A new eviphidid mite genus (Parasitiformes; Mesostigmata;
Eviphididae) associated with the dung beetle Scarabaeus
transcaspius Stolfa (Coleoptera, Scarabaeidae) in
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(PARASITIFORMES; MESOSTIGMATA; EVIPHIDIDAE)
IN TURKMENISTAN
SCARABAEUS
PHORESY
SUMMARY: A new eviphidid mite, Cryptoseius petrovae gen. et sp.
nov., was found under elytra of the beetle Scarabaeus transcaspius
Stolfa from Turkmenistan. Descriptions of the female, male and
deutonymph are given. The new genus displays a significant
superficial resemblance to Scamaphis Karg (Eviphididae) and
Coleolaelaps Berlese (Laelapidae), also associated with large
scarabaeid beetles. A new combination, Coleo laelaps anoxiae comb.
n. (Laelapidae) for Pelethiphis anoxiae Koyumdjieva, 1977 (Evi
phididae), is suggested.
EVIPHIDIDAE
SCARABAEUS
PHORESIE
RF:suMil : Un nouvel eviphidide, Cryptoseius petrovae n. gen., nov.
sp., s'est revele sous les elytres de Scarabaeus transcaspius
Stolfa, au Turkmenistan. On don ne les descriptions de la femelle,
du male et de la deutonymphe. Le nouveau genre presente une
ressemblance superficielle significative avec Scamaphis Karg
(Eviphididae) et Coleolaelaps Berlese (Laelapidae), eux aussi
associes a de grands coleopteres scarabaeides. Pour Pelethiphis
anoxiae Koyumdjieva, 1977 (Eviphididae), une combinaison nouvelle,
Coleo/ae/aps anoxiae comb. n. (Laelapidae), est suggeree.
Gamasid mites of the family Eviphididae occupy various substrates
and biotopes, but most are asso ciated with discrete temporary
habitats, such as ver tebrate droppings, nests and carrion, as
well as sea debris. Many species are stenobiontic and occupation of
the proper substrate is facilitated by phoresy on appropriate
arthropods.
Phoretic specificity among eviphidid species varies. For example,
Crassicheles holsaticus (Willmann) has been found phoretic both on
different dipterans and on staphylinid beetles. Some species of
Alliphis Hal bert and Scarabaspis Womersley, which live in drop
pings, may be distributed by any coprophilous scara baeid beetles.
Crassicheles concentricus Karg can be
found on many different small dipt~rans. On the contrary, Scamaphis
equestris (Berlese) and Alliphis necrophilus Christie are evidently
associated with beetles of particular genera (respectively
Geotrupes Latreille and Nicrophorus F.), whereas phoresy of
Pelethiphis gurei (Costa), Pelethiphis opacus Koyu mdjieva and
Eviphis pterophilus Berlese is apparently species specific.
During a study of mites occurring on different species of
Scarabaeus, several new eviphidid species were found, including a
representative of a new, mor phologically unusual genus,
Cryptoseius gen. n., which is described below. It was found only on
Sca rabaeus transcaspius Stolfa, collected at two loca-
* Severtsov Institute of Ecology and Evolution, Russian Academy of
Sciences, Leninsky pr. 33, Moscow 117071, Russia.
Acarologia, t. XXXIX, fasc. 2, 1998.
- 116- tions in Turkmenistan in different years. It is worth noting
that Scarabaeus transcaspius is also character ized by significant
morphological and ecological spe cificity, being considered as the
species most adapted to the conditions of sandy deserts among
Scarabaeus L. (KABAKOV, 1980).
The description presented below is based on mate rial collected by
the coleopterologists T. A. CHER NYAKHOVSKAYA and K. V. MAKAROV.
All measure ments are given . in micrometers. The setal
designations follow those of LINDQUIST & EvANS (1965) and
LINDQUIST (1995) for the idiosoma, and those of EvANS (1963) for
the legs. Setal homologies were established regarding the
disposition in relation to location of pores and the variations in
setal arran gements. The holotype and part of the paratypes series
are deposited in the collection of the Zoological Institute of the
Russian Academy of Sciences, St. Petersburg. Other paratypes are in
the author's col lection.
Cryptoseius gen. nov.
Type species: Cryptoseius petrovae sp. nov. Mono typic.
