forms of diapause in
nymphs and peculiarities of their regulation in
ticks I. ricinus of Leningrad population]. Parazi-
tologiya, 1: 279–287. [In Russian]

Belozerov, V.N. 1968. [Photoperiodic control of sea-
sonal development in ixodid ticks]. In: A.S. Dani-
levsky (Ed.). Fotoperiodicheskie adaptatsii u
nasekomykh i kleshchey. Leningrad, p. 100–128.
[In Russian]

Belozerov, V.N. 1969. [Photoperiodic reaction of larvae
and nymphs of the tick Haemaphysalis concinna
Koch.]. Parazitologiya, 3: 17–21. [In Russian]

Belozerov, V.N. 1970. [Nymphal diapause in the tick
Ixodes ricinus L. (Acarina, Ixodidae). 3. Photope-
riodic reaction of unfed nymphs]. Parazitologiya,
4: 139–145. [In Russian]

Belozerov, V.N. 1971. [Nymphal diapause in the tick
Ixodes ricinus L. (Acarina, Ixodidae). 5. Impact of
changes in photoperiodic conditions of unfed
nymphs on their development after engorgement].
Parazitologiya, 5: 481–487. [In Russian]

Belozerov, V.N. 1972. [An inversion of photoperiodic
reaction in the control of development and dia-
pause of nymphs Ixodes ricinus L. (Acarina, Ixo-
didae) and mechanisms of this phenomenon]. In:
N.I. Goryshin (Ed.). Problemy fotoperiodisma i
diapausy u nasekomykh. Leningrad, p. 175–192.
[In Russian]

Belozerov, V.N. 1973a. [Egg diapause in Ixodes rici-
nus and its dependence on photoperiodic condi-
tions of unfed adult females]. Vestnik Leningrad-
skogo universiteta, 9 (Biol. 2): 33–37. [In

Belozerov, V.N. 1973b. [Experimental analysis of sea-
sonal adaptations of Deracentor silvarum Ol.].
Parazitologiya, 7: 14–18. [In Russian]

Belozerov, V.N. 1974c. [Experimental study of sea-
sonal adaptations of Haemaphysalis concinna
ticks from different sites of the Far East]. Med.
parasitologiya i parasit. bolezni. 43: 31–38. [In

Belozerov, V.N. 1976a. [Life cycles and seasonal adap-
tations in ixodid ticks]. In: Doklady 28 yezhegod-
nykh chteniy pamyati N.A. Kholodkovskogo.
Leningrad, pp. 53–101. [In Russian] [In English:
NAMRU-3, T1124]

Belozerov, V.N. 1976b. [Seasonal cycle’s adaptations
in the tick Rhipicephalus turanicus B. Pom.]. In:
Tret’ya vsesoyuznaya konferentsiya po teoret-
icheskoy i prikladnoy akarologii (Abstracts).
Tashkent. pp. 41–42. [In Russian]

Belozerov, V.N. 1977. [Events of oligomerization in
transformations of life cycles of ixodid ticks]. In:
Znachenie polimerizatsii i oligomerizatsii v evo-
lutsii. Leningrad, pp. 25–28. [In Russian]
Belozerov, V.N. 1981. [Ecological rhythms in ixodid
ticks and their regulation]. Parasitol. sbornik, 30:
22–46. [In Russian]

Belozerov, V.N. 1982. Diapause and biological rhythms
in ticks. In: F.D. Obenchain and R. Galun (Eds.).
Physiology of Ticks. Pergamon Press, Oxford, pp.

Belozerov, V.N. 1985a. [Diapause, its place and role in
life cycles, mechanisms]. In: N.A. Filippova (Ed.).
Tayezhny kleshch Ixodes persulcatus. Leningrad,
pp. 214–219. [In Russian]

Belozerov, V.N. 1985b. [Data on biology of the tick
Rhipicephalus pumilio]. In: Pyataya vsesoyuznaya
akarol. konferentsiya (Abstracts). Frunze, p.
30–31. [In Russian]

Belozerov, V.N. 1988. [Photoperiodism and seasonal
development of ixodid ticks]. Abstract of the Doc-
tor of Biological Sciences thesis. St. Petersburg
State University, 40 pp. [In Russian]

Belozerov, V.N. 1990. [Peculiarities of induction and
termination of egg diapause in Ixodes ricinus]. In:
Dostizheniya meditsinskoy entomologii i
akarologii (Materialy X kongressa vsesoyuznogo
entomol. obshchestva). Leningrad, pp. 72–74. [In

Belozerov V.N. 1991. Evolution of life cycles in ticks
(Ixodidae) due to climate seasonality. In: F.
Dusbabek and V. Bukva (Eds). Modern Acarolo-
gy. Prague. Vol. 2, p. 135–139.

