Chrysidid wasps (Hymenoptera, Chrysididae)—Parasites of eumenine wasps (Hymenoptera, Vespidae: Eumeninae) in Crimea

Abstract

Chrysidid wasps can be considered the most abundant and sometimes the most important enemies of eumenine wasps. We investigated the nesting biology of 36 species of Eumeninae in Crimea (47% of the Crimean fauna) in 2002–2013. As a result we found 15 species of chrysidid wasps (Chrysis cylindrica, Ch. rutilans, Ch. splendidula, Ch. graelsii, Ch. valesiana, Ch. brevitarsis, Ch. ignita, Ch. impressa, Ch. longula, Ch. pseudobrevitarsis, Ch. ruddii, Ch. sexdentata, Ch. ambigua, Ch. taczanovskii, and Stilbum calens) associated with 16 species of hosts of the subfamily Eumeninae. Thus, 13 new host-parasite associations were discovered and 8 previously known ones were confirmed by direct rearing of chrysidids from the nests of eumenine wasps. The data on the host of Chrysis ambigua were obtained for the first time, and the species itself was recorded for Eastern Europe for the first time. Most of the revealed chrysidid wasps develop as inquilines in the nests of eumenines. The reports by early authors of bees as hosts of these chrysidid species are disputable. At least one of the species reared, Stilbum calens, developed as an orthoparasite in the nests of eumenine wasps.

Full text

ISSN 0013-8738, Entomological Review, 2015, Vol. 95, No. 4, pp. 472–485. © Pleiades Publishing, Inc., 2015.
Original Russian Text © K.V. Martynova, A.V. Fateryga, 2015, published in Entomologicheskoe Obozrenie, 2015, Vol. 94, No. 2, pp. 379–396.
472
Chrysidid Wasps (Hymenoptera, Chrysididae)—Parasites
of Eumenine Wasps (Hymenoptera, Vespidae: Eumeninae)
in Crimea
K. V. Martynova
a
and A. V. Fateryga
b
a
Donetsk National University, Vinnitsa, Ukraine
e-mail: martynov_av@ukr.net
b
Karadag Nature Reserve, Feodosiya, Russia
e-mail: fater_84@list.ru
Received August 25, 2014
Abstract—Chrysidid wasps can be considered the most abundant and sometimes the most important enemies of
eumenine wasps. We investigated the nesting biology of 36 species of Eumeninae in Crimea (47% of the Crimean
fauna) in 2002–2013. As a result we found 15 species of chrysidid wasps (Chrysis cylindrica, Ch. rutilans,
Ch. splendidula, Ch. graelsii, Ch. valesiana, Ch. brevitarsis, Ch. ignita, Ch. impressa, Ch. longula,
Ch. pseudobrevitarsis, Ch. ruddii, Ch. sexdentata, Ch. ambigua, Ch. taczanovskii, and Stilbum calens) associated
with 16 species of hosts of the subfamily Eumeninae. Thus, 13 new host-parasite associations were discovered and
8 previously known ones were confirmed by direct rearing of chrysidids from the nests of eumenine wasps. The
data on the host of Chrysis ambigua were obtained for the first time, and the species itself was recorded for Eastern
Europe for the first time. Most of the revealed chrysidid wasps develop as inquilines in the nests of eumenines. The
reports by early authors of bees as hosts of these chrysidid species are disputable. At least one of the species reared,
Stilbum calens, developed as an orthoparasite in the nests of eumenine wasps.
DOI: 10.1134/S0013873815040090
Cuckoo wasps (Hymenoptera, Chrysididae) have
a wide range of hosts (Kimsey and Bohart, 1991).
Their larvae may develop at the expense of eggs of
phasmids, larvae and/or pupae of sawflies, slug moths,
solitary bees and wasps, and also at the expense of
food (moth and beetle larvae, aphids, spiders, bugs,
and thrips) stored in the nests of wasps of the families
Vespidae, Sphecidae, and Crabronidae. Members of
the tribe Chrysidini (subfamily Chrysidinae) are the
only cuckoo wasps known to parasitize potter wasps of
the subfamily Eumeninae (family Vespidae), and they
may be considered the most common enemies of these
wasps (Boesi et al., 2005). Host-parasite associations
between eumenine and chrysidid wasps were estab-
lished by a number of authors (Blüthgen, 1961; Krom-
bein, 1967; Spradbery, 1973; Iwata, 1976; Cowan,
1991; Kimsey and Bohart, 1991).
By the latest estimations, there are over 3500 spe-
cies of Eumeninae in the world fauna (Hermes et al.,
2013). These wasps are ubiquitous and possess a great
variety of nest-building instincts. Most species are
solitary; a few are characterized by primitive eusocial-
ity. The females hunt lepidopteran larvae or, rarer,
beetle and sawfly larvae. Their nests are most often
built in preexisting cavities; however, wasps of many
species gnaw out their own nest tunnels in the soil or
construct nests of mineral or plant material on the
surface of the substrate (Spradbery, 1973; Iwata, 1976;
Cowan, 1991). The known fauna of Crimea includes
76 species of Eumeninae (Fateryga, 2010); the nesting
biology of 20 species was studied in detail in this re-
gion (Ivanov and Fateryga, 2003, 2004, 2006; Ivanov
et al., 2005a, 2005b; Fateryga, 2009, 2012a, 2012b,
2012c, 2013a, 2013b; Fateryga and Ivanov, 2009,
2013; Fateryga and Matushkina, 2010; Fateryga et al.,
2013). Cuckoo wasps were listed among the nest para-
sites of only six of them (Ivanov and Fateryga, 2004,
2006; Fateryga, 2009, 2012a; Fateryga and Ivanov,
2009, 2013), and these were not always identified to
species. This communication presents data on the spe-
cies composition of cuckoo wasps parasitizing the
nests of potter wasps in Crimea and summarizes the
data on their host-parasite relations.
