J Pest Sci (2008) 81:35–42

DOI 10.1007/s10340-007-0182-9

ORIGINAL PAPER

Phoretic mites of three bark beetles (Pityokteines spp.) on Silver Wr

Milan Pernek · Boris Hrasovec · Dinka Matosevic ·
Ivan Pilas · Thomas Kirisits · John C. Moser

Received: 23 March 2007 / Revised: 6 September 2007 / Accepted: 16 September 2007 / Published online: 5 October 2007
© Springer-Verlag 2007

Abstract The species composition and abundance of relative abundance of mite associates were similar for all
phoretic mites of the bark beetles Pityokteines curvidens, three Pityokteines species. Another species, Pleuronectoc-
P. spinidens, and P. vorontzowi on Silver Wr (Abies alba) elaeno barbara was commonly found phoretic on P. curvi-
were investigated in 2003 at two locations (Trakoscan and dens, captured in pheromone traps in 2005 at the location
Litoric) in Croatia. Stem sections and branches from Litoric. Furthermore, two previously collected mite speci-
A. alba trees infested by Pityokteines ssp. were collected mens from Switzerland, phoretic on P. curvidens, were
and incubated in rearing cages. Bark beetles emerging from identiWed as Nanacarus sp. and Bonomia sp. The records
the stem sections and branches were examined for photetic from Croatia and Switzerland in the present study increase
mites. A total of ten mite species were documented for the the number of known mite associates of Pityokteines spp.
Wrst time as associates of Pityokteines spp. on A. alba. from one previously documented species to 14 species.
These included Dendrolaelaps quadrisetus, Ereynetes None of the phoretic mites found in the survey in Croatia
scutulis, Histiostoma piceae, Paraleius leontonychus, Pleu- appear to have the potential to be used for biological con-
ronectocelaeno japonica, Proctolaelaps hystricoides, trol of Pityokteines spp., although the feeding habits are
Schizostethus simulatrix, Tarsonemus minimax, Trichouro- unknown for many species recorded.
poda lamellose, and Uroobovella ipidis. T. minimax was
the most frequent phoretic mite of all the three scolytines Keywords Abies alba · Acarina · Scolytinae · Forest pests
and U. ipidis was also common, whereas, the other mite
species occurred less frequently. The species spectrum and
Introduction

M. Pernek (&) · D. Matosevic · I. Pilas

Four bark beetle species of the genus Pityokteines (Coleop-
Forest Research Institute, Jastrebarsko,
Cvjetno naselje 41, Jastrebarsko, Croatia tera, Curculionidae, Scolytinae) occur in the Palaearctic
e-mail: milanp@sumins.hr region (PfeVer 1995; Knizek 1998). Among these, P. curvi-
dens (Germar), P. spinidens (Reitter), and P. vorontzowi
B. Hrasovec
(Jakobson) commonly occur in Europe where they infest
Faculty of Forestry, University of Zagreb,
Svetosimunska 25, Zagreb, Croatia Silver Wr (Abies alba Mill.) and occasionally other conifers
(Schwerdtfeger 1981; PfeVer 1995). P. curvidens and
T. Kirisits P. spinidens usually breed in the lower trunk of Silver Wr
Institute of Forest Entomology,
trees, while P. vorontzowi usually occupies only the upper
Forest Pathology and Forest Protection (IFFF),
Department of Forest and Soil Sciences (DFS), part of the crown (Pernek 2005). The three Pityokteines
University of Natural Resources and Applied Life Sciences, species have been reported as an important factor in Silver
Vienna (BOKU), Hasenauerstrasse 38, 1190 Vienna, Austria Wr decline (widely known as “Tannensterben” in German-
speaking countries) in some parts of Europe, a syndrome
J. C. Moser
Southern Research Station, USDA Forest Service, caused by a variety of abiotic and biotic factors
2500 Shreveport Hwy., Pineville, LA, USA (Schwerdtfeger 1981). P. curvidens, P. spinidens, and