Belozerov, V.N. 1995a. [Characteristics of parameters
in two-step photoperiodic reaction controlling
seasonal development of Ixodes ricinus nymphs
(Acarina: Ixodidae)]. Parazitologiya, 29: 240–249.
[In Russian]

Belozerov, V.N. 1995b. [Photoperiodic control of de-
velopment and diapause in nymphs of the Taiga
tick Ixodes persulcatus (Ixodidae)]. Parazitologi-
ya, 29: 101–104. [In Russian]

Belozerov, V.N. 1995c. Preliminary data on the influ-
ence of photoperiod on development of Haemaph-
ysalis parva (Acarina, Ixodidae). Entomol. oboz-
renie, 74: 152–154. [In Russian]

Belozerov, V.N. 1998. Role of two-step photoperiodic
reaction in the control of development and dia-
pause in the nymphs of Ixodes persulcatus. Rus-
sian J. Zool., 2: 414–418.

Belozerov, V.N. 1999. Dormancy in the life cycles of
ixodid ticks and their adaptations to predictable
and unpredictable environmental changes. In: Ac-
arology IX. Vol. 2 (Symposia), USA, Columbus,
Ohio, p. 53–56.

Belozerov, V.N. 2006. Seasonal adaptations in the life
cycles of the Acari (mites and ticks): comparative
and evolutionary aspects. In: XII International
congress of Acarology (Book of Abstracts). Am-
sterdam, p. 29.

Belozerov, V.N. 2007. [Distribution of dormant stages
in the life cycles of the Acari (Chelicerata: Arach-
nida) in comparison with mandibulate arthropods
(Insects and Crustaceans)]. Trudy biol. nauchno-
issled. instituta St. Peterburgskogo gos. univer-
siteta, 53: 193–233. [In Russian, Engl. Summary]

Belozerov, V.N. 2008. Diapause and quiescence as two
main kinds of dormancy and their significance in
life cycles of mites and ticks (Chelicerata: Arach-
nida: Acari). Part 1. Acariformes. Acarina, 16:

Belozerov, V.N. 2009. New aspects in investigations of
diapause and non-diapausing types of dormancy
in insects and other arthropods. Entomol. Review,
89: 127–136.

Belozerov, V.N., Bogdanov, V.E. and Kvitko, N.V.
1966. [Seasonal changes in temperature reactions
of Ixodes ricinus engorged females]. Vestnik Len-
ingrad. universiteta (Ser. biology), 3: 37–44. [In

Belozerov, V.N., Fourie, L.J., Van der Lingen, F.J., and
Kok, D.J. 1996. Photoperiodic control of develop-
mental diapause in nymphs of the Karoo paralysis
tick Ixodes (Afrixodes) rubicundus Neum. (Ixodi-
dae). In: Acarology IX (Proc. IX Intern. Congr.
Acarol.), USA, 1996, vol. 1, p. 677–680.

Belozerov, V.N., Fourie, L.J., and Kok, D.J. 2003. Pho-
toperiodic control of developmental diapause in
nymphs of prostriate ixodid ticks (Acari: Ixodi-
dae). Exp. Appl. Acarol., 28: 163–168.

Belozerov, V.N. and Il’in, Yu.A. 1974. [Combination
of long-day and short-day processes in photoperi-
odic control of development of larvae and nymphs
in the tick Ixodes ricinus]. In: Materialy VII kon-
ferentsii vsesoyuznogo entomol. obshchestva.
Leningrad, pt. 1: 204–205. [In Russian]

Belozerov, V.N. and Il’in, Yu.A. 1981. [Pecualirities of
regulation of larval-nymphal metamorphosis in
the tick Ixodes ricinus L. (Acarina, Ixodidae) un-
der replacement of photoperiodic regimens]. Tru-
dy biol. nauchno-issled. instituta Leningradskogo
universiteta, 31: 143–160. [In Russian]

Belozerov, V.N. and Kvitko, N.V. 1965. [Main features
of photoperiodic reaction in ticks Dermacentor
marginatus Sulz. (Ixodidae)]. Zool. zhurnal, 44:
363–372. [In Russian]

Belozerov, V.N. and Lamanova, A.I. 1967. [Initiation
of winter diapause and the reaction to day length
in Dermacentor marginatus in Central Kazakh-
stan]. Med. parasitologiya i parazit. bolezni, 36:
464–469. [In Russian]

Belozerov, V.N. and Luzev, V.V. 1974. [Relationship
between the behaviour and development of larvae
and nymphs of Haemaphysalis longicornis Neum.
and photoperiodic conditions]. Parazitologiya, 8:
515–523. [In Russian]

Belozerov, V.N. and Mikhailova, Z.A. 1966. [Diapause
of larvae in the tick Ixodes ricinus L. and its de-
pendence on environmental conditions. 2. Interac-
tion of tick maintenance before and after their
feeding in processes controlling the diapause of
engorged larvae]. Zool. zhurnal, 45: 543–550. [In