Within the framework of this publication, it also
seems expedient to discuss some aspects of parasitism
of cuckoo wasps associated with Eumeninae. Depend-

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473
ing on their way of life, cuckoo wasps may be divided
into inquilines and orthoparasites (see Malyshev,
1966). Inquilines, such as Pseudospinolia neglecta
(Shuckard, 1837), Chrysis ignita (Linnaeus, 1761),
Ch. nitidula Fabricius, 1775 (= Ch. coerulans Fabri-
cius, 1804), and Ch. cembricola Krombein, 1958, use
the food stored in the host cells (Chapman, 1869;
Maréchal, 1923; Krombein, 1967; Sears et al., 2001;
Veenendaal, 2013). The females of these cuckoo
wasps lay an egg in the host cell at the last stages of
provisioning. The few existing data point to the fact
that the young inquiline larvae kill the egg or the
young host larva and only after this set to feeding on
the stored provisions (Adlerz, 1910; Krombein, 1967;
Veenendaal, 2013). Orthoparasites, such as Chrysis
viridula Linnaeus, 1761 (= Ch. bidentata Linnaeus,
1767), Ch. grandis Brullé, 1846 (= Ch. carinata
Guérin, 1842), Neochrysis viridis (Guérin, 1842), and
Stilbum cyanurum (Forster, 1771), lay eggs directly at
the host stage on which their larvae would develop: on
the last-instar larva already cocooned or completing its
cocoon, the prepupa (Chapman, 1869; Adlerz, 1910;
Malyshev, 1966) or the pupa (Matthews and Mat-
thews, 2009). To lay eggs, the females of these cuckoo
wasps have to penetrate through the cell walls and also
through the cocoon of the host.
The ways of penetration of cuckoo wasps into the
potter wasp nests as well as the ways of oviposition
and larval feeding are described only in several works.
The publication of Chapman (1860) which contains
data on the development of two cuckoo wasp species,
Chrysis viridula and Pseudospinolia neglecta on one
host, Odynerus spinipes (Linnaeus, 1758) is worth of
special mention, since these data allow one to consider
the traits distinguishing inquilines from orthoparasites.
In inquilines, emergence of adults coincides with that
of the host, whereas the adults of orthoparasites start
emerging after the last host adult has left the nest.
Thus, the flight periods of orthoparasites and their
hosts do not coincide (Kilimnik, 1993). The activity
peak of inquilines falls on the period of nest construc-
tion and provisioning by the host. They may infest any
cell in the nests but rarely infest the nests built during
the late phase of their host’s flight. Such orthopara-
sites as Ch. viridula become active at the stage of nest
provisioning but lay eggs when the host larvae have
reached their maximum size and begin to spin co-
coons. It is interesting that females of orthoparasites
can penetrate only into unfinished cells or into cells
lacking the additional protective layer that covers the
whole nest (Chapman, 1869), whereas their progeny
may perish if this layer is subsequently added
(Fateryga and Ivanov, 2009). Kilimnik (1993) consid-
ers the location of the infested cells only in the outer
section of the nest to be the indication of orthopara-
sitism. Females of Ch. viridula lay from 6 to 19 eggs
in each host cocoon (Chapman, 1869), which may be
related to the fact that its eggs are likely to be dam-
aged by the actively moving host larva during cocoon
spinning. The great number of the eggs laid may be
also considered an indication of orthoparasitism. Thus,
inquilines and orthoparasites developing on the same
host may be spatially isolated and also separated in
terms of their flight time.
There are no metaparasites among the cuckoo wasps
associated with Eumeninae. The specific trait of
metaparasites is delayed development of the larvae:
although these wasps lay an egg into the host cell al-
ready at the provisioning stage (almost simultaneously
with the oviposition of the host), the larva of the
metaparasite begins to eat the host larva only after it
has consumed all the provender, reached its maximum
size, and completed its cocoon, i.e., at the prepupal
stage. This is characteristic of cuckoo wasps develop-
ing in bee nests. This behavior of the cuckoo wasp
larvae is adaptive because, being carnivorous, they
cannot consume the provender of plant origin (pollen
and nectar) themselves and have to wait until it has
been eaten by the host. Since metaparasites eat the
already cocooned host larva, their own cocoons occur
inside those of their hosts (Ferton, 1905; Krombein,
1958, 1967; Grandi, 1961; Martynova, 2014). The
same position of cocoons is characteristic of ortho-
parasites. But, in spite of this shared feature, meta- and
orthoparasites can be differentiated by the position of
infested cells in the linear host nests. Orthoparasites
may infest only one or (rarely) several endmost cells
since they usually lay eggs after completion of the host
nesting period, whereas metaparasites may be found in
any host cell (or in several cells located in any part of
the nest).
The structure of potter wasp nests excludes the pos-
sibility of development of reptoparasitic cuckoo wasps
(Martynova and Fateryga, 2014), i.e., those capable of
penetrating through thin partitions in linear nests and
consuming several host larvae of different instars.
A number of problems associated with establish-
ment of parasite-host relations between representatives
of these groups require special attention. Analysis of

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474
the published data is difficult since some of them are
clearly erroneous. For example, Lepeletier (1806, cited
after Maréchal, 1925) wrote that cuckoo wasps did not
usually construct cocoons. The description by
Maneval (1932) of Pseudospinolia neglecta develop-
ing as an endoparasite inside the larva of Hoplitis vil-
losa (Schenck, 1853) was most probably based on
misidentification of the insect or misinterpretation of
the observations (Kunz, 1994).
In some cases, the cuckoo wasps could have been
misidentified, for instance, those of the ignita group of
the genus Chrysis Linnaeus, 1761, where some species
were quite recently described (e.g., Chrysis leptoman-
dibularis Niehuis, 2000). Between the late XIX and
the middle of the XX century, when most data on the
host-parasite associations of potter and cuckoo wasps
were published, many authors could not distinguish
the closely related and sibling species because their
diagnoses were insufficiently detailed. Examination of
the type material has recently provided additional di-
agnostic characters for such taxa (Rosa, 2005, 2009;
Smissen, 2010).