Diagnosis. Idiosoma of adults elongate-oval, sli ghtly convex;
sclerites of body and extremities yellow. Dorsal shield reduced,
not covering whole body, with incomplete set of setae. Setae s2,
r2, r3 inserted ante rodorsally on shield elements which are fused
with peritremat~c extremities; main part of shield bears 20 pairs
of smooth setae of various lengths. Setae jl small, not standing
out against background of other frontal setae. Female sterna!
shield well sclerotized, bearing three pairs of setae and three
pairs of pores; first pair of pores oval, oriented transversally,
located behind setae Stl . Metasternal setae free on integu ment.
Peritremes strongly shortened rather broad; peritrematic shields
rudimentary; exopodal shields lost. Tectum triangular, with dentate
margin, not pilose. Hypognathal groove narrow, with 8 rows of
denticles. Labrum, supralabral process and internal malae of
hypostome strongly developed, apically pointed. Female chela small,
gracile, with some sharp denticles as illustrated (Fig. A 6). Male
d. m. tube-like, spermatodactyl canal extends inside digit.
Palpal
apotele two-tined. Chaetotactic formula for legs I, II, II, IV
respectively: femora (2-311, 2/3-2) (2-311, 2/2-1) (1-3/2-1)
(1-2/1, 1/1-1); genua (1-3/2, 2/1-2) (2-3/1, 2/1-2) (1-2/1, 211-1)
(1-2/1, 2/1-1); tibiae (1-3/2, 2/1-2) (2-2/1, 2/1-2) (1-1/1, 2/1-1)
(1-111, 2/1-1). Femur and tarsus II in male each supplied with a
small, roundish apophysis of setal origin. Sexual dimorphism not
sharply expressed. Deutonymphs resemble adults. Females, males and
deutonymphs phoretic. Associa ted with lamellicorn
Scarabaeidae.
At present, the new genus is placed in the subfamily Eviphidinae
Berlese (sensu EVANS, 1957). The diffe rential diagnosis given
here is rather formal because of the necessity to clarify the
subfamily structure of Eviphididae in general (MAKAROVA, 1996a) and
to revise some compounded taxa (e.g. Pelethiphis Ber lese and
Alliphis Halbert). It is given in comparison with genera of the
subfamily Eviphidinae recognized by EVANS & TILL (1979) and
MAMN (1993), exclu ding Iphidosoma Berlese, Rhodacaridae (KETHLEY,
1983).
Differential diagnosis. The degree of the reduction of the dorsal
shield and its setal set in both females and males, the
chaetotactic pattern of femur and genu IV (Fig. A 3), as well as
the tecta! structure are unique in Eviphididae. In contrast to most
Eviphidi dae (except Scamaphis equestris), the peritreme is
shortened and the peritrematic shield is rudimentary. As distinct
from Scamaphis Karg, which shares with the new genus some other
marked characteristics (the elongated form of the body, the
reduction of the dorsal shield and its setal set, the broad
shortened peritremes, etc.), Cryptoseius gen. n. shows a different
orientation of the first sterna! pores (perpendicular to the body
axis), a narrow hypognathal groove, posses sing 8 rows of
denticles, and strong expressed cheli cera! sexual dimorphism; in
the female the third ster na! pores are situated on the sterna}
shield, the setae Mst are inserted on the integument, and the
opistho gastric cuticle has only 4 pairs of setae (larval pat
tern); in the male the leg apophyses are only of setal origin,
present on the femur and tarsus II (in Scama phis in addition to
the enlarged setae on tarsi II-IV, there is a large cuticular
process on trochanter IV).
Etymology. The generic name applies to the locali zation of the
mite during phoresy-under the elytra of beetles.
I j3
,, ~ ;,..., s2. r/·~v-· .· .,...... - ---s: .;r·./ OQw Qo ,.,
t
zS ( f
.14 I •
FIG. A: Cryptoseius petrovae gen. sp. n., female.
l. - Dorsum of idiosoma. 2. - Venter of idiosoma. 2a. - Variant of
posterior margin structure of sterna! shield. 3.- Femur and genu
IV. 4. - Hypostome. 5.- Telotarsus IV with ambulacrum. 6. -
Chelicera. Scales: 1 & 2 = 100 ~m; 3- 6 = 50 ~m.
Cryptoseius petrovae sp. n.
Material: Holotype: female-Turkmenistan, Amu Darya R., 110 km
lower Chardzhou, Nargyz Isl. (Amu-Darya Reserve), under elytra of
Scarabaeus
transcaspius Stolfa, 16 June 1988 (T. A. CHER NYAKHOVSKAYA);
paratypes: 17 females, 6 males, 6 deutonymphs, same locality and
data, 2 females, 1 male- Turkmenistan, East Kara-Kum, env. Repetek
railway station (Repetek Biosphere Reserve), under elytra of
Scarabaeus transcaspius, 9 June 1989 (K. V. MAKAROV).