Belozerov, V.N. and Mourad, M.G. 1977. [Photoperi-
odic regulation of nymphal diapause in the tick
Hyalomma anatolicum Koch]. Entomol. obozre-
nie, 56: 495–504. [In Russian]

Belozerov, V.N. and Naumov, R.L. 2002b. Nymphal
diapause and its photoperiodic control in the tick
Ixodes scapularis. Folia parasitol. (Prague). 49:

Belyaeva, N.S. and Ryabova, I.N. 1971. [Life cycles of
ixodid ticks in the southern part of Khabarovsk
territory]. In: Voprosy geografii Dalnego Vostoka.
Khabarovsk, pp. 302–326. [In Russian]

Berdyev, A.B. 1973. [Materials on the developmental
cycle of Rhipicephalus turanicus B. Pom. in natu-
ral biotopes of Turkmenia]. In: Parasity zhivot-
nykh v Turkmenii. Ashkhabad, pp. 5–15. [In Rus-

Berdyev, A.B. 1974. [About developmental cycle of
Hyalomma anatolicum excavatum in Turkmenia].
Med. parasitologiya i parasit. bolezni, 43: 38–43.
[In Russian]

Berkvens, D.L., Pegram, R.G. and Brandt, J.R.A. 1995.
A study of the diapausing behaviour of Rhipiceph-
alus appendiculatus and Rhipicephalus zam-
beziensis under quasi-natural conditions in Zam-
bia. Medical and Veterinary Entomology, 9:

Bregetova, N.G. 1956. Gamazovye kleshchi (Gama-
zoidea). [Gamasid mites]. Moscow-Leningrad,
247 pp. [In Russian]

Brendonck, L. 1996. Diapause, quiescence, hatching
requirements: what we can learn from large fresh-
water branchiopods (Crustacea: Branchiopoda:
Anostraca, Notostraca, Conchostraca). Hydrobio-
logia, 320: 85–97.

Broufas, G. 2002. Diapause induction and termination
in the predatory mite Euseius finlandicus in peach
orchards in northern Greece. Exper. Appl. Acarol-
ogy, 25: 921–932.

Burg, J.G. 2001. Seasonal activity and spatial distribu-
tion of hostseeking adults of the tick Dermacentor
variabilis. Medical and Veterinary Entomology,
15: 413–421.

Danks, H.V. 1987. Insect dormancy: an Ecological
Perspective. Biol. Survey of Canada, Monograph
Series, No. 1, 439 pp.

Danks, H.V. 1991. Life cycle pathways and the analy-
sis of complex life cycles in insects. Canad. Ento-
mologist, 123: 23–40.

Danks, H.V. 1999. The diversity and evolution of in-
sect life cycles. Entom. Science, 2: 651–660.
Dautel, H. and Knülle, W. 1998. The influence of phys-
iological age of Argas reflexus larvae (Acari: Ar-
gasidae) and of temperature and photoperiod on
induction and duration of diapause. Oecologia,
113: 46–52.

Dautel, H. and Knülle, W. 2006. Investigation into the
diapause and cold hardiness in Ixodes ricinus eggs.
In: XII International congress of Acarology (Book
of Abstracts). Amsterdam, p. 46–47.

Dautel, H., Dippel, C., Kämmer, D., Werkhausen, A.,
and Kahl, O. 2008. Winter activity of Ixodes rici-
nus in a Berlin forest. International Journal of
Medical Microbiology. 298, Suppl. 1: 50–54.

Dogiel, V.A. 1954. Oligomerizatsiya gomologichnykh
organov kak odno iz glavnykh napravleniy evo-
lutsii zhyvotnikh [Oligomerization of homologous
organs as one of the main pathways of animal evo-
lution]. Leningrad, 368 pp. [In Russian]

Doube, B.M. 1975. The biology of the Kangaroo tick,
Ornithodoros (Pavlovskyella) guerneyi Warbur-
ton (Acarina: Argasidae), in the laboratory. J.
Med. Entomol. 12: 240–243.

Drew, M.L. and Samuel, W.M. 1985. Factors affecting
transmission of larval winter ticks, Dermacentor
albipictus (Paccard), to moose, Alces alces L., in
Alberta, Canada. Journal of Wildlife Diseases, 21:

Drew, M.L. and Samuel, W.M. 1989. Instar develop-
ment and disengagement rate of engorged female
winter ticks, Dermacentor albipictus (Acari: Ixo-
didae), following single- and trickle-exposure of
moose (Alces alces). Exper. Appl. Acarology, 6:

Drummond, R.O. 1967. Seasonal activity of ticks (Ac-
arina: Metastigmata) on cattle in Texas. Ann. Ent.
Soc. Amer., 60: 439–447.

Emme, A.M. 1953. [Some problems in the theory of
insect diapause]. Uspekhy sovremennoy biologii,
35: 395–421. [In Russian]

Estrada-Peňa, A., Pegram, R.G., Barré, N., and Venzal
J.M. 2007. Using invaded range data to model the
climate suitability for Amblyomma variegatum
(Acari: Ixodidae) in the New World. Exp. Appl.
Acarol., 41: 203–214.