It should be also taken into account that in some
cases mistakes may arise in establishing the host-
parasite relationships due to the fact that hosts of
many cuckoo wasps make their nests in abandoned
nests of other wasps and bees or else, in the tunnels of
xylophagous insects. For instance, Trautmann (1927,
cited after Berland and Bernard, 1938) reported the
finding of Chrysis sexdentata Christ, 1791 in the nest
of the mason bee Megachile (Chalicodoma) sicula
(Rossi, 1792) on Sicily. Other authors (Berland and
Bernard, 1938) mentioned potter wasps of the genus
Ancistrocerus as the hosts of this cuckoo wasp. Since
these wasps sometimes nest in abandoned cells of bees
of the subgenus Chalicodoma Lepeletier, 1841 (see
Blüthgen, 1961), association between Ch. sexdentata
and M. sicula is doubtful (Kunz, 1994). Another ex-
ample of such a mistake may be indication of the bu-
prestid beetle Ptosima undecimmaculata (Herbst,
1784) (= Buprestis novemmaculata Fabricius, 1775) as
the host of Chrysis ignita (Berland and Bernard,
1938). In this case, the actual host was probably some
hymenopteran nesting in the empty tunnels of this
xylophagous beetle (Rosa, 2006). The situation may
be aggravated if such data are cited by subsequent
authors without critical consideration.
Observations pointing to the host associations of
cuckoo wasps are not equally significant. Already
Berland and Bernard (1938) suggested classification
of such data depending on their reliability, in particu-
lar, on whether only penetration of the cuckoo wasps
into the nest and oviposition were recorded, or all the
development stages were observed, etc. Data or sug-
gestions based on observations of cuckoo wasps which
showed activity near the host nesting places or pene-
trated into the host nests deserve special consideration.
Such data should be interpreted with great caution, all
the more so that, as pointed out by Buysson (1891–
1896), adults of many cuckoo wasp species feed as
“marauders” on the mixture of pollen and nectar
stored in provisioned cells of bees. For example, Ar-
nold (1908) observed Chrysis fulgida (Linnaeus, 1761)
flying around the nests of the mason bee Osmia nive-
ata (Fabricius, 1804) [= O. fulviventris (Panzer,
1798)]. Numerous subsequent studies showed that
Ch. fulgida could develop in nests of many wasps
(potter wasps of the genera Odynerus Latreille, 1802,
Ancistrocerus Wesmael, 1836, and Symmorphus
Wesmael, 1836, and also sand wasps of the genera
Trypoxylon Latreille, 1796, Crabro Fabricius, 1775,
and Argogorytes Ashmead, 1899), but no bees were
recorded among its hosts (Mocsáry, 1912; Blüthgen,
1961; Doronin, 1996). As far as it is known, only Ber-
land and Bernard (1938) and Rosa (2006) considered
the circumstances suggesting this association when
citing Arnold (1908). Many other authors ignored this
information and simply included the mason bee into
the list of hosts of Ch. fulgida (Nikolskaya, 1978; Ba-
naszak, 1980; Zvantsov, 1988), thus distorting the
knowledge of its host associations.
Thus, when analyzing the relations between Chry-
sididae and Eumeninae, one should proceed from reli-
able data and consider the original sources to specify
the circumstances in which observations were made.
This paper is dedicated to the memory of Professor
E.S. Sugonyaev who devoted a considerable part of his
long scientific work to the study of parasitism by the
example of chalcid wasps.
MATERIALS AND METHODS
Research was carried out in Crimea in 2002–2013
(1 cuckoo wasp was reared in 2001), mainly based on
the material of the completely finished potter wasp
nests. The nests were found under natural conditions
or obtained from artificial constructions (trap-nests).
The traps were installed in 80 sites embracing all the
landscape zones of Crimea (Priority-Setting in Con-
servation, 1999): 12 sites in the light forests of the

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475
The material of eumenine wasp nests studied in Crimea in 2002–2013
Number of nests
Wasp species
in natural substrate
in artificial nest
constructions
*Discoelius dufourii Lepeletier, 1841 – 23
D. zonalis (Panzer, 1801) – 9
*Paravespa rex (von Schulthess, 1924) 12 –
*Tropidodynerus interruptus (Brullé, 1832) 12 –
*Gymnomerus laevipes (Shuckard, 1837) 5 26
Odynerus albopictus calcaratus (Morawitz, 1885) 5 –
O. femoratus de Saussure, 1856 10 –
Alastor mocsaryi (André, 1884) 1 10
Leptochilus alpestris (de Saussure, 1855) 1 –
L. regulus (de Saussure, 1855) – 2
*Syneuodynerus egregius (Herrich-Schaeffer, 1839) 1 107
*Euodynerus dantici (Rossi, 1790) 1 28
*Eu. disconotatus (Lichtenstein, 1884) 16 43
*Eu. posticus (Herrich-Schaeffer, 1841) – 245
Eu. quadrifasciatus (Fabricius, 1793) – 11
Eu. velutinus Blüthgen, 1951 – 6
Stenodynerus bluethgeni van der Vecht, 1971 – 10
S. chevrieranus (de Saussure, 1855) 1 3
S. steckianus (von Schulthess, 1897) – 2
Allodynerus rossii (Lepeletier, 1841) – 1
Symmorphus crassicornis (Panzer, 1798) – 2
*Ancistrocerus antilope (Panzer, 1798) – 181
*A. auctus (Fabricius, 1793) 29 17
*A. gazella (Panzer, 1798) 3 53
*A. nigricornis (Curtis, 1826) 2 161
A. oviventris (Wesmael, 1836) 2 –
*A. parietinus (Linnaeus, 1761) – 40
A. trifasciatus (Müller, 1776) – 15
*Eumenes coarctatus lunulatus (Fabricius, 1804) 18 –
*Eu. coronatus (Panzer, 1799) 16 –
Eu. dubius de Saussure, 1852 4 –
Eu. papillarius (Christ, 1791) – 6
Eu. punctaticlypeus kostylevi (Kurzenko, 1976) 1 –
Eu. sareptanus André, 1884 2 –
Katamenes dimidiatus (Brullé, 1832) 4 –
*K. flavigularis (Blüthgen, 1951) 29 –
Total 175 1001
Asterisk (*) marks the species in whose nests cuckoo wasps were found.