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P. vorontzowi are commonly found in Croatia and have spp. originated from natural Silver Wr stands (at two loca-
been associated with increased levels of Silver Wr mortality tions in Croatia: one in the Pannonic biogeographic region
since the beginning of 2000 (Pernek 2005). (Trakoscan, 400 m a.s.l., 46°05(⬘28⬘⬘ N and 15°56⬘55⬘⬘ E)
Bark beetles are known to be associated with diverse and the other one in the Dinaric biogeographic region
guilds of arthropods of which phoretic mites are amongst (Litoric, 550 m a.s.l., 45°27⬘00⬘⬘ N and 15°04⬘06⬘⬘ E). On
the best-known associates (Kinn 1983; Moser and Macias- 6 October 2003, one stem section (37 cm in length and
Samano 2000; Klepzig et al. 2001; Lombardero et al. 2003; 36 cm in diameter) and four branches (1 m long) were cut
Moser et al. 2005). Phoretic mites use bark beetles for from the trunk of a Silver Wr tree infested by Pityokteines
transportation to new, suitable habitats and their ecological spp. in Trakoscan. In Litoric, sampling was conducted on
roles are diverse (Lindquist 1969). It is of particular practi- 20 October 2003. Here, another stem section (46 cm in
cal interest that some phoretic mites act as predators and length and 34 cm in diameter) and four branches (1 m long)
parasitoids of bark beetles and their immature stages, espe- were taken from a bark beetle infested Silver Wr tree. Bolts
cially egg and early larval stages (Moser 1975; Moser et al. and branches were collected when broods were in the pupal
1978). They are thus potential agents for biological control stage. The sampled material was incubated in separate rear-
of scolytine forest pests (Kinn 1983), although there is ing cages at 23(§2)°C and under a 16L:8D photoperiod.
presently no successful example of any biological control All bark beetles emerging from the stem sections and
program against bark beetles using phoretic mites. branches were collected daily from the cage screens during
Mites phoretic on bark beetles can also be involved in the entire emergence period.
complex symbiotic interactions with bark beetles and fungi, A total of 192 beetles belonging to the three Pityokte-
especially ophiostomatoid fungi belonging to the genera ines species were randomly chosen and placed into vials
Ceratocystis, Ceratocystiopsis, Ophiostoma, Grosmannia with 70% ethanol (Table 1). Bark beetle specimens
and related genera of anamorphic fungi (WingWeld et al. belonging to diVerent species and originating from diVer-
1993; Kirisits 2004; Zipfel et al. 2006) and their host trees ent localities were placed in separate vials. The beetles
(Klepzig et al. 2001; Hofstetter et al. 2005). The occurrence were then transferred to lactophenol for clearing. Their
of hyperphoretic fungal spores (both conidia and asco- accompanying mites were counted and plucked from their
spores) on phoretic mites of bark beetles suggests that they can bodies. Mites were tallied separately according to whether
to some extent be involved in the transmission of fungi to they were: (1) still attached to the beetle, (2) separated
their host trees, including ophiostomatoid fungi and espe- from the beetles in lactophenol, or (3) separated from the
cially also tree pathogens (Moser et al. 1989b, 1997, 2005). beetles in alcohol.
Phoretic females of some species in the minimax group of The identiWcation of two mite specimens phoretic on
Tarsonemus even possess so-called sporothecae, specialized P. curvidens, collected by B. Forster (see acknowledgments)
integumental, pocket-like structures, in which ascospores in Switzerland and sent to J. C. Moser for determination,
and conidia of ophiostomatoid fungi are deposited and was also included in this study, as they represent hitherto
transported (Moser 1985; Bridges and Moser 1986). unpublished records. Voucher specimens (slides) of all mite
Increased local populations of P. curvidens, P. spinidens species detected in this study are stored in the collections of
and P. vorontzowi have been recently noted in conifer for- the authors (M. Pernek and J. C. Moser), and in other col-
ests in various parts of Croatia, resulting in high damage lections of mite taxonomists (H. Klompen and E. E. Lind-
levels on Silver Wr (Pernek 2005). This increase in impor- quist; see acknowledgments).
tance of Pityokteines species as forest pests of A. alba in
Croatia prompted a research on the biology of these bark
beetles species, their natural enemies (Pernek 2005) and Results and discussion
also on their mite associates. The purpose of this study was
to investigate the species spectrum and abundance of pho- A total of 504 individuals of P. spinidens emerged from 8
retic mites associated with P. curvidens, P. spinidens, and November to 10 December 2003 from the collected stem
P. vorontzowi in natural Silver Wr stands in Croatia. Previ- sections incubated in separate rearing cages at Trakoscan,
ously unpublished records of two phoretic mites of P. cur- while 512 specimens of P. spinidens and 55 indviduals of
videns in Switzerland are also reported. P. curvidens emerged from 25 November to 13 December
2003 from the stem sections collected at Litoric (Table 1).
From the branches collected at Trakoscan, 231 individuals
Materials and methods of P. vorontzowi emerged from 23 November to 2 Decem-
ber 2003, and 416 specimens of this scolytine species
Bark beetle specimens and respective bark and wood sam- emerged from 25 November to 6 December 2003 from the
ples included in this study of phoretic mites of Pityokteines branches collected at Litoric (Table 1).