Evans, G.O. 1992. Principles of Acarology. CAB Inter-
national, UK, 563 pp.

Faasch, H. 1967. Beitrage zur Biologie der einheimis-
chen Uropodiden Uroobovella marginatra Koch
und Uropoda orbicularis Müller, und experiment-
elle Analyse ihres Phoresieverhalten. Zool. Jahrb.,
Syst., 94: 521–608.

Filippova, N.A. (Ed.). 1985. Tayezhny kleshch Ixodes
persulcatus Schulze. Morphologiya, sistematika,
ekologiya, meditsinskoe znachenie. Leningrad,
416 p. [In Russian]

Fourie, L.J., Belozerov, V.N. and Needham, G.R. 2001.
Ixodes rubicundus nymphs are short-day diapause-
induced ticks with termolabile sensitivity and des-
iccation resistance. Medical and Veterinary Ento-
mology, 15: 335–341.

Fujimoto, K. 1993. Effect of photoperiod on the attach-
ment and development of immature Ixodes persul-
catus. Japan J. Sanit. Zool., 44: 271–277.

Fujimoto, K. 2003. The host-seeking and feeding ac-
tivities of a new generation of Amblyomma testu-
dinarium nymphs (Acari : Ixodidae) in summer
observed under experimental conditions. Med.
Entomol. Zoology, 54: 395–397.

Goddard, J. 1992. Ecological studies of adult Ixodes
scapularis in central Missisipi: questing activity in
relation to time of year, vegetation type and me-
teorological conditions. J. Med. Entom., 29 (3):

Gotoh, T., Yamaguchi, K. and Mori, K. 2004. Effect of
temperature on life history of the predatory mite
Amblyseius (Neoseiulus) californicus (Acari: Phy-
toseiidae). Exper. Appl. Acarology, 32: 15–30.
Guglielmone, A.A. 1994. The seasonal occurrence of
Amblyomma triguttatum triguttatum. Acarologia,
35: 107–113.

Guglielmone, A.A. and Moorhouse, D.E. 1986. The ef-
fect of photoperiod on the development of Ambly-
omma triguttatum. J. Med. Entom., 23: 274–278.

Gurney, W.S.C., Crowley, P.H. and Nisbet, R.M. 1991.
Locking life cycles onto seasons: Circle-map
models of population dynamics and local adapta-
tion. Journ. Mathemat. Biology, 30: 251–279.

Gurney, W.S.C., Crowley, P.H. and Nisbet, R.M. 1994.
Stage-specific quiescence as a mechanism for syn-
chronizing life cycles to seasons. Theoretical pop-
ulation biology, 46: 319–343.

Gilyarov, M.S. and Bregetova, N.G. (Eds.). 1977.
Opredelitel’ kleshchey, obitayushchikh v potchve.
Mesostigmata [Identification Keys of soil-dwell-
ing mites Mesostigmata]. Leningrad, 718 pp. [In

Harris, J.A. 1973. Structure and dynamics of a cave
population of the Guano mite, Uroobovella copro-
phila (Womersley). Australian Journal of Zoolo-
gy, 21: 239–275.

Hartenstein, R. 1962. Life histories of Pergamasus
crassipes and Amblygamasus septentrionalis (Par-
asitidae). Ann. Ent. Soc. Amer., 55: 196–202.

Helle, W. and Sabelis, M.W. (Eds.). 1985. Spider mites.
Their biology, Natural Enemies and Control. Am-
sterdam. Elsevier. Vol. 1B, 458 p.

Hodek, I. 1996. Diapause development, diapause ter-
mination and the end of diapause. Europ. Journal
of Entomology, 93: 475–487.

Hodek, I. 2002. Controversial aspects of diapause de-
velopment. Europ. Journ. of Entomology, 99:

Hoogstraal, H. 1956. African Ixodoidea. I. Ticks of Su-
dan. Wash., 1101 pp.

Houck, M.A. and OConnor, B.M. 1991. Ecological and
evolutionary significance of phoresy in the Astig-
mata. Annual Review of Entomology, 36:

Karg, W. 1971. Acari (Acarina), Milben. Unterordung
Parasitiformes (Anactinochaeta). Die freilebenden
Gamasina (Gamasides), Raubmilben. In: Die
Tierwelt Deutschlands. Teil 59. VEB G. Fischer
Verlag, Jena. 475 s.

Karg, W. 1989. Acari (Acarina), Milben. Unterordung
Parasitiformes (Anactinochaeta). Uropodina
Kramer, Schildkrötenmilben. In: Die Tierwelt
Deutschlands. Teil 67. VEB G. Fischer Verlag,
Jena. 203 s.

