2011. 19 (1)



Filimonova S.A.

About author:

Filimonova S.A., Zoological Institute, Russian Academy of Sciences, Universitetskaya emb., 1, 199034, St. Petersburg


The study summarizes data on the internal anatomy of the mites belonging to the parvorder Eleutherengona (Acariformes: Trombidiformes) mainly based on the families Tetranychidae, Cheyletidae, Syringophilidae, Myobiidae, and Demodicidae. The arrangement and functioning of the digestive tract, propodosomal glands, connective tissue, and reproductive system of both sexes are taken into account to reveal common features of the Eleutherengona as well as possible phylogenetically informative characters in particular families. The study shows that all the examined eleutherengone families demonstrate a mixture of relatively primitive and advanced features. The following most striking internal characters are common to all studied eleutherengone species. The postventricular midgut is represented by a simple tube-like excretory organ, which is usually in open connection with the ventriculus. The anal and genital
openings of the females are close to each other and located at the terminal end of the body. The glandular component of the testicular epithelium is absent or highly reduced, as well as the male accessory glands. The number of salivary glands is reduced or these glands are lost in some parasitic forms. The coxal glands are either devoid of the proximal filter sacculus or provided by a sacculus with a considerably reduced lumen. The coxal gland epithelium does not possess a regular brush border, showing high pinocytotic activity.


Akimov, I.A. and Gorgol, V.T. 1984. [Structure and
functions of the digestive system of Bakericheyla
chanayi (Cheyletidae, Trombidiformes), an ec-
toparasite of birds]. Parasitologiya, 18: 15–22. [In

Akimov, I.A. and Gorgol, V.T. 1990. Khishchnye i
paraziticheskie kleshchi cheyletidy. [Predaceous
and Parasitic Cheyletid Mites]. Naukova Dumka,
Kiev, 119 pp. [In Russian]

Alberti, G. 1973. Ernährungsbiologie und Spinnvermö-
gen der Schnabelmilben (Bdellidae, Trombidifor-
mes). Zeitschrift fűr Morphologie der Tiere, 76:

Alberti, G. 1974. Fortpflazungsverhalten und Fortpfla-
zungsorgane der Schnabelmilben (Acarina: Bdel-
lidae; Trombidiformes). Zeitschrift fűr Morpholo-
gie der Tiere, 78: 111–157.

Alberti, G. 1980a. Zur Feinstruktur der Spermien und
Spermiocytogenese der Milben (Acari). 2. Anacti-
notrichida. Zoologische Jahrbücher Abteilung fűr
Anatomie und Ontogenie der Tiere, 104: 77–138.

Alberti, G. 1980b. Zur Feinstruktur der Spermien und
Spermiocytogenese der Milben (Acari). 2. Actino-
trichida. Zoologische Jahrbücher Abteilung fűr
Anatomie und Ontogenie der Tiere, 104: 144–

Alberti, G. 2005. On some fundamental characteristics
in acarine morphology. Atti della Accademia Na-
zionale Italiana di Entomologica. R.A., 53: 315–

Alberti, G. and Coons, L.B. 1999. Acari: Mites. In:
F.W. Harrison and R.F. Foelix (Eds.). Microscop-
ic Anatomy of Invertebrates. Vol. 8C, Wiley-Liss,
New York, pp. 515–1217

Alberti, G. and Crooker, A.R. 1985. Internal anatomy.
In: W. Helle and M.W. Sabelis (Eds.). Spider
mites. Their Biology, Natural Enemies and Con-
trol. World Crop Pests. Vol. 1A. Elsevier, Amster-
dam, pp. 29–62.

Alberti, G., Seniczak, A. and Seniczak, S. 2003. The
digestive system and fat body of an early-deriva-
tive oribatid mite, Archegozetes longisetosus,
Aoki (Acari: Oribatidae, Trhypochthoniidae). Ac-
arologia, 43: 149–219.

Alberti, G. und Storch, V. 1973. Zur Feinstruktur der
Munddrüsen von Schnabenmilben (Bdellidae,
Trombidiformes). Zeitschrift für wissenschaftliche
Zoologie, 186: 149–160.

Alberti, G. und Storch, V. 1974. Über Bau und Func-
tion der Prosoma-Drüsen von Spinnmilben (Tet-
ranychidae, Trombidiformes). Zeitschrift fűr Mor-
phologie der Tiere, 79: 133–153.