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Table 1 Overview about the number of Pityokteines spinidens, Trakoscan and incubated in breeding cages in the laboratory, the
P. curvidens, and P. vorontzowi emerging from the stem sections and number of beetles investigated for the presence of phoretic mites, and
branches collected in 2003 at the two Croatian localities Litoric and number and percentage of investigated beetles carrying phoretic mites
Observations Pityokteines Pityokteines Pityokteines Total
spinidens vorontzowi curvidens

Litoric Trakoscan Litoric Trakoscan Litoric

Total number of bark beetles emerging from stem 512 504 416 231 55 1,718
sections/branches incubated in the laboratory
Number of beetles sampled for identiWcation 56 21 53 20 42 192
of phoretic mites
Number of sampled beetles carrying mites 34 11 28 11 30 114
Sampled beetles carrying mites (%) 60.7 52.4 52.8 55.0 71.4 59.4

The percentages of Wr bark beetles carrying mites on Pityokteines in Croatia (Tables 2, 3, 4, 5, 6). Overall, Tar-
their bodies ranged from about 52 to 71%, depending on sonemus minimax (Vitzthum) (Fig. 1g) and Uroobovella
the Pityokteines species and the collection site (Table 1). ipidis (Vitzthum) (Fig. 1f) were relatively common, whereas,
The overall average of insects carrying mites was about Dendrolaelaps quadrisetus (Berlese) (Fig. 1d), Paraleius
60%. These values considerably underestimate the actual leontonychus (Berlese) (Fig. 1j), and Histiostoma piceae
portion of individuals carrying phoretic mites. This is sug- (Scheucher) (Fig 1h) occurred occasionally. Trichouropoda
gested by the large number of mites that fell-oV the beetles, lamellosa (Hirschmann) (Fig. 1e), Proctolaelaps hystrico-
as indicated by their presence in the alcohol and lactophe- ides (Lindquist and Hunter) (Fig. 1c), Schizostethus simula-
nol sediments (Tables 2, 3, 4, 5, 6). trix (Athias–Henriot) (Fig. 1b), Ereynetes scutulis (Hunter)
Ten phoretic mite species are documented here for the (Fig. 1i), and Pleuronectocelaeno japonica (Kinn) (Fig. 1a)
Wrst time as associates of Wr bark beetles in the genus were rare. In addition, Pleuronectocelaeno barbara