Klompen, H., Lekveishvili, M. and Black, W.C. 2007.
Phylogeny of parasitiform mites (Acari) based on
rRNA. Molecular Phylogenetics and Evolution,
43: 936–951.

Kolonin, G.V., Pikunov, D.G., Basylnikov, V.I. et al.
1976. [Hibernation of ixodid ticks on wild ungu-
late animals of the Maritime territory]. Zool. zhur-
nal, 55: 1253–1255. [In Russian]

Koštal, V. 2006. Eco-physiological phases of insect
diapause. Journal of Insect Physiology, 52:

Koulianos, S. and Schwarz, H.H. 1999. Reproduction,
development and diet of Parasitellus fucorum
(Mesostigmata, Parasitidae), a mite associated
with bumblebees (Hymenoptera, Apidae). Journ.
Zool., 248: 267–269.

Krasinskaya, A.L. 1961. [Morpho-biological pecual-
iarities of the post-embbryonic development of
uropodid mites in the Leningrad Region]. Parazi-
tol. sbornik, 20: 108–147. [In Russian]

Labruna, M.B., Amaku, A., Metzner, J.A., Pinter, A.,
and Ferreira, F. 2003. Larval Behavioral Diapause
Regulates Life Cycle of Amblyomma cajennense
(Acari: Ixodidae) in Southeast Brazil. J. Med. En-
tomol. 40: 170–178.

Lange, A.B. 1984. [Subtype Chelicerata]. In: M.S. Gil-
yarov (Ed.). Zhizn’ zhivotnykh. Moscow. Vol. 3:
8–108. [In Russian]

Lehtinen, P.T. 1991. Phylogeny and zoogeography of
the Holothyrida. In: F. Dusbabek and V. Bukva
(Eds.). Modern Acarology, vol. 2. SPB Academic,
Prague, pp. 101–113.

Lindquist, E.E. 1984. Current theories on the evolution
of major groups of Acari and on their relationship
with other groups of Arachnida, with consequent
implications for their classification. In: D.A. Grif-
fiths and C.E. Bowman (Eds.). Acarology VI, vol.
1. Ellis Horwood, pp. 28–36.

Lindsay, L.R., Barker, I.K., Surgeoner, G.A. et al.
1995. Survival and development of Ixodes scapu-
laris under various climate conditions in Ontario.
Journ.Med. Entomol., 32: 143–152.

Liu, J.Z., Liu, Z.N., Zhang, Y. et al. 2005. Biology of
Dermacentor silvarum (Acari: Ixodidae) under
laboratory conditions. Exper. Appl. Acarology,
36: 131–138.

Loew, J. 1964. Über den Einfluss der Photoperiode auf
die Metamorphose von Ixodes ricinus. Angew.
Parasitol., 5: 3–13.

Lord, C.C. 1995. Seasonal activity of nymphal Ixodes
scapularis in different habitats of New Jersey.
Journ. Med. Entomol., 32: 66–70.

MacDonald, I.L. and Zuchini, W. 1997. Hidden Mark-
ov and other models for discrete-valued time se-
ries. Chapman and Hall, London.

Madder, M., Speybroeck, N., Brandt, J., Tirrry, L.,
Hodek, I., and Berkvens, D. 2002. Geographic
variation in diapause response of adult Rhipiceph-
alus appendiculatus ticks. Exper. Appl. Acarol.,
27: 209–221.

Madder, M., Speybroeck, N., Brandt, J., and Berkvens,
D. 1999. Diapause induction in adults of three
Rhipicephalus appendiculatus stocks. Exp. Appl.
Acarogy, 23: 961–968.

Mihm, J.A. and Chiang, H.C. 1976. Laboratory studies
of the life cycle and reproduction of some soil-
and manure-inhabiting predatory mites (Laelapti-
dae). Pedobiologia, 16: 353–363.

Mourad, M.G. and Belozerov, V.N. 1976. [Photoperi-
odical control of adult diapause in ticks Hyalomma
anatolicum]. In: Tretiya vsesoyuznaya konferen-
tsiya po teoreticheskoy i prikladnoy akarologii
(Abstracts). Tashkent, p. 73–74. [In Russian]

Müller, H.J. 1970. Formen der Dormanz bei Insekten.
Nova Acta Leopoldina, N.F., 35: 7–27.
Müller, H.J. 1992. Dormanz bei Arthropoden. G. Fis-
cher Verlag, Jena. 289 s.

Murrell, A., Dobson, S.J., Walter, D.E., Campbell,
N.J.H., Shao, R., and Barker, S.C. 2005. Relation-
ships among the three major lineages of the Acari
(Arthropoda : Arachnida) inferred from small sub-
unit rRNA: paraphyly of the parasitiformes with
respect to the opilioacariformes and relative rates
of nucleotide substitution. Invertebrate. Syst., 19,

Nawar, M.S., Shereef, G.M. and Ahmed, M.A. 1993.
Effect of food on development, reproduction and
survival of Chiropturopoda bakeri (Acarina:
Uropodidae). Exp. Appl. Acarol., 17: 277–281.