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476
south coast, 3 in the forests of the south macroslope,
4 in the mountain meadows and yaila steppes, 5 in the
forests of the north macroslope, 16 in the piedmont
forest-steppes, 19 in the true steppes, and 21 sites
in the semi-desert steppes and saline lands. The trap-
nests were sheaves of hollow stem segments of
the common reed Phragmites australis (Cav.) Trin.
ex Stend., wrapped in paper and polythene film. The
most commonly used standard trap was a sheaf of
19–25 reed stem segments 15–25 cm long and 3 to
12 mm in diameter. Besides trap-nests, we used spe-
cial Fabre’s hives (Ivanov et al., 2009) and also single
reed stems which were dug into the vertical surface of
a steep with their openings level with the soil surface.
Under natural conditions, the nests were found at-
tached to plant stems and rocks, in the soil, in hollow
plant stems, and in the nest tunnels and abandoned
cells of other wasp and bee species, mainly Sceliphron
destillatorium (Illiger, 1807) (Hymenoptera, Spheci-
dae).
The structure and composition of nests were studied
in the laboratory, in winter. The reed stems containing
nests were split with a knife, and the upper part was
removed. The nests located in the soil and in cells of
S. destillatorium were opened by cutting off consecu-
tive earth layers. After this, a schematic drawing of the
general outlook of the opened nest and arrangement of
cells in the nest cavity was made on a sheet of paper.
Larvae and prepupae of cuckoo wasps in their cocoons
and also prepupae of the host removed from the cells
were placed in short glass tubes with dense cottonwool
stoppers on both ends, labeled with the number of the
nest and cell. After eclosion, the adults were identified
to species and this information was added to the nest
scheme. Some data were obtained from direct observa-
tions of adult cuckoo wasps close to the host nests
(Fateryga, 2009).
The table contains the list of all the potter wasp
species whose nests were studied by us in Crimea,
with the exception of Eustenancistrocerus amadanen-
sis (de Saussure, 1855), whose nests were abandoned
and did not contain a single cell (Fateryga et al.,
2013). The number of nests found under natural condi-
tions and obtained from nest traps in 2002–2013 (ex-
cluding the abandoned nests with no cells in them) is
given for each of the 36 species. The total number of
the nests examined was 1176.
The cuckoo wasps were identified using a number
of published keys (Mocsáry, 1889; Linsenmaier, 1959,
1968, 1987, 1997; Rosa, 2006; Smissen, 2010). The
identity of some specimens was confirmed by P. Rosa
(Bernareggio, Italy). The material studied is kept in the
collections of Vernadsky Taurida National University,
Simferopol (VTNU) and Donetsk National University,
Vinnitsa (DonNU).
RESULTS
As the result of research, 127 adults of 15 species of
Chrysididae were obtained from nests of 16 species of
Eumeninae. The annotated list of cuckoo wasp species
and the material studied are given below. The list does
not include the records of empty cocoons and those
from which adults did not emerge for different rea-
sons. These cocoons were found only in the nests of
those host species from which at least one adult
cuckoo wasp was reared; they most probably belonged
to the same species.
Genus CHRYSIS Linnaeus, 1761
Subgenus Chrysis s. str.
Ch. viridula group
Chrysis cylindrica Eversmann, 1857.
Material. Tarkhankut Peninsula, Kipchak ravine,
near nests of Tropidodynerus interruptus (in the proc-
ess of excavating the “blockage” of earth lumps cover-
ing the nest entrance, in the absence of host wasp fe-
males), 22.VI.2007 (A. Fateryga), 3 ♀ (DonNU).
Notes. This species was identified only to genus
and listed as “Chrysis sp. 1” by Fateryga (2009).
Distribution. The species was recorded in the coun-
tries of Central, Southern, and Eastern Europe (includ-
ing Crimea) and Asia Minor (Linsenmaier, 1968;
Kimsey and Bohart, 1991; Rosa and Soon, 2004).
Host range. The only known host of this species is
Tropidodynerus interruptus (Brullé, 1832) (Linsen-
maier, 1968).
Specific biological traits. The nests of
T. interruptus are vertical burrows in the soil contain-
ing a single cell each. In the process of gnawing out
the nest, the female makes lumps of earth and regurgi-
tated liquid, which it removes from the nest canal and
piles up by the entrance. Every time before flying out
to hunt, the female covers the nest entrance with these
lumps (Arens, 1999; Fateryga, 2009). Females of
Ch. cylindrica can dismantle this “blockage” in the
absence of the host females (Fateryga, 2009). Such

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477
behavior should evidently be regarded as confirmation
of the host-parasite association.
Ch. splendidula group
Chrysis rutilans Olivier, 1790
Material. Simferopol, in nest of Gymnomerus
laevipes, 4.VIII.2002 (A. Fateryga), 1 ♀ (dead)
(VTNU); Simferopol, Fountains, in nest of Katamenes
flavigularis, 2007 (V. Zhidkov), 1 ♀ (dead) (DonNU);
Simferopol Distr., Krasnolesye, from nest of Ancistro-
cerus sp., 23.V.2009 (A. Fateryga), 1 ♀ (VTNU); env.
of Perevalnoe, Ayan area, in nest of G. laevipes, 2008
(A. Fateryga), 1 ♀ (dead) (DonNU); same locality,
from nest of G. laevipes, 2.VI.2009 (A. Fateryga), 1 ♀
(VTNU).
Notes. In the work of Fateryga and Ivanov (2009)
this species was identified only to genus and listed as
“Chrysis (Tetrachrysis) sp”.
Distribution. The species is widely distributed in
the Palaearctic, from Europe and North Africa to
Southwest Asia (Semenov-Tian-Shansky and Nikol-
skaya, 1954; Linsenmaier, 1959, 1968; Móczár, 1967;
Rosa, 2006).
Host range. The diversity of hosts of this species is
difficult to assess since many authors who cited data
on its biology did not distinguish Ch. rutilans from
Ch. splendidula Rossi, 1790, Ch. insperata Chevrier,
1870, and Ch. continentalis Linsenmaier, 1959 (Rosa,
2006). The previously recorded nominal hosts are the
potter wasps Gymnomerus laevipes (Shuckard, 1837),
including G. l. scandinavus (de Saussure, 1856),
Microdynerus exilis (Herrich-Schaeffer, 1839),
M. nugdunensis (de Saussure, 1855), M. timidus
(de Saussure, 1856), Allodynerus rossii (Lepeletier,
1841), Anthepipona orbitalis (Herrich-Schaeffer,
1839), Ancistrocerus parietum (Linnaeus, 1758),
Eumenes pomiformis Fabricius, 1781; the sand wasps
Trypoxylon figulus (Linnaeus, 1758), T. scutatum
Chevrier, 1867, and the mason bees Osmia andre-
noides Spinola, 1808 and Heriades truncorum (Lin-
naeus, 1758) (Buysson, 1891–1896; Mocsáry, 1912;
Blüthgen, 1961; Kilimnik, 1993; Rosa, 2006).