Table 2 Species spectrum and abundance of mites phoretic on 42 individuals of Pityokteines curvidens at the locality Litoric
Mite species Phoretic stage On Location on beetle Lactophenol Alcohol Totals
beetles sediments sediments

Pleuronectocelaeno japonica Female 2 Under elytra 0 0 2

Schizostethus simulatrix Deutonymph 0 0 1 1
Dendrolaelaps quadrisetus Deutonymph 10 Under elytra 2 1 13
Trichouropoda lamellose Deutonymph 1 Elytral declivity 0 2 3
Uroobovella ipidis Deutonymph 18 Coxa, elytral declivity, 12 0 30
and ventral thorax
Tarsonemus minimax Female 28 Elytral declivity, under elytral, 60 128 216
and ventral thorax
Histiostoma piceae Deutonymph 4 Under elytra 6 2 12
Paraleius leontonychus Female 1 Coxa 1 4 6
Total 64 81 138 283

Table 3 Species spectrum and abundance of phoretic mites on 56 individuals of Pityokteines spinidens at the locality Litoric
Mite species Phoretic stage On Location on beetle Lactophenol Alcohol Totals
beetles sediments sediments

Dendrolaelaps quadrisetus Deutonymph 8 Under elytra 0 0 8

Uroobovella ipidis Deutonymph 32 Ventral thorax, elytral declivity, 0 0 32
coxa, and dorsal thorax
Tarsonemus minimax Female 29 Under elytra and ventral thorax 0 6 35
Histiostoma piceae Deutonymph 9 Under elytra 0 2 11
Paraleius leontonychus Female 4 Ventral thorax and leg 0 0 4
Total 82 0 8 90

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Table 4 Species spectrum and abundance of phoretic mites on 21 individuals of Pityokteines spinidens at the locality Trakoscan
Mite species Phoretic stage On Location on beetle Lactophenol Alcohol Totals
beetles sediments sediments

Proctolaelaps hystricoides Female 0 1 1 2

Dendrolaelaps quadrisetus Deutonymph 4 Under elytra 0 0 4
Trichouropoda lamellose Deutonymph 0 1 0 1
Uroobovella ipidis Deutonymph 5 Under elytra, elytral declivity, 15 2 22
and coxa
Tarsonemus minimax Female 28 Under elytra and ventral thorax 63 356 447
Histiostoma piceae Deutonymph 4 Under elytra 3 0 7
Ereynetes scutulis Female 0 0 1 1
Paraleius leontonychus Female 0 1 7 8
Total 41 84 367 492

Table 5 Species spectrum and abundance of phoretic mites on 53 individuals of Pityokteines vorontzowi at the locality Litoric
Mite species Phoretic stage On Location on beetle Lactophenol Alcohol Totals
beetles sediments sediments

Proctolaelaps hystricoides Female 0 0 1 1

Dendrolaelaps quadrisetus Deutonymph 4 Under elytra 0 0 4
Uroobovella ipidis Deutonymph 29 Under elytra, elytral declivity, 19 0 48
and coax
Tarsonemus minimax Female 64 Under elytra and ventral thorax 86 159 309
Histiostoma piceae Deutonymph 1 Under elytra 1 0 2
Ereynetes scutulis Female 0 1 3 4
Paraleius leontonychus Female 1 Ventral thorax 2 5 8
Total 99 109 168 376

Table 6 Species spectrum and abundance of phoretic mites on 20 individuals of Pityokteines vorontzowi at the locality Trakoscan
Mite species Phoretic stage On Location on beetle Lactophenol Alcohol Totals
beetle sediments sediments

Dendrolaelaps quadrisetus Deutonymph 5 Under elytra 1 0 6

Uroobovella ipidis Deutonymph 8 Ventral thorax, elytral declivity, 3 1 12
coax, and dorsal thorax
Tarsonemus minimax Female 1 Under elytra and ventral thorax 10 52 63
Histiostoma piceae Deutonymph 1 Under elytra 0 0 1
Paraleius leontonychus Female 0 Ventral thorax and leg 0 3 3
Total 15 14 56 85