Alberti, G. und Storch, V. 1976. Ultrastruktur — Un-
tersuchungen am männlichen Genitaltrakt und an
Spermien von Tetranychus urticae (Tetranychi-
dae, Acari). Zoomorphologie, 83: 283–296.

Alberti, G. und Storch, V. 1977. Zur Ultrastruktur der
Coxaldrüsen actinotricher Milben (Acari, Acti-
notrichidae). Zoologische Jahrbücher Abteilung
für Anatomie und Ontogenie der Tiere, 98: 394–

Alberti, G. und Storch, V. 1983. Zur Ultrastruktur der
Mitteldarmdrüsen von Spinnentieren (Scorpiones,
Araneae, Acari) unter verschiedenen Ernährungs-
bedingungen. Zoologischer Anzeiger, 3/4: 145–

Amosova, L.I. 1983. Tissues of the internal environ-
ment. In: Y.S. Balashov (Ed.). Atlas elektronno-
mikroskopicheskoi anatomii iksodovich kleschei.
[An atlas of ixodid tick ultrastructure]. Nauka, Le-
ningrad, pp. 147–174.

Anwarullah, M. 1963. Beiträge zur Morphologie und
Anatomie einiger Tetranychiden (Acari, Tetrany-
chidae). Zeitschrift für Angewandte Zoologie, 50:

Bader, C. 1938. Beiträge zur Kenntnis der Verdauungs-
vorgängebei Hydracarinen. Revue Suisse de Zoo-
logie, 45: 721–806.

Bader, C. 1954. Das Darmsystem der Hydracarinen.
Eine vergleichend anatomische Untersuchung.
Revue Suisse de Zoologie, 61: 505–549.

Berridge, M.J. 1970. Osmoregulation in Terrestrial Ar-
thropods. In: M. Florkin and B.T. Cheer (Eds.).
Chemical Zoology. Vol. 5A. Academic Press,
London, pp. 287–319.

Blauvelt, W.E. 1945. The internal anatomy of the com-
mon red spider mite Tetranychus telarius (L.).
Cornell University Experiment Station Memoirs,
270: 1–46.

Bochkov, A.V. 2002. [The classification and phylogeny
of the mite superfamily Cheyletoidea (Acari, Pros-
tigmata)]. Entomologicheskoye obozreniye, 81:
488–513. [In Russian]

Bochkov, A.V. 2009. A review of mites of the par-
vorder Eleutherengona (Acariformes: Prostigma-
ta) — permanent parasites of mammals. Acarina,
Supplement 1, 149 pp.

Bochkov, A.V., O’Connor, B.M. and Wauthy, G. 2008.
Phylogenetic position of the mite family Myobii-
dae within the infraorder Eleutherengona (Acari-
formes) and origins of parasitism in eleutheren-
gone mites. Zoologischer Anzeiger, 247: 15–45.

Brown, J.R.C. 1952. The feeding organs of the adult of
the common “chigger”. Journal of Morphology,
91: 15–52.

Castagnoli, M. 1974. La spermateca in alcune specie di
Brevipalpini (Acarina, Tenuipalpidae). Redia, 55:

Coons, L.B., Tarnowski, B. and Ourth, D.D. 1982. Rhi-
picephalus sanguinius: Localization of vitello-
genin synthesis by immunological methods and
electron microscopy. Experimental Parasitology,
54: 331–339.

Coons, L.B., L’Amoreaux, W.J., Rossel-Davis, R., and
Star-Spires, L. 1990. Fine structure of the fat body
and nephrocytes in the life stages of Dermacentor
variabilis. Experimental and Applied Acarology,
8: 125–142.

Crooker, A.R. and Cone, Jr. W. 1979. Structure of the
reproductive system of the adult female two-spot-
ted spider mite Tetranychus urticae. In: J.G. Ro-
driguez (Ed.). Recent Advances in Acarology.
Vol. II. Academic Press, New York, Saint-Fran-
cisco, London, pp. 405–411

Desch, C. E. Jr. 1984. The reproductive anatomy of De-
modex folliculorum. In: D.A. Griffiths and C.E.
Bowman (Eds.). Acarology. Vol.VI. Ellis Hor-
wood, Chichester, pp. 464–469.