Specific biological traits. According to our data,
Ch. rutilans develops at the expense of a number of
potter wasp species varying strongly in their nesting
bi
ology, in particular, Katamenes flavigularis (Blüth-
gen, 1951) building open nests on stone surfaces, An-
cistrocerus sp. nesting in preexisting cavities, and
Jucancistrocerus caspicus Giordani Soika, 1970 nest-
ing in the earth on clay slopes (Fateryga and Amolin,
2013); the latter is absent in Crimea. Chrysis rutilans
occurs most often in the nests of G. laevipes, infesting
1.5% of its progeny (Fateryga, 2012a).
Chrysis splendidula Rossi, 1790
Material. Lake Sasyk, from nests of Eumenes
coarctatus lunulatus, VIII.2007 and 1–10.VI.2008
(D. Puzanov), 1 ♀, 1 ♂ (DonNU); Cape Martyan Re-
serve, from nests of Eu. c. lunulatus, 6–14.V.2012
(A. Fateryga), 1 ♀, 1 ♂ (DonNU).
Distribution. The species is known from Europe,
North Africa, Asia Minor, and Central Asia (Radosz-
kowski, 1891; Buysson, 1891–1896; Linsenmaier,
1959; Móczár, 1967; Kimsey and Bohart, 1991; Rosa,
2006).
Host range. The list of known hosts of this species
should nominally include Gymnomerus laevipes,
Paragymnomerus spiricornis (Spinola, 1808),
Euodynerus variegatus (Fabricius, 1793), Eumenes
mediterraneus Kriechbaumer, 1879, and Eu. pomi-
formis (Buysson, 1891–1896; Enslin, 1933; Berland
and Bernard, 1938; Kilimnik, 1993; Tussac and
Blasco-Zumeta, 1999–2004). Considering the original
sources, we find that Trypoxylon figulus, T. scutatum,
and Osmia andrenoides listed earlier as hosts of this
species were in fact named by Buysson (1891–1896)
as hosts of Ch. rutilans (which he referred to as
Ch. splendidula Dahlbom nec Rossi). Then these spe-
cies were mistakenly included into the list of hosts of
Ch. splendidula Rossi by a number of authors (Moc-
sáry, 1912; Móczár, 1967; Nikolskaya, 1978; Zvan-
tsov, 1988; Kilimnik, 1993; Rosa, 2006). On the other
hand, Buysson (1891–1896) noted that Ch. splendi-
dula Rossi (as Ch. cyanopyga Dahlbom) had been
reared from the nests of “Eumenes pomiformis L. var.
mediterraneus Kriechb.,” i.e., Eu. mediterraneus ac-
cording to the modern classification, which earlier had
not been listed among the hosts of this species. The
association with Eu. pomiformis, mentioned in most
works, was observed only by Ferton (1910). A number
of other reports await confirmation. For example,
Lamprecht (1881) mentioned Ch. splendidula as
a possible parasite of Symmorphus allobrogus (de
Saussure, 1855) (= S. bifasciatus auct. non Linnaeus,
1761); Blüthgen (1961) named the potter wasp Lepto-
chilus alpestris (de
Saussure, 1855) as a possible host;
Fabre (cited after Berland and Bernard, 1938) said the

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478
same of a wool bee of the tribe Anthidiini. Thus,
Eumenes coarctatus lunulatus (Fabricius, 1804) is
indicated herein as a host of Ch. splendidula for the
first time.
Ch. graelsii group
Chrysis graelsii Guérin-Meneville, 1842
Material. Lisya Bay, from nests of Euodynerus dis-
conotatus in abandoned cells of Sceliphron destillato-
rium, 12.IV.2014 (A. Fateryga), 2♀ (DonNU).
Distribution. The species was recorded from
Europe, North Africa, Western and Central Asia (Ber-
land and Bernard, 1938; Balthasar, 1954; Linsenmaier,
1959; Rosa and Soon, 2004).
Host range. There are published data on develop-
ment of this species at the expense of Odynerus reni-
formis (Gmelin, 1790) and Euodynerus notatus
(Jurine, 1807), and also the bees Osmia bicornis (Lin-
naeus, 1758) [= O. rufa (Linnaeus, 1758)] and Hoplitis
anthocopoides (Schenck, 1853) (= O. caementaria
Gerstaecker, 1869) (Mocsáry, 1912; Berland and Ber-
nard, 1938; Grandi, 1961; Rosa and Soon, 2004; Pärn
et al., 2015). Euodynerus disconotatus (Lichtenstein,
1884) is reported here for the first time as a host of
Ch. graelsii.
Specific biological traits. This species seems to be
specialized to infesting potter wasp nests located in the
soil or close to the soil surface. Its previously known
host, O. reniformis, builds nests in the earth on clay
steeps and erects a curved earthen tube by the nest
entrance (Fabre, 1993). Euodynerus disconotatus, like
Eu. notatus, usually nests above the soil level in vari-
ous available cavities, often in old nests of other wasps
and bees (Fateryga, 2012c). In our material, nests in-
fested with cuckoo wasps were located in abandoned
cells of a nest of Sceliphron destillatorium positioned
under a stone at the soil surface level. Thus, the nests
of Eu. disconotatus did not essentially differ from
those of O. reniformis, except for the absence of the
earthen tube by the entrance.
Chrysis valesiana Frey-Gessner, 1887
Material. Tarkhankut Peninsula, Kipchak ravine,
near nests of Tropidodynerus interruptus (in the proc-
ess of excavating the “blockage” of earth lumps cover-
ing the nest entrance, in the absence of the host wasp
females), 22.VI.2007 (A. Fateryga), 1 ♀ (DonNU);
Lisya Bay, from nest of Paravespa rex, 10–12.VI.2011
(A. Fateryga), 1 ♀ (DonNU).