(Athias–Henriot) was commonly found phoretic on P. cur- tonymphs (S. simulatrix, D. quadrisetus, T. lamellosa,
videns collected from pheromone traps installed in 2005 U. ipidis, H. piceae) (Tables 2, 3, 4, 5, 6).
in a Silver Wr stand in Litoric, while further specimens of The most abundant mite species on all of the three bark
P. japonica could not be detected (M. Pernek and J. C. Moser, beetle species was T. minimax which comprised 39–90% of
unpublished data). With P. barbara, which was not found all mite individuals per collection, depending on the Pityo-
on Pityokteines beetles emerging from the logs in 2003 kteines species and the site where samples were collected
(Tables 2, 3, 4, 5, 6), the number of known mite associates (Tables 2, 3, 4, 5, 6). The species spectrum and relative
of Pityokteines spp. in Croatia increased from 10 to 11 abundance of phoretic mites were similar for all the
species (Tables 2, 3, 4, 5, 6). Mites were either phoretic on three Pityokteines species, especially regarding the most
Pityokteines spp. as females (P. hystricoides, P. japonica, common species, T. minimax and U. ipidis (Tables 2, 3, 4,
T. minimax, E. scutulis, P. leontonychus) or as deu- 5, 6). The diVerences in the assemblages and the abundance

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of phoretic mites between Pityokteines species and between 1997). The three mite species from Switzerland were, how-
diVerent sampling localities thus refer mainly to those mite ever, not found in the present survey in Croatia. Summariz-
species that were generally rare (P. hystricoides, S. simula- ing the previous record from Switzerland (Nierhaus-
trix, T. lamellosa, H. piceae, E. scutulis). Care should be Wunderwald 1997) and those from Croatia and Switzerland
taken in interpreting the results of this study, as only a sin- made in the present study, the known assemblage of pho-
gle tree at each of the two study sites were sampled. How- retic mites of Pityokteines spp. on A. alba consists of 14
ever, previous experience in a number of phoretic mite bark species.
beetle systems has shown that intensive sampling of a sin-
gle tree will reveal most or all of the common mite species,
although many of the uncommon associates may be missed Phoretic mites from Pityokteines curvidens
(Moser and Roton 1971; Moser and Bogenschütz 1984;
Moser et al. 1989a, 1997). A total of 42 individuals of P. curvidens from Litoric,
The two mite specimens from P. curvidens, collected by consisting of 23 males (55%) and 19 females (45%), were
B. Forster in Switzerland, were identWed as Nanacarus sp. collected after their emergence from logs, and examined for
(Fig. 1k) and Bonomia sp. (Fig. 1l), respectively. Another mites. Eight mite species, P. japonica, S. simulatrix,
phoretic mite recorded from P. curvidens, again in Switzer- D. quadrisetus, T. lamellosa, U. ipidis, T. minimax,
land, is Pyemotes dryas (Vitzthum) (Nierhaus-Wunderwald H. piceae, and P. leontonychus were detected among the

Fig. 1 Phoretic mites from Pityokteines spp. in Croatia: a Pleuronec- f Uroobovella ipidis, g Tarsonemus minimax, h Histiostoma piceae, i
tocelaeno japonica, b Schizostethus simulatrix, c Proctolaelaps Ereynetes scutulis, j Paraleius leontonychus, and from Switzerland: k
hystricoides, d Dendrolaelaps quadrisetus, e Trichouropoda lamellosa, Nanacarus sp., l Bonomia sp.