Nel’zina, E.N. 1951. Krysiny kleshch [The rat mite].
Moscow, 100 pp. [In Russian]
Nel’zina, E.N.and Danilova, G.M. 1960. [Rhipicepha-
lus schulzei as inhabitant of the little suslik bor-
rows (Citellus pygmaeus Pall.)]. Med. parasi-
tologiya i parasit. bolezni, 29: 291–300. [In

Norton, R.A., Kethley, J.B., Johnston, D.E., and
OConnor, B.M. 1993. Phylogenetic perspectives
on genetic systems and reproductive modes of
mites. In: D.L. Wrensch and M.A. Ebbert (Eds.).
Evolution and Diversity of Sex Ratio in Insects
and Mites. Chapman and Hall, New York, pp.

Norval, R.A.I. 1977a. Ecology of the tick Amblyomma
hebraeum in the eastern province of South Africa.
1. Distribution and seasonal activity. J. Parasi-
tol., 63: 734–739.

Norval, R.A.I. 1977b. Ecology of the tick Amblyomma
hebraeum in the eastern province of South Africa.
2. Survival and development. J. Parasitol., 63:

Nosek, J., Lichard, M., Sztankay, M. 1967. The ecol-
ogy of ticks in the Tribec and Hronsky Inovec
mountains. Bull. Org. Mond. Sante, 36 (Suppl. 1):

Ogden, N.H., Lindsay, L.R., Beauchamp, G, Charron,
D., Maarouf, A., O’Callaghan, C.J., Waltner-
Toews, D., and Barker, I.K. 2004. Investigation
of relationships between temperature and devel-
opmental rates of tick Ixodes scapularis (Acari:
Ixodidae) in the laboratory and field. J. Med. En-
tomol., 41: 622–633.

Oliveira, P.R., Borges, L.M.F., Leite, R.C., and Freit-
as, C.M.V. 2003. Seasonal dynamics of the Cay-
enne tick, Amblyomma cajennense, on horses in
Brazil. Medical and Veterinary Entomology, 17:

Overmeer, W.P.J. 1985. Diapause. In: W. Helle and
M.W. Sabelis (Eds.). Spider Mites. Their Biolo-
gy, Natural Enemies and Control. Vol. 1B, El-
sevier, Amsterdam, pp.95–102.

Ozman-Sullivan, S.K. 2006. Life history of Kampimo-
dromus aberrans as predator of Phytoptus avel-
lanae (Acari: Phytoseiidae, Phytoptidae). Exper.
Appl. Acarology, 38:15–23.

Patrick, C.D. and Hair, J.A. 1975. Ecological observa-
tions on Dermacentor albipictus in eastern Okla-
homa. J. Med. Entomol., 12: 393–394.

Pegram, R.G., Mwaze, E.T., Zivkovic, D. and Jonge-
jan, F. 1988. Morphognetic diapause in Ambly-
omma variegatum (Acari: Ixodidae). Medical and
Veterinary Parasitology, 2: 301–307.

Pomerantzev, B.I. 1948. [Main directions of the evolu-
tion in Ixodoidea]. Parasitol. sbornik, 10: 5–19.
[In Russian]

Pomerantzev, B.I. 1950. Iksodovye kleshchi [Ixodid
ticks (Ixodidae)]. Fauna of the USSR, Paukoo-
braznyye 4 (2). Leningrad, 224 pp. [In Russian]

Putman, W.L. 1962. Life-history and behaviour of the
predacious mite Typhlodromus caudiglans (Phy-
toseiidae) in Ontario, with notes on the prey of
related species. Can. Entomol., 94: 163–177.

Radinovsky, S., 1965. The biology and ecology of
granary mites of the Pacific Northwest. Life his-
tory and development of Leiodinychus krameri
(Uropodidae). Ann. Ent. Soc. Amer., 58: 259–267.

Radovsky, F.J. 1994. The evolution of parasitism and
the distribution of some Dermanyssoid mites (Me-
sostigmata) on vertebrate hosts. In: M.A. Houck
(Ed.). Mites: Ecological and evolutionary analy-
ses of life-history patterns. Chapman & Hall, p.

Randolph, S E. 1993. Climate, satellite imagery and the
seasonal abundance of the tick Rhipicephalus ap-
pendiculatus in southern Africa: a new perspec-
tive. Medical and Veterinary Entomology, 7:

Randolph, S.E. 1997. Abiotic and biotic determinants
of the seasonal dynamics of the tick Rhipicephalus
appendiculatus in South Africa. Medical and Vet-
erinary Entomology, 11: 25–37.

Randolph, S.E. 2004. Tick ecology: processes and pat-
terns behind the epidemiological risk posed by
ixodid ticks as vectors. Parasitology, 129: 37–65.