Notes. This species was identified only to genus
and listed as “Chrysis sp. 2” by Fateryga (2009).
Distribution. The species was recorded in the coun-
tries of Southern Europe (France, Portugal, Spain, and
Switzerland) and North Africa (Algeria, Morocco)
(Berland and Bernard, 1938; Linsenmaier, 1959; Kim-
sey, Bohart, 1991; Rosa and Soon, 2004).
Host range. The author of the original description
named the potter wasp Paragymnomerus
spiricornis as
t
he host of this species (Frey-Gessner, 1887, 1892). In
our material, the association with Paravespa rex (von
Schulthess, 1924) was reliably established, and the
fact of “excavation” of the nests of Tropidodynerus
interruptus was recorded.
Specific biological traits. Similar to Ch. cylindrica,
the females of this species can remove the “blockage”
covering the entrance to the nest of T. interruptus in
the absence of the host. Excavation of one of the nests
after the cell had been sealed by the wasp revealed
a cuckoo wasp egg on the nest wall 2 mm below the
plug. It was also observed that the cuckoo wasp fe-
males examined with their antennae and ovipositor the
weevil larvae placed by the T. interruptus female close
to the burrow entrance (Fateryga, 2009). The nests of
the second host species, P. rex, have no cover of earth
lumps and instead of it, an earthen tube is erected by
the nest entrance; the females of Ch. valesiana can
easily find this tube to get inside the nest (Fateryga
and Ivanov, 2013). It should be noted that the nests of
P. spiricornis, earlier mentioned as host of
Ch. valesiana, are generally similar to those of P. rex
in structure (Móczár, 1939).
Ch. ignita group
Chrysis brevitarsis Thomson, 1870
Material. Simferopol Distr., env. of Krasnolesye,
Kharab-Tavel, from nest of Ancistrocerus antilope,
18–21.V.2011 (V. Zhidkov), 1 ♀ (VTNU); same local-
ity, from nest of Discoelius dufourii, 18–21.V.2011
(V. Zhidkov), 1 ♀ (DonNU).
Distribution. The species was recorded in a number
of European countries: Sweden, Norway, Denmark,
the Netherlands, Finland, Switzerland, Spain, Austria,
Czechia, Estonia, Germany, Poland, and Ukraine (Lin-
senmaier, 1959; Móczár, 1967; Rosa and Soon, 2004;
Brustilo and Martynov, 2009).
Host range. Potter wasps of the genus Discoelius
Latreille, 1809: D. zonalis (Panzer, 1801) and

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479
D. dufourii Lepeletier, 1841 (= D. priesneri Mader,
1936), were mentioned earlier as hosts of this species
(Blüthgen, 1961; Gauss, 1966). We observed its de-
velopment not only on D. dufourii but also on Ancis-
trocerus antilope (Panzer, 1798).
Chrysis ignita (Linnaeus, 1758)
Material. Krasnoperekopsk Distr., Pochetnoe, from
nests of Ancistrocerus gazella, 4–30.IV.2010
(V. Zhidkov), 1 ♀, 4 ♂ (VTNU); Tarkhankut Penin-
sula, Kipchak ravine, from nest of A. gazella,
23.IV.2005 (A. Fateryga), 2 ♂ (VTNU); Simferopol,
from nests of Ancistrocerus nigricornis, 12–
19.VI.2003 and 1–10.V.2008 (A. Fateryga and
S. Ivanov), 3 ♀, 1 ♂ (VTNU); Simferopol Distr.,
Gvardeiskoe, from nest of A. gazella, 15.VIII.2003
(A. Milovanov), 1 ♂ (VTNU); same locality, from
nest of Euodynerus disconotatus, 4.VI.2004 (A. Mi-
lovanov), 2 ♂ (DonNU); Simferopol Distr., Zhivopis-
noe, in nest of A. nigricornis, 2005 (S. Ivanov), 13 ♀,
2 ♂ (dead) (VTNU); env. of Perevalnoe, Ayan, from
nests of A. nigricornis, 27.VI.2006 and 9–10.VI.2007
(A. Fateryga), 3 ♂ (VTNU); Yalta, from nest of Ancis-
trocerus auctus in abandoned nest cell of Sceliphron
destillatorium, VI.2001 (A. Fateryga), 1 ♀ (VTNU).
Notes. In the work of Ivanov and Fateryga (2004)
the species was mistakenly listed as Ch. cerastes
Abeille de Perrin, 1877.
Distribution. The species is trans-Palaearctic (Ber-
land and Bernard, 1938; Balthasar, 1954; Linsenmaier,
1959; Móczár, 1967; Kimsey and Bohart, 1991).
Host range. Chrysis ignita belongs to a taxonomi-
cally complicated group of species which are still hard
to differentiate despite the recent revisions (Smissen,
2010). Therefore, the reported host-parasite associa-
tions, especially those discovered in the XVIII–XX
centuries, should be called in question. Potter wasps
are most often reported as the hosts of Ch. ignita, but
there are also indications to representatives of the su-
perfamily Apoidea. The list of nominal hosts includes
the following species of Eumeninae: Odynerus
spinipes, Gymnomerus laevipes, G. l. scandinavus,
Euodynerus posticus (Herrich-Schaeffer, 1841),
Eu. variegatus, Allodynerus delphinalis (G
iraud,
1866), Symmorphus allobrogus, S. angustatus (Zet-
terstedt, 1838), S. bifasciatus (Linnaeus, 1761),
S. debilitatus (de Saussure, 1855), S. gracilis (Brullé,
1832), S. murarius (Linnaeus, 1758), Ancistrocerus
antilope, A. parietum, A. parietinus (Linnaeus, 1761),
A. nigricornis (Curtis, 1826), A. gazella (Panzer,
1798), A. oviventris (Wesmael, 1836), A. scoticus
(Curtis, 1826), A. trifasciatus (Müller, 1776), Eumenes
pedunculatus (Panzer, 1799), and Delta unguiculatum
(Villers, 1789). We have confirmed development on
two of the above species (see Material), whereas the
fact of development at the expense of Ancistrocerus
auctus (Fabricius, 1793) and Euodynerus disconotatus
has been established for the first time.