Desch, C. E. Jr. 1988. The digestive system of Demo-
dex folliculorum (Acari: Demodicidae) of man: A
light and electron microscope study. In: G.V.
Channabasavanna and C.A. Viraktamath (Eds.).
Progress in Acarology. Vol. 1. New Delhi, Ox-
ford, pp. 187–195.

Desch, C.E. and Nutting, Wm. B. 1977. Morphology
and functional anatomy of Demodex folliculorum
(Simon) of Man. Acarologia, 19: 422–462.

Di Palma, A., Nuzzaci, G. and Alberti A., 2009. Mor-
phological, ultrastructural and functional adapta-
tions of the mouthparts in cheyletid mites (Acari:
Actinedida: Prostigmata). International Journal
of Acarology, 35, 521–532.

Dosse, G. and Langenscheidt, M. 1964. Morphologi-
sche, biologische und histologische Untersuchun-
gen an Hybriden aus dem Tetranychus urticae —
complex (Acari, Tetranychidae). Zeitschrift für
Angewandte Entomologie, 54: 349–359.

El Shoura, S.M. 1986. Fine structure of haemocytes
and nephrocytes of Argas (Persicargas) arboreus
(Ixodoidea: Argasidae). Journal of Morphology,
189: 17–24.

El Shoura, S.M. 1989. Ultrastructure of larval haemo-
cytes and nephrocytes in the tick Ornythodoros
(Pavlovskyella) erraticus (Ixodoidea: Argasidae).
Acarologia, 30: 35–40.

Ehrnsberger, R. 1984. Anatomie des Verdauungstrak-
tes der Rhagidiidae (Acari, Trombidiformes). Os-
nabrücker Naturwissenschaftliche Mitteilungen,
11: 67–90.

Feiertag-Koppen, C.C.M. and Pijnacker, L.P. 1982.
Development of the female germ cells and process
of internal fertilization in the two-spotted spider
mite Tetranychus urticae Koch. (Acariformes,
Tetranychidae). International Journal of Insect
Morphology and Embryology, 11: 271–284.

Filimonova, S.A. 1999. [Organization of the female re-
productive system in the parasitic mite Myobia
murismusculi (Trombidiformes: Myobiidae)].
Parasitologiya, 33: 377–386. [In Russian]

Filimonova, S.A. 2001a. Internal anatomy of the fe-
males of Myobia murismusculi (Schrank) (Acari:
Trombidiformes, Myobiidae). Entomological Re-
view, 81: 1049–1058.

Filimonova, S.A. 2001b. [The fine structure of the
midgut in the mite Myobia murismusculi]. Tsy-
tologiya, 43: 425–431. [In Russian]

Filimonova, S.A. 2001c. [The cells — analogous to fat
body in parasitic mites Myobia murismusculi
(Schrank) (Trombidiformes: Myobiidae)]. Zhur-
nal evolutsionnoy biokhimii i fisiologii, 37: 329–
334. [In Russian]

Filimonova, S.A. 2004. The fine structure of the coxal
glands in Myobia murismusculi (Schrank) (Acari:
Myobiidae). Arthropod Structure and Develop-
ment, 33: 149–160.

Filimonova, S.A. 2006. An ultrastructural investigation
of the testes and spermiogenesis in Myobia muris-
musculi (Schrank) (Acari, Actinedida: Myobii-
dae). Invertebrate Reproduction and Develop-
ment, 49: 273–283.

Filimonova, S.A. 2008a. The fine structure of the midg-
ut in the mite Anystis baccarum (L.) (Acari, Actin-
edida: Anystidae). Arthropod Structure and De-
velopment, 37: 299–309.

Filimonova, S.A. 2008b. [Peculiarity of the internal
anatomy in the mite family Syringophilidae by the
example of Syringophilopsis fringilla a parasite of
chaffinch]. Parazitologiya, 42: 395–404. [In Rus-

Filimonova, S.A. 2009. The ultrastructural investiga-
tion of the midgut in the quill mite Syringophilop-
sis fringilla (Acari, Trombidiformes: Syringophil-
idae). Arthropod Structure and Development, 38:

Filimonova, S.A. and Mironov, S.V. 2010. Functional
morphology of the gnathosoma in the quill mite
Syringophilopsis fringilla Fritsch (Acari, Prostig-
mata: Syringophilidae). Zoologischer Anzeiger,
249 (3–4): 165–180.