Randolph, S.E. and Rodgers, D.J. 1997. A generic pop-
ulation model for the African tick Rhipicephalus
appendiculatus. Parasitology, 115: 265–279.

Randolph, S.E., Green, R.M., Hoodless, A.N., and
Peace, M.F. 2002. An empirical quantitative
framework for the seasonal population dynamics
of the tick Ixodes ricinus. Intern. Journal for Par-
asitology, 32: 979–989.

Razumova, I.V. 1965. [Seasonal course of diapause ap-
pearance in Dermacentor pictus and factors of its
stimulation]. Med. parasitologiya i parasit. bolez-
ni, 1965, 34: 46–52. [In Russian]

Rechav, Y. 1981. Ecological factors affecting the sea-
sonal activity of the brown ear tick Rhipicephalus
appendiculatus. In: G.B. Whitehead and J.D. Gib-
son (Eds). Tick Biology and Control, South Afri-
ca, Grahamstown University, pp. 187–191.

Rubina, M., Hadani, A. and Ziv, M. 1982. The life cy-
cle of the tick Hyalomma anatolicum excavatum
maintained under field conditions in Israel. Rev.
Elev. Med. Vet. Pays Trop., 35: 255–264.

Rudzinska, M. 1998. Life history of the phoretic preda-
tory mite Arctoseius semiscissus (Acari: Ascidae)
on a diet of sciarid fly eggs. Exper. Appl. Acarol-
ogy, 22: 643–648.

Saito Y, 1990. Life-history and feeding habit of Typhlo-
dromus bambusae, a specific predator of Schizo-
tetranychus celarius. Exp. Appl. Acarology, 10:

Sapozhnikova, F.D. 1964. [Photoperiodic response of
the mite Typhlodromus (Amblyseius) similis (Ac-
arina: Phytoseiidae)]. Zool. zhurnal, 43:
1140–1144. [In Russian]

Saulich, A.Kh. and Musolin, D.L. 2007. [Yearth sea-
sons: diversity of seasonal adaptations and eco-
logical mechanisms controlling seasonal develop-
ment of true bugs (Heteroptera) in the temperate
climate]. Trudy biol. nauchno-issled. instituta St.
Peterburgskogo gos. universiteta, 53: 25–106. [In
Russian, Engl. Summary]

Schwarz, H.H. and Koulianos, S. 1998. When to leave
the brood chamber? Routes of dispersal in mites
associated with burying beetles. Exper. Appl. Ac-
arology, 22: 621–631.

Serdyukova, G.V. 1951. [Hibernation of Ixodes ricinus
eggs in conditions of Karelian peninsula]. Dokla-
dy AN SSSR, 81: 1171–1173. [In Russian]

Short, N.J. and Norval, R.A.I. 1981. Regulation of sea-
sonal occurrence in the tick Rhipicephalus appen-
diculatus Neum. Tropical Animal Health and Pro-
duction, 13: 19–26.

Smith, C.N. and Cole, M. 1941. Effect of length of day
on the activity and hibernation of the American
dog tick Dermacentor variabilis. Ann. Ent. Soc.
Amer., 34: 426–431.

Sonenshine, D.E. 1988. Diapause in tick vectors of
desease. In: The Arboviruses: epidemiology and
ecology, vol.1, p. 219–243.

Sonenshine, D.E. 1991. Biology of Ticks. Oxford. Vol.
1, 472 pp; 1993, Vol. 2, 488 pp.

Speybroeck, N., Madder, M., Van den Bossche, P.
Mtambo, J., Berkvens, N., Chaka, G., Mulumba,
M., Brandt, J., Tirry, L., and Berkvens D. 2002.
Distribution and phenology of ixodid ticks in
southern Zambia. Medical and Veterinary Ento-
mology, 16: 430–441.

Speybroeck, N., Lindsey, P.J., Billiouw, M., Madder,
M., Lindsey, J. K., and Berkvens, D.L. 2006.
Modeling diapause termination of Rhipicephalus
appendiculatus using statistical tools to detect
sudden behavioral changes and time dependen-
cies. Environ. Ecol. Stat., 13: 69–87.

Sutherst, R.W. and Bourne, A.S. 1991. Development,
survival, fecundity and behavior of Haemaphysa-
lis longicornis at two locations in Southeast
Queensland. Intern. J. Parasitol., 21: 661–672.

Sutherst, R.W. and Moorhouse, D.E. 1972. The sea-
sonal incidence of ixodid ticks on cattle in South-
eastern Queensland. Austral. J. Agr. Res., 23:
Tauber, M.J., Tauber, C.A. and Masaki, S. 1986. Sea-
sonal Adaptations of Insects. Oxford Univ. Press,
411 pp.