Specific biological traits. In Crimea, this species
seems to be specialized to infesting the nests of small
wasps of the genus Ancistrocerus positioned in exist-
ing cavities. It is most closely associated with
A. nigricornis and in some years destroys up to 4.0%
of its progeny (Fateryga, unpublished data).
Chrysis impressa Schenck, 1956
Material. Karadag Nature Reserve, from nest of
Ancistrocerus parietinus, 22.IV.2013 and 8.IV.2014
(V. Zhidkov), 1 ♀, 2 ♂ (DonNU).
Distribution. The species is known only from
Europe (Rosa and Soon, 2004; Agnoli and Rosa,
2012).
Host range. Morgan (1984) reared this species
from the nests of Ancistrocerus parietinus and A. tri-
fasciatus in Great Britain. Our research confirmed the
association with A. parietinus.
Chrysis longula Abeille de Perrin, 1879
Material. Bakhchisarai Distr., Vilino, from nests of
Euodynerus posticus, 11.V–19.VI.2009 (S. Ivanov),
18 ♀, 12 ♂ (25 in VTNU, 5 in DonNU); Bakhchisarai
Distr., Nauchnyi, from nests of Ancistrocerus antilope,
21–22.IV.2013 (V. Zhidkov), 2 ♀ (VTNU); Simfer-
opol Distr., Lozovoe, from nest of A. antilope,
9.V
.2011 (S. Ivanov), 1 ♀ (VTNU); env. of Krasnole-
sye, Kharab-Tavel, from nests of A. antilope, 9–
16.V.2011 (V. Zhidkov), 2 ♀, 6 ♂ (VTNU); Crimea
Nature Reserve, Buzinova barracks, from nests of
A. antilope, 23–29.IV.2010, 26.IV–11.V.2011, 12–
23.IV.2013 (V. Zhidkov), 6 ♀, 6 ♂ (8 in VTNU, 4 in
DonNU); Karadag Nature Reserve, from nests of
A. antilope, 19–24.V.2006 (S. Ivanov), 1 ♀, 2 ♂
(VTNU).
Distribution. This is a trans-Palaearctic species oc-
curring from Europe and North Africa to Japan (Lin-
senmaier, 1959, 1968; Kimsey and Bohart, 1991;
Rosa, 2006).

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480
Host range. There are published data on the devel-
opment of this species at the expense of the potter
wasps Ancistrocerus antilope, A. parietinus, and Sym-
morphus murarius. Indication by Doronin (1996) of its
development on Diodontus tristis Vander Linden,
1829 (Hymenoptera, Crabronidae) and Osmia bicornis
(= O. rufa) seems doubtful. One more species,
Euodynerus posticus, is added herein to the host list.
Specific biological traits. In spite of the most ex-
tensive material obtained, we revealed only two hosts
of this species, A. antilope and Eu. posticus. Although
these hosts belong to different genera, they possess
a similar nest structure including close size parameters
of the cells (Fateryga, 2012b), which are larger than
those of the hosts of Ch. ignita. It can be stated that
Ch. longula is a rather specialized parasite of
Eu. posticus and A. antilope which in some years de-
stroys up to 9.6% of the host progeny in Crimea
(Fateryga, unpublished data).
Chrysis pseudobrevitarsis Linsenmaier, 1951
Material. Bakhchisarai Distr., Nauchnyi, from nests
of Ancistrocerus antilope, 23–24.IV.2013
(V. Zhidkov), 4 ♀ (DonNU); Karadag Nature Reserve,
from nest of A. antilope, 24.V.2006 (S. Ivanov), 1 ♀
(DonNU).
Distribution. This is a European species (Linsen-
maier, 1959; Kimsey and Bohart, 1991; Rosa and
Soon, 2004; Rosa, 2006).
Host range. Morgan (1984) indicated Ancistrocerus
antilope as a host of this species in the territory of
Great Britain. In Estonia it was reared from the nest of
Euodynerus notatus (Pärn et al., 2015). Suggestions on
the possibility of development at the expense of
Euodynerus quadrifasciatus (Fabricius, 1793) were
also published (Kunz, 1994; Saure, 1998). We have
confirmed its association with A. antilope.
Chrysis ruddii Shuckard, 1836
Material. Demerdzhi Mt, from nest of Eumenes
coronatus, 22.V.2008 (A. Fateryga), 1 ♀ (DonNU).
Distribution. The species was recorded in Europe,
North Africa, Middle East, and Southern Asia (Ber-
land and Bernard, 1938; Linsenmaier, 1959, 1968;
Móczár, 1967; Rosa and Soon, 2004).
Host range. According to the published data,
a number of host species are known including the pot-
ter wasps and mason bees: Ancistrocerus oviventris,
A. parietum, A. scoticus, Odynerus spinipes,
O. reniformis, Delta unguiculatum, Eumenes coarc-
tatus (Linnaeus, 1758), Hoplitis adunca
(Panzer,
1798) (= O
smia spinolae Lepeletier, 1841),
H. anthocopoides (= O. caementaria), and H. ravouxi
Pérez, 1902 (Mocsáry, 1912; Berland and Bernard,
1938; Blüthgen, 1961; Morgan, 1984; Rosa, 2006).
Development of Ch. ruddii at the expense of Eumenes
coronatus (Panzer, 1799) is recorded herein for the
first time.
Ch. sexdentata group
Chrysis sexdentata Christ, 1791
Material. Krasnoperekopsk Distr., Pochetnoe, in
nest of Euodynerus dantici, 30.IV.2010 (V. Zhidkov),
1 ♂ (dead) (VTNU); Tarkhankut Peninsula, Kipchak
ravine, in nest of Eu. dantici in abandoned nest cell of
Sceliphron destillatorium, 2004 (A. Fateryga), 1 ♀
(dead) (DonNU); Bakhchisarai Distr., Tabachnoe,
from nest of Eu. dantici, 4.VI.2010 (V. Zhidkov), 1 ♀
(VTNU); Simferopol Distr., Urozhainoe, from nests of
Eu. dantici, 16–19.VI.2003 (A. Fateryga), 4 ♀
(DonNU).
