A new eviphidid mite genus (Parasitiformes; Mesostigmata; Eviphididae) associated with the dung beetle Scarabaeus transcaspius Stolfa (Coleoptera, Scarabaeidae) in TurkmenistanAcarologia is proudly non-profit, with no page charges and free open access
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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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