Gorgol, V.T. and Barabanova, V.V. 1979. [Activity of
some digestive enzymes as a sign of predation of
the mite Cheyletus eruditus (Trombidiformes,
Cheyletidae)]. Trudy Ukrainskoy Akademii Nauk.
Ser. B., 10: 855–858. [In Russian]

Grandjean, F. 1938. Observations sur les Bdelles. An-
nales de la Sociéte Entomologique de France,
107: 1–24.

Hafiz, H.A. 1935. The embryological development of
Cheyletus eruditus. Proceedings of the Royal So-
ciety of London, 117: 174–201.

Jonczy, J. and Kropczynska, D. 1974. Fine structure of
the cuticle of Aceria macrorhynchus (Nal.) (Ac-
arina: Eriophyidae). Acta Biologica Cracov. Ser.
Zool., 17: 227–223.

Kethley, J. 1982. Acariformes. In: S.P. Parker (Ed.).
Synopsis and Classification of Living Organisms.
Vol. 2. McGraw-Hill, New York, pp. 119–146.

Klumpp, W. 1954. Embryologie und Histologie der Bi-
enenmilbe Acarapis woodi Rennie 1921. Zeitschrift
fűr Parasitenkunde, 16: 407–442.

Krantz, G. W. and Walter, D. E. 2009. A manual of
acarology. Third edition. Lubbock: Texas Tech
University Press, 807 pp.

Langenscheidt, M. 1973. Zur Wirkungsweise von Ster-
ilität erzeugenden Stoffen bei Tetranychus urticae
Koch (Acari, Tetranychidae). 1. Entwicklung der
weiblichen Genitalorgane und Oogenese bei un-
behandelten Spinnmilben. Zeitschrift für Ange-
wandte Entomologie, 73: 103–106.

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-
ume 1. Ellis Horwood, Limited, Chichester, pp.

Mc Enroe, W.D. 1961. Guanine excretion by the two-
spotted spider mite Tetranychus telarius. Annals
of the entomological society of America, 54: 925–

Mathur, S.N. and Le Roux, E.J. 1970. The reproductive
organs of the velvet mite, Allotrombium lerouxi
Moss (Trombidiformes, Trombidiidae). Canadian
Entomologist, 102: 144–157.

Matsubara, T., Ohta, Y. and Ehara, Sh. 1992. Fine
structure of female and male internal reproductive
organs in a spider mite Tetranychina harti (Ew-
ing) (Acari: Tetranychidae). Applied Entomology
and Zoology, 27: 65–78.

Mills, L.R. 1973. Morphology of glands and ducts in
the two-spotted spider mite, Tetranychus urticae
Koch, 1836. Acarologia, 15: 218–236.

Mironov, S.V. and Bochkov, A.V. 2009. [Modern con-
ceptions of macrophylogeny in acariform mites
(Chelicerata, Acariformes)]. Zoologichesky Zhur-
nal, 88: 922–937. [In Russian]

Mitchell, R.D. 1955. Anatomy, Life History and Evolu-
tion of the Mites Parasitizing Fresh Water Mus-
sels. Miscellaneous Publications of Museum of
Zoology. Vol. 89. University of Michigan, USA,
41 pp.

Mitchell, R.D. 1964. The anatomy of the adult chigger
mite Blankaartia ascoscutellaris (Walch.). Jour-
nal of Morphology, 114: 373–391.

Mitchell, R. 1970. The evolution of a blind gut in trom-
biculid mites. Journal of Natural History, 4: 221–

Moss, W.W. 1962. Studies on the morphology of the
trombidiid mite Allotrombium lerouxi Moss (Ac-
ari). Acarologia, 4: 313–345.

Mothes, U. and Seitz, K. 1980. Licht- und elektronmik-
roskopische Untersuchungen zur Funktionsmor-
phologie von Tetranychus urticae (Acari, Tetrany-
chidae). Exkretionssysteme.Zoologische
Jahrbücher für Anatomie, 104: 500–529.

Mothes, U. and Seitz, K. 1981a. Functional microscop-
ic anatomy of the digestive system of Tetranychus
urticae (Acari, Tetranychidae). Acarologia, 22:

Mothes, U. and Seitz, K.A. 1981b. Fine structure and
function of the prosomal glands of the two-spotted
spider mite, Tetranychus urticae (Acari, Teytrany-
chidae). Cell and Tissue Research, 221: 339–349.