Description: Medium-sized ; elongate mites. Scleri tes of body and
extremities yellow in adults. Dorsal shield not covering entire
body, leaving broad pleu rae. Reticulate ornamentation of shields
poorly deve loped; they are very slightly granulated.
FEMALE. Dorsum (Fig. A 1): Anterolateral parts of dorsal shield
separated from its main part and fused with peritrematic
extremities. Main part of dorsal shield (496-568 x 264-304)
commonly with 20 pairs of smooth, heteromorphic setae. Dorsal
chaetotaxy unstable, often asymmetrical; certain setae may be
absent (for example, seta J4 asymmetrically lost in one in four
females), doubled, or have migrated on shield and behind it. Setal
length varies greatly. Setae z3, i6, sl, s3, r4, r6, J2, J3, Zl,
Z2, SI, S3, R2 and R4 always absent. Anterolateral shield elements
bear setae s2, r 2, r 3 (24-72). Shortest setae on dorsal shield
are z5, Jl, J4, J5, Z3, Z4 (9-17), zl 16-22 andjl-2 (21-40). Most
dorsal macrosetae range from 94-176, including setae s6 and Z5
inserted on soft cuticle, which are conspicuously longer than r5
(35-68), RI (28-56), R3 (28-50) and R5 (46-74).
Venter (Fig. A 2): Tritosternum with massive base (length 28-40)
and pilose laciniae (84-1 08). Praester nal region weakly
sclerotized. All setae on ventral surface hair-like distally.
Sternum normally develo ped, fused with endocoxal shields, with 3
pairs of setae and 3 pairs of oval pores; form of posterior margin
of sternal shield variable (Fig. B2a); reticula tion on its
wrinkled surface only visible anteriorly. Metasternal setae free on
integument; Stl, St2 (30-42) always shorter than St3 and M.st
(40-58). Genital shield widened and rounded posteriorly, bea ring
a pair of setae (48-66), its anterior hyaline exten-
118 - si on roundish, not extending to sternal shield margin.
Solenostomes situated posteriorly in acetabula of coxae Ill.
Metapodal shields irregularly oval. Opis thogastric cuticle with
only 4 pairs of setae, JVJ and JV2 ( 40-58) always longer than Z V2
(32-48) and JV5
(20-24). Anal shield pear-shaped, twice longer than broad (117 x
64), its granulation developed mainly laterally, pores (2 pairs)
hardly visible, cribrum very extensive, perianal setae 28-40.
Peritremes rather broad (width 14-16), strongly shortened, reaching
only midlevel of coxae II; peritrematic shields nar row.
Gnathosoma: Tectum triangular (Fig. B 6a, b), with dentate margin,
highly variable. Hypognathal groove (Fig. A 4) narrow (width 7-9),
with 8 rows of deQticles (1-4 in a row). Hypostomatic setae needle
like, Cl 24-28, CJ 32-38, C2, C4 14-21. Corniculi triangular, 37-40
x 13-15, with lateral gutters containing salivary styli". In!ernal
malae of hypes tome long, reaching midlevei of genu of palp,
fringed laterally. Labrum well developed, almost reaching level of
palp apex, pilose; supralabral process narrow, long, sharply
pointed. Setae a! of palp femur and genu needle-like. Chelicera
small, delicate (Fig. A 6), length of chela 56-61; d.f with two
teeth of different size, its tip split, pilus dentilis short,
setiform; d.m. with one large tooth and one small subapical
denticle.
Legs: All legs shorf~r than dorsal shield (I 382-420, II 374-394,
Ill 370-406, IV 462-516); length of tar sus I 104-108, tarsus IV
140-160. Chaetotactic for mula legs I, II, Ill, IV, respectively:
femora (2-3/1, 2/3-2) (2-3/1, 2/2-1) (1-3/2-1) (1-2/1, 1/1-1);
genua (1-3/2, 2/1-2) (2-3/1, 211-2) (1-211, 211-1) (1-2/1, 211-1);
tibiae (1-3/2, 2/1-2) (2-2/1, 2/1-2) (1-111, 2/1-1) (1-111, 2/1-1).