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283 mite individuals examined (Table 2). Two mite species, detected on certain locations. Individuals of P. leontony-
P. japonica and S. simulatrix, were detected only on this chus were seen on the thorax or legs, while those of
Pityokteines species. The discovery of P. japonica in Europe P. japonica occurred only under the elytra. D. quadrisetus
is puzzling and raises questions concerning the hosts, eco- and H. piceae were found under the elytra and T. lamellosa on
logical niches, and geographic distribution of this species. the elytral declivity. U. ipidis occurred on various loca-
Kinn (1991) and Moser et al. (1997) found this mite phoretic tions, including under the elytra, on the elytral declivity,
on the spruce bark beetle Ips typographus japonicus (Niij- and the ventral thorax. Although numerous individuals of
ima) in Japan. However, in all collections of Ips typogra- the most abundant species, T. minimax, were present under
phus L. in Europe only phoretic individuals of P. barbara the elytra, the most common locations of this mite likely
where detected, while P. japonica never occurred (Moser were somewhere on the beetle’s body surface. Most indi-
and Bogenschütz 1984; Moser et al. 1989a). Hence, based viduals of this mite species fell-oV from the beetles, which
on the diVerences in phoretic hosts and geographic distribu- is more likely to occur when they are carried on the beetle’s
tion, it is possible that the populations of P. japonica on P. body surface, where attachment of the mite to the insect is
curvidens in Croatia and on I. typographus japonicus in relatively weak. Locations of E. scutulis, P. hystricoides
Japan may be morphologically identical, but genetically and and S. simulatrix could not be assessed, because of that all
ecologically separate sibling species. individuals of these species were collected from lactophe-
nol or alcohol sediments.

Phoretic mites from Pityokteines spinidens

Ecological roles of the phoretic mites
A total of 77 individuals of P. spinidens (21 from Trako- on Pityokteines spp.
scan, 56 from Litoric), consisting of 36 males (47%) and 41
females (53%), were collected after their emergence from The ecological roles of most of the phoretic mites associ-
logs and examined for mites. Five mite species, D. quadris- ated with Pityokteines spp. are poorly known. However,
etus, U. ipidis, T. minimax, H. piceae, and P. leontonychus, based on the knowledge of other bark beetle–mite–fungal
were detected among the 90 mite individuals from the loca- systems and previous records of the biology of these mites
tion Litoric (Table 3). Eight mite species were detected in the scientiWc literature, it is justiWed to assume that their
among the 492 mite individuals sampled from the location biology and ecology are diverse (Moser et al. 1995; Klep-
Trakoscan (Table 4). These included all the species zig et al. 2001; Lombardero et al. 2003). H. picea, like most
recorded from P. spinidens at Litoric and additionally also members of this genus may occur in liquid, “soupy” sub-
P. hystricoides, T. lamellose, and E. scutulis. strates, and may be a Wlter feeder of bacteria and yeasts
(Oconnor 1984). D. quadrisetus and T. lamellosa prey on
nematodes (Kinn 1967, 1987). Likewise, P. japonica and
Phoretic mites from Pityokteines vorontzowi P. barbara may also feed on nematodes (Kinn 1971). No
records exist about the feeding habits of U. ipidis. The anal
A total of 73 individuals of P. vorontzowi (20 from Trako- opening of P. hystricoides is enlarged, suggesting that it
scan, 53 from Litoric), consisting of 40 males (55%) and 33 may feed on fungal spores. S. simulatrix, like S. lyriformis
females (45%), were collected after their emergence from (Moser 1975), may prey on subcortical arthropods, as it is a
branches and examined for mites. Seven mite species, member of the predatory mite family Parasitidae. Most ori-
P. hystricoides, D. quadrisetus, U. ipidis, T. minimax, batids are detritivores (Jacot 1934; Walter and Proctor
H. piceae, E. scutulis, and P. leontonychus, were detected 1999), and P. leontonychus is probably no exception to the
among the 376 mite individuals from Litoric (Table 5). general trophic pattern of this group of mites. Similar to
Five species were detected from the 85 mite individuals other Tarsonemus species, T. minimax may feed on fungal
from Trakoscan (Table 6). These included all mite species mycelia and spores (Oconnor 1984; Lombardero et al.
recorded from P. vorontzowi at Litoric, except P. hystrico- 2003). E. scutulis is phoretic on many scolytines on coni-
ides and E. scutulis (Table 6). fers and it preferably inhabits drier and older subcortical
galleries. Although no feeding preferences of E. scutulis
have so far been observed, Walter and Proctor (1999) noted
Location of phoretic mites on Pityokteines beetles that Ereynetes macquariensis (Fain), at least, may feed on
(Tables 2, 3, 4, 5, 6) small mites, eggs, and immature stages of larger arthropods
or on nematodes. Only S. simulatrix, as it is a member
Locations of the diVerent mite species on the bark beetle of the predatory mite family Parasitidae and probably
bodies varied widely. Some species were consistently E. scutulis could have some potential for biological control