Mothes, U. and Seitz, K.A. 1981c. The transformation
of male sex cells of Tetranychus urticae K. (Acari,
Tetranychidae) during passage from the testis to
the oocytes: an electron microscopic study. Inter-
national Journal of Invertebrate Reproduction, 4:

Mothes, U. and Seitz, K.A. 1981d. Licht und elek-
tronenmikroskopische Untersuchungen zur Funk-
tionsmorphologie von Tetranychus urticae (Acari,
Tetranychidae). II. Weibliches Geschlechtssystem
und Oogenese. Zoologische Jahrbücher Abteilung
für Anatomie und Ontogenie der Tiere (Jena),
105: 106–134.

Mothes-Wagner, U. 1985. Fine structure of the
‘hindgut’ of the two-spotted spider mite, Tetrany-
chus urticae, with special reference to origin and
function. Experimental and applied acarology, 1:

Mothes-Wagner, U. and Seitz, K.A. 1984. Ultrahistol-
ogy of oogenesis and vitellogenesis in the spider
mite Tetranychus urticae. Tissue and Cell, 16:

Norton, R.A., Kethley, J.B., Johnston, D.E., and OCon-
nor, B.M. 1993. Phylogenetic perspectives on ge-
netic 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 & Hall, NewYork, London, England,
pp. 8–99.

Nuzzaci, G. and Alberti, G. 1996. Internal anatomy and
physiology In: E.E. Lindquist, M. W. Sabelis and
J. Bruin (Eds.). Eriophyoid Mites — Their Biolo-
gy, Natural Enemies and Control World Crop
Pests. Vol. 6. Elsevier, Amsterdam, pp. 101–150.

Oliver, J.H.Jr. 1983. Chromosomes, Genetic variance
and reproductive strategies among mites and ticks.
Bulletin of the Entomological Society of America,
29: 8–17.

Paran, T.P. 1979. Evolution of the excretory duct in
Myobia murismusculi (Schrank, 1781), Myobii-
dae (Trombidiformes — Acarina). Acarologia,
21: 55–60.

Paran, T.P. 1982. Functional morphology of the gna-
thosoma in Myobia musculi (Schrank, 1781)
(Trombidiformes — Acari). Acarologia, 23: 347–

Pijnacker, L.P. 1985. Spermatogenesis. In: W. Helle and
M.W. Sabelis (Eds.). Spider mites. Their Biology,
Natural Enemies and Control World Crop Pests.
Vol. 1A. Elsevier, Amsterdam, pp. 109–115.

Pijnacker, L.P. and Drenth-Diephuis, L.J. 1973. Cyto-
logical investigations on the male reproductive
system and sperm track in the spider mite Tetrany-
chus urticae Koch (Tetranychidae, Acarina).
Netherland Journal of Zoology, 23: 446–464.

Pijnacker, L.P., Ferwerda, M.F. and Helle, W. 1981.
Cytological investigations on the female and male
reproductive system of the parthenogenetic privet
mite Brevipalpus obovatus Donnadieu (Phytopti-
palpidae, Acari). Acarologia, 22: 157–163.

Shatrov, A.B. 1989a. [The principal peculiarities of the
midgut structure of the active stages of the life
cycle of trombiculid mites (Acariformes, Tromb-
iculidae)]. Zoologichesky Zhurnal, 68: 28–37. [In

Shatrov, A.B. 1989b. [Salivary glands organization in
fed deutonymphs and adult mites Hirsutiella zach-
vatkini Schlug. (Acariformes, Trombiculidae)].
Entomologicheskoye obozreniye, 68: 882–893. [In

Shatrov, A.B. 1995. [Fine structural organization and
functional peculiarities of tubular glands in tromb-
iculid mites (Acariformes, Trombiculidae)]. Para-
sitologiya, 29: 449–458. [In Russian]

Shatrov, A.B. 1997. Vitellogenesis and egg-shell forma-
tion in ovipositing females of the trombiculid mite
Hirsutiella zachvatkini (Schluger) (Acariformes:
Trombiculidae). Acarologia, 38: 141–151.