Unlike the general type of leg chaetotaxy in Eviphididae (EVANS,
1963); femur Ill in this species bears additional spiniform seta
v2, femur IV and genu IV with additional spiniform setae pv (Fig. A
3). External spurs on distal margins of coxae II-IV lar ger than
internal ones. Most leg setae thickened, some dorsal ones on femora
and genua I-IV are macrosetae as shown in Fig. B 2,5. Femora I-II
with 2 macrosetae (adl ,pdl), femora III-IV with 1 macro seta
(adl). Genu I with 2-4 macrosetae (always pdl, pd2, sometimes adl,
ad2), g~nua II-IV with 2 macro setae (genu II with pdl, pdi; genua
III-IV with adl, pdl). Length of longest leg macro seta (pdl on
the
ll H> (JQ ..... CON
FIG. B: Cryptoseius petrovae gen. sp. n.
1- 5: Male; 6: Female; 7: Deutonymph. 1. - Venter of idiosoma.
2.-Trochanter, femur, genu and tibia I. 3.- Supralabral process. 4
a, b. Chelicera in different positions. 5. - Leg II without coxa.
6 a, b. - Variants of tectum form. 7. - ldiosoma. Scales: 1, 5
& 7 = 100 flm; 2-4
& 6 = 50 f!m.
-120- femur II) 72-100. One of the subapical ventral setae on tarsi
II-IV modified into strongly sclerotized, api cally tridentate
blade, evidently used as support (Fig. 1, 5). All leg claws
weak.
MALE. Dorsum: Main part of dorsal shield 456-480 x 256-282. Dorsal
chaetotaxy generally as in female, but most of those setae inserted
on soft cuticle much shorter (r5, RI, R3 12-20).
Venter (Fig. B 1): Tritosternum short (16-20), laci niae 72-80. On
sternogenital shield, setae Stl-3 (20-27) always shorter than St4-5
(24-38); opistho gastric cuticle with 4 pairs of setae; JVJ, JV2
(22-34) always longer than ZV2 (19-21) and JV5 (13-18); perianal
setae 22-30. Left and right peritremes of different length in some
individuals.
Gnathosoma: Tectum as in female. Deutosternal groove broadens
anteriorly. Labrum consisting of two laminae; wide pellicle-like
blade lies under long narrow one (as in female) . Subcheliceral
plate nar rows sharply in supralabral process (Fig. B 3). Chela
length 52-54; d.f toothless, with beak-like tip (Fig. B 4); d.m.
larger than df, cylindrical, bearing one tooth; tip of
spermatodactyl blade-like, its canal extending inside digit.
Legs: Legs much shorter than in female; length of tarsus I 76-88,
tarsus IV 98-116; set of setae the same, but many ventral setae
very strong (Fig. B 2, 5). Femoral macrosetae as in female. Genua
I-IV with 4 macrosetae (adl, ad2, pdl, pd2). Longest leg macro
seta (adl) situated on femur IV, 92-100. Femur and tarsus II each
bear a small roundish apophysis of · setal origin (Fig. B 5).
Specialized supporting seta on tarsi II-IV strongly swollen
basally; long hair-like seta situated ventrally near it (Fig. B
5a).
DEUTONYMPH. Dorsum (Fig. B 7): Dorsal shield weakly sclerotized,
440-450 x 240-250, bearing 21 pairs of setae (S2 on shield margin,
r2 and r3 on integument). Only 3 pairs lateral setae (s4, s5, S2)
long (up to 80), all dorsocentral setae small.
Venter (Fig. B 7): All shields poorly sclerotized. Tritosternum as
in female. Sternal shield bears 4 pairs of setae and 3 pairs of
pores, st5 on integument, opisthogastric cuticle with 4 pairs of
setae. Length of most setae on ventral surface 20-23, only JV5
(8-9). With rounded peritreme fragment in front of peri treme
proper in most deutonymphs, situated at level of border between
coxae I and II.
Gnathosoma: generally as in female. Length of d.m. 40-44.
Legs: Specialized three-tined setae on tarsi II-IV, as in
female.
Etymology: The species is named in honour of the prominent Russian
acarologist Dr Adelaida D. PETROVA-NIKITINA.