Ushatinskaya, R.S. 1976. Insect dormancy and its clas-
sification. Zool. Jahrbücher, Abt. Systematik, 103:

Vail, S., Smith, G. and Lord, C. 1994. Population biol-
ogy of lxodes scapularis, the vector of Lyme Dis-
ease in the Eastern and North Central United
States. Parasitic and Infectious Diseases, 19:

Van Houten, Y.M., and Veenendaal, R.L. 1990. Effects
of photoperiod, temperature, food and relative hu-
midity on the induction of diapause in the preda-
tory mite Amblyseius potentillae. Exp. Appl. Ac-
arol., 10: 111–128.

Veerman, A. 1985. Diapause. In: W. Helle and M.W.
Sabelis (Eds). Spider Mites. Their Biology, Natu-
ral Enemies and Control. Amsterdam, Vol. 1A,
pp. 279–316.

Veerman, A. 1992. Diapause in phytoseiid mites: a re-
view. Exper. Appl. Acarology, 14: 1–60.

Wainstein, B.A. 1977. [Family Phytoseiidae (Gamasi-
na)]. In: M.S. Gilyarov and N.G. Bregetova (Eds.).
Opredelitel’ kleshchey, obitayushchikh v potchve.
Mesostigmata. Leningrad, pp. 226–243. [In Rus-

Walker, J.B., Keirans, J.E. and Horak, I.G. 2000. The
Genus Rhipicephalus (Acari, Ixodidae). A guide to
the brown ticks of the world. Cambridge Univ.
Press, 635 pp.

Walter, D.E. and Proctor, H.C. 1999. Mites. Ecology,
Evolution and Behaviour. UNSW Press, CABI
Publishing, 322 pp.

Weigmann, G. 1996. Zonation and phenology of
Uropodina (Uropodidae) in marine littoral salt-
marches. In: Acarology IX (Proceedings), vol. 1,
Prague, Academia, p. 603–605.

Wilkinson, P.R. 1968. Phenology, behaviour and host-
relations of Dermacentor andersoni in outdoor
“rodentaria” and in nature. Canad. J. Zool., 46:

Wilkinson, P.R. 1973. Termination of diapause in lab-
oratory-reared Dermacentor andersoni adults.
Proc. 3rd Intern. Congr. Acarology. Praha, pp.

Wright, J.E. 1969. Photoperiodic induction of diapause
in larvae of the winter tick, Dermacentor albipic-
tus. Ann. Entomol. Soc. Am., 62: 285–287.

Wright, J. E. 1971. Relationship of seasonal activity of
Dermacentor albipictus (Acarina: Ixodidae) to
photoperiod: a laboratory study. Ann. Entomol.
Soc. Am., 64: 1456–1458.

Yuval, B. and Spielman, A. 1990. Duration and regula-
tion of the developmental cycle of Ixodes dam-
mini. Journ. Med. Entomol., 27: 196–201.

Zakhvatkin, A.A. 1948. [Systematics of the genus Lae-
laps (Acarina, Parasitiformes) and problems of its
epidemiological significance]. Parazitol. sbornik,
Leningrad, 10: 51–76. [In Russian]

Zakhvatkin, A.A. 1952. [Division of Acarina in orders
and their places in the system of the Chelicerata].
Parazit. sbornik, Leningrad, 14: 5–46. [In Rus-

Zaslavsky, V.A. 1972. [Two-step photoperiodic reac-
tions as the basis for elaboration of a model of
photoperiodic control of development in arthro-
pods]. Entomol. obozrenie, 52: 217–239. [In Rus-

Zaslavsky, V.A. 1974. [The principles of photoperiodic
control of arthropod development]. Zhurnal obsh-
chey biologii, 35: 717–736. [In Russian]

Zaslavsky, V.A. 1984. Photoperiodicheskiy i tempera-
turny control razvitiya nasekomych [Photoperiod-
ic and temperature control of insect development].
Leningrad, 180 pp. [In Russian] [In English:
Springer, 1987]

Zein-Eldin, E.A. 1956. Studies on the legume mite,
Petrobia apicalis. Journal of Economical Ento-
mology, 49: 291–296.

Zemskaya, A.A. 1951. [Biology and development of
the poultry mite Dermanyssus gallinae due to its
epidemiologic importance]. Zool. zhurnal, 30:
51–63. [In Russian]

Zemskaya, A.A. 1973. Parazyticheskye gamazovye
kleshchy i ikh meditsinskoye znachenie [Parasitic
gamasid mites and their medical importance].
Moscow, 167 pp. [In Russian]

Zhang, Y.X., Zhang, Z.Q., Liu, Q.Y., and Lin, J.Z.
1999. Biology of Typhlodromus bambusae (Acari:
Phytoseiidae), a predator of Schizotetranychus
nanjingensis (Acari: Tetranychidae) injurious to
bamboo in Fujian, China. Systematic and Applied
Acarology, 4: 57–62.

Zhmaeva, Z.M. 1948. [The tick Haemaphysalis con-
cinna in Primor’e]. In: Voprosy kraevoy, obsh-
chey i exper. parazitologii. Moscow, pp. 46–55.
[In Russian]