Shatrov, A.B. 1998. The ultrastructure and possible
functions of nephrocytes in the trombiculid mite
Hirsutiella zachvatkini (Acariformes: Trombiculi-
dae). Experimental and Applied Acarology, 22:

Shatrov, A.B. 2000. Krasnotelkovye kleshchi i ich
parazitizm na pozvonochnykh zhivotnykh. [Trom-
biculid mites and their parasitism on vertebrate
hosts]. Proceedings of Zoological Institute. Vol.
285, Saint Petersburg, 276 pp. [In Russian]

Shatrov, A.B. 2002. Oogenesis in ovipositing females
of the microtrombidiid mite Platytrombidium fas-
ciatum (C.L. Koch) (Acariformes: Microtrombi-
diidae). Invertebrate Reproduction and Develop-
ment, 42: 1–15.

Shatrov, A.B. 2003. Comparative midgut ultrastructure
of unfed larvae and adult mites of Platytrombidi-
um fasciatum (C.L. Koch, 1836) and Camerotrom-
bidium pexatum (C.L. Koch, 1837) (Acariformes:
Microtrombidiidae). Arthropod Structure and De-
velopment, 32: 227–239.

Shatrov, A.B. 2005. Ultrastructural investigation of the
salivary glands in adults of the microtrombidiid
mite Platytrombidium fasciatum (C.L.Koch, 1836)
(Acariformes: Microtrombidiidae). Arthropod
Structure and Development, 34: 49–61.

Shatrov, A.B. 2006. Ultrastructure of coxal glands in
the unfed microtrombidiid larvae, Platytrombidi-
um fasciatum (C.L. Koch, 1836) and Camerotrom-
bidium pexatum (C.L. Koch, 1837) (Acariformes:
Microtrombidiidae). Abhandlungen und Berichte
des Naturkundemuseums Görlitz, 78: 57–78.

Shatrov, A.B. 2007. [Electron-microscopic organiza-
tion of the midgut and excretory organ in the adult
water mite, Teutonia cometes (Koch, 1837) (Para-
sitengona: Hydrachnidia: Teutonidae) and its evo-
lutionary implication]. Zoologiya bespoz-
vonochnych, 4: 195–208. [In Russian]

Shatrov, A.B. 2010. Comparative morphology, ultra-
structure and functions of the excretory organ
(postventricular midgut) in the Parasitengona (Ac-
ariformes). Acarologia, 50: 93–112.

Schmidt, U. 1935. Beiträge zur Anatomie und Histolo-
gie der Hydracarinen, besonders von Diplodontus
despiciens O.F.Müller. Zeitschrift für Morpholo-
gie und Ökologie der Tiere, 30: 99–176.

Smith, J. W. and Boudreaux, H.B. 1972. An autoradio-
graphic search for the site of fertilization in spider
mites. Annals of Entomological Society of Ameri-
ca, 65: 69–74.

Šobotnik, J., Alberti, G., Weyda, F., and Hubert, J.
2008. Ultrastructure of the digestive tract in Ac-
arus siro (Acari, Acaridida). Journal of Morphol-
ogy, 269: 54–71.

Summers, F.M. and Witt, R.L. 1971. The gnathosoma
of Cheyletus cachuamilpensis Baker (Acarina:
Cheyletidae). Proceedings of Entomological Soci-
ety of Washington, 73: 158–168.

Vistorin, H.E. 1980. Ernährungsbiologie und Anatomie
des Verdauungstraktes der Nicoletiellidae (Acari,
Trombidiformes). Acarologia, 21: 204–215.

Vistorin-Theis, G. 1977. Anatomische Untersuchungen
an Calyptostomiden (Acari, Trombidiformes). Ac-
arologia, 19: 242–257.

Weyda, F. 1980. Reproductive system and oogenesis in
active females of Tetranychus urticae (Acari, Tet-
ranychidae). Acta Entomologica Bohemoslovica,
77: 375–377.

Whitmoyer, R.E., Nault, L.R. and Bradfute, O.E. 1972.
Fine structure of Aceria tulipae (Acarina: Erio-
phyidae). Annals of the Entomological Society of
America. 65: 201–215.

Witte, H. 1975. Funktionsanatomie des weiblichen
Genitaltraktes und Oogenese bei Erythraeiden
(Acari, Trombidiformes). Zoologische Beiträge,
21: 247–277.