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of Pityokteines bark beetle pests, but they were very rarely Acknowledgments This research was supported by an USDA Forest
recorded in the present survey. However, further Weld and Service Federal Financial Assistance Award (SRC 06–IC-11330129-
038). Many thanks go to Kier Klepzig (USDA Forest Service, South-
laboratory studies are required to precisely assess the feed- ern Research Station, Pineville, Louisiana, USA) for his help to pro-
ing habits of the phoretic mites of Pityokteines spp. and vide Milan Pernek with this award and for his collaboration. Financial
their potential use as biocontrol agents against Wr bark support from the cooperation project “Key issues in forest protection
beetles. research in Austrian and Croatian forest stands” (ScientiWc and Tech-
nological Cooperation between Austria and Croatia, Project no. 22/
2004, 2004–2005), funded by the Austrian Exchange Service (ÖAD)
and the Croatian Ministry of Science, Education and Sports is also
Hyperphoretic fungal spores on phoretic mites gratefully acknowledged. We thank Zeljko Kauzlaric, Matija Plese
of Pityokteines spp. (Hrvatske nume Ltd., Croatia), and Beat Forster (Federal Research
Institute, Birmensdorf, Switzerland) for their assistance in beetle sam-
pling, Stacy Blomquist (USDA Forest Service, Southern Research Sta-
Hyperphoretic fungal spores were attached to the body sur- tion, Pineville, Louisiana, USA) for technical support, Rudolf
faces of many individuals of phoretic mites of Pityokteines Wegensteiner (Institute of Forest Entomology, Forest Pathology and
spp. with the exception of specimens of P. japonica, Forest Protection, BOKU University, Vienna, Austria) for permitting
us to use his laboratory facilities, Hans Klompen (Museum of Biolog-
S. simulatrix, E. scutulus, and P. leontonychus, which gen- ical Diversity, Ohio State University, Columbus, OH, USA) for deter-
erally occurred rarely. These spores appeared to be conidia mination of P. japonica, Evert Lindquist (Biodiversity Theme
of several fungi and were of various sizes and shapes. None Research Branch, Agriculture and Agri-Food Canada, OT, Canada) for
could be unambiguously assigned to individual fungal spe- his help with the determination of several mite species, and Axel
Schopf (Institute of Forest Entomology, Forest Pathology and Forest
cies, except for large, four-celled, Alternaria-like conidia. Protection, BOKU University, Vienna, Austria) for his constructive
The assemblages of ophiostomatoid fungi associated comments and suggestions on an earlier version of the manuscript.
with any of the three Pityokteines species has thus-far not
been studied (Kirisits 2004). Examination of 20 gallery sys-
tems of P. spinidens on a Silver Wr tree in Litoric in April References
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unpublished data). None of the spores found on the body J Econ Entomol 27:858–859
surfaces of mites phoretic on Pityokteines spp. were identi- Kinn DN (1967) Notes on the life cycle and habits of Digamasellus
Wed as ascospores of Ceratocystis, Ceratocystiopsis, Gros- quadrisetus (Mesostigmata: Digamasellidae). Ann Entomol Soc
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