DISCUSSION
Phoretic specificity of mesostigmatic mites often is associated
with their development in the nests of insects or on their larvae
in soil and other substrates (COSTA, 1969, 1971; COSTA &
HUNTER, 1971; SAM~I NAK, 1991; ATHIAS-BINCHE, 1991, etc.). The
close cohabitation with insects, considering also the similar
peculiarities of the mouth apparatus organization (strongly
developed pointed mouth appendages labrum, supralabral process,
malae ), shown by some entomophilous representatives of different
gamasid families (Macrochelidae, Laelapidae, Eviphididae,
Pachylaelapidae), may often be accompanied by direct trophic
relations with the brood of hosts. But the nutrition of these mites
has practically never been studied. There is some information
(COSTA, 1966, 1969) showing that the feeding on saprobiontic
nematodes is insufficient for the full development (Neopodocinum
caputmedusae (Berlese) and Pelethi phis gurei (Costa)), or for
normal reproduction (Pachylaelaps hispani Berlese) of mites
intimately associated with the beetle Copris lunaris L. At the same
time, laelapid mites of the genus Co/eo/ae/aps Berlese, which
possess a similar mouth apparatus, are considered to feed on skin
exudates of the larvae of scarabaeid beetle subfamily Melolonthinae
(VITZTHUM, 1940-1943). The same diet was proven for mites of the
laelapid genus Dinogamasus Kramer, in which development is related
to the brood of car penter bees (SCAIFE, 1952, cited in COSTA,
1969; MADEL, 1975, cited in SCHERF, 1975).
The collecting of mites from large scarabaeids and burying beetles
often results in the discovery of new species, including sibling
species (CosTA, 1967; MULLER & SCHWARZ, 1990; ATHIAS-BINCHE,
SCHWARZ & MEIERHOFER, 1993; BROWN & WILSON, 1992; MAKAROVA,
1996b, etc.). The species richness
of these mites is evidently due to the isolation of the underground
beetle nests and differences in the eco logical niches of the
hosts. Despite this, strict one host preferences may not come into
existence (CosTA, 1971; COSTA & ALLSOPP, 1979, 1981; KRANTZ
& MELLOT, 1972; KRANTZ, 1991; MAKAROVA, 1995, etc.). It is
important to notice that even in case of highly specific
association, the nutrition of the mites may be not related directly
to their hosts (NEUMANN, 1943; COSTA, 1964, 1967; KRANTZ &
MELLOT, 1972; KRANTZ & RoYCE, 1994).
Cryptoseius gen. n. displays a significant superficial resemblance
to Scamaphis Karg, associated with the beetles of the genus
Geotrupes (HYATT, 1990; HAlT LINGER, 1990, 1993; MAsAN, 1994; new
data). This similarity is expressed in the elongated form of the
body, reduction of dorsal shield and its setal set, the
heterogeneity of dorsal shield setae (lengthened mar ginal setae
and shortened dorsocentral ones), the broad shortened peritremes,
the strongly developed mouth processes and sensilla on the
palptarsus, and the spine-like setae on the legs. These external
simila rities could be considered as parallelisms, due to a
similar mode of life.
In different combinations, most of these morpho logical features
also appear in entomophilous gama sids of other families (HUNTER
& ROSARIO, 1988), such as representatives of the genera
Dinogamasus Kramer, Coleolaelaps Beriese, Hypoaspis Canestrini s.
str., Dynastaspis Costa (Laelaptidae ), some species of
Neopodocinum Berlese (Macrochelidae), Cosmeto lalelaps Womersley,
Paradoxiphis Berlese, etc., which coexist with large insects that
take care of their brood. These circumstances probably explain the
des cription of Coleolaelaps anoxiae (KOYUMDJIEVA, 1977) comb. n.
(Laelapidae) as a member of Pelethi phis (Eviphididae).
It is noteworthy that the morphological instability of characters
in the new species parallels the indivi dual variability displayed
by representatives of other gamasid families (Paradoxiphis,
Coleolaelaps) asso ciated with large scarabaeids (CosTA &
HUNTER, 1971; COSTA & ALLSOPP, 1979).
At present, a discussion about the relations of Cryptoseius gen. n.
with others genera of Eviphididae would be untimely (see
diagnosis). The plesiomor phic condition of the leg chaetotaxy,
tectum and
121
sterna! porotaxy suggests a rather archaic character of the new
genus.
ACKNOWLEDGEMENTS
The author is most grateful to Dr E. E. LINDQUIST, who kindly
provided comments and corrections to the manuscript, particularly
concerning setal desi gnations, to Dr A. D. PETROVA-NIKITINA and
Dr A. B. BABENKO for stimulating discussions, to Dr A. V. ToLSTIKOV
for improving the first English version of the paper, as well as to
Mrs T. A. CHERNY AKHOVS
KA Y A and Dr K. V. MAKAROV for providing material for study. The
work was supported by the "Biodiver sity" Programme, established
by the Russian Govern ment, and the Russian Foundation for Basic
Research.
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