Distribution. The species was recorded in Europe,
North Africa, and West Asia (Berland and Bernard,
1938; Linsenmaier, 1959; Móczár, 1967; Kimsey and
Bohart, 1991).
Host range. There are published data on develop-
ment of Ch. sexdentata at the expense of the potter
wasps Ancistrocerus parietum and Paragymnomerus
spiricornis, and also the mason bees Hoplitis adunca,
Osmia caerulescens (Linnaeus, 1758), and
O. brevicornis (Fabricius, 1798) (Berland and Ber-
nard, 1938; Balthasar, 1954; Blüthgen, 1961; Kunz,
1994). At the same time, Cerceris arenaria (Linnaeus,
1758) should not probably be listed among the hosts
since the author who published this observation
merely noted the presence of the cuckoo wasp near the
nest of this potential host (Abeille de Perrin, 1878,
cited after Berland and Bernard, 1938). In Crimea,
Ch. sexdentata is a specialized parasite of Euodynerus
dantici (Rossi, 1790).
Subgenus Cornuchrysis Balthasar, 1953
Ch. cerastes group
Chrysis ambigua Radoszkowski, 1897
Material. Simferopol Distr., Urozhainoe, from nest
of Syneuodynerus egregius, 20.VI.2003 (A. Fateryga),
1 ♀ (DonNU).

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481
Notes. In the work of Ivanov and Fateryga (2006)
this species was indicated as Chrysis cerastes.
Distribution. This species is known by the few
findings in Southern Europe, West and Middle Asia
(Radoszkowski, 1891; Linsenmaier, 1959, 1968; Kim-
sey and Bohart, 1991; Rosa and Soon, 2004); it was
not previously recorded in Eastern Europe.
Host range. No data on the hosts of this species
have been published earlier. We reared it for the first
time from the nests of Syneuodynerus egregius (Her-
rich-Schaeffer, 1839).
Ch. taczanovskii group
Chrysis taczanovskii Radoszkowski, 1876
Material. Tarkhankut Peninsula, Kipchak ravine,
from nests of Euodynerus disconotatus in abandoned
nest cells of Sceliphron destillatorium, 18–22.VI.2008
(A. Fateryga), 2 ♂ (VTNU, DonNU).
Distribution. The species was recorded in Southern
and Eastern Europe, Middle East, and Middle Asia
(Balthasar, 1954; Linsenmaier, 1959; Móczár, 1967;
Kimsey and Bohart, 1991; Rosa and Soon, 2004).
Host range. There are published data on the devel-
opment of Ch. taczanovskii at the expense of the wasp
Sceliphron destillatorium (Fateryga and Kovblyuk,
2014).
Specific biological traits. An interesting feature of
this species is the fact that it was reared from the nest
cells of Eu. disconotatus located in abandoned nest
cells of S. destillatorium. The nests of Eu. disco-
notatus can also be located in other available cavities,
for instance, in hollow plant stems (Fateryga, 2012c);
nevertheless, the only finding of Ch. taczanovskii in
the potter wasp nests was made in the nests of
Eu. disconotatus constructed in the nests of S. de-
stillatorium. Additional observations are needed to
specify the relations of Ch. taczanovskii with such
different hosts.
Genus STILBUM Spinola, 1806
Stilbum calens Fabricius, 1781
Material. Env. of Katsiveli, in nest of Katamenes
flavigularis, VII.2006 (A. Fateryga), 2 ♀ (dead)
(DonNU).
Distribution. The species is known from Central,
Southern, and Eastern Europe, North Africa, and Mid-
dle East (Buysson, 1891–1896; Semenov-Tian-
Shansky and Nikolskaya, 1954; Linsenmaier, 1959,
1968; Rosa and Soon, 2004; Rosa, 2006).
Host range. A number of authors indicated the pot-
ter wasps Delta unguiculatum and Katamenes arbus-
torum (Panzer, 1799), the dauber Sceliphron madras-
patanum tubifex (Latreille, 1809), and also the bee
Megachile (Chal
icodoma) parietina (Geoffroy, 1785)
as hosts of S. calens (Buysson, 1891–1896; Berland
and Bernard, 1938; Blüthgen, 1961; Rosa, 2006). One
more species, K. flavigularis, is added herein to this
list.
Specific biological traits. Dead females of this spe-
cies were found in the cells of the nest which had a
heavy common cover made of stones and earth paste
(Fateryga and Ivanov, 2009). Empty cocoons from
which cuckoo wasps had emerged were found in the
cells of other nests with a weaker common cover or
without a cover. Stilbum calens seems to be the main
parasite of K. flavigularis, destroying up to 8.7% of its
progeny (Fateryga and Ivanov, 2009); in the latter
publication S. calens was erroneously called an inqui-
line.
Some biological features of a closely related spe-
cies, S. cyanurum (Forster, 1771), are known. For ex-
ample, Móczár (1961) noted oviposition on the co-
cooned prepupa of S. destillatorium. Further develop-
ment of the S. cyanurum larva evidently proceeded in
an orthoparasitic way, since the cited author photo-
graphed the host cocoon with an opening and the
cuckoo wasp cocoon removed from it. A female of
S. cyanurum was also observed to make an opening in
the cell wall of a potter wasp Abispa ephippium (Fab-
ricius, 1775) and to lay an egg on the late pupa of this
host (Matthews and Matthews, 2009). The same type
of development seems to be realized in S. calens de-
veloping in the cells of K. flavigularis. Cocoons of
K. flavigularis were found by us in the cells where
larvae of S. calens were developing; this means that
the host larva consumed the stored provisions and
successfully completed its development, and was eaten
by the cuckoo wasp larva only after it had completed
its cocoon.
DISCUSSION
The 15 species of Chrysididae recorded by us were
obtained from the nests of 16 species of Eumeninae,
which comprised 44% of the species whose nests were
studied by us in Crimea and 21% of the entire Crimean

MARTYNOVA, FATERYGA
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482
fauna of the subfamily. As a result, 13 new host-
parasite associations were discovered, and the previ-
ously known hosts were confirmed for 8 species of
cuckoo wasps. Data on the biology of Chrysis am-
bigua were obtained for the first time, and the species
itself was recorded for Eastern Europe for the first
time.
The nests of most potter wasp species, such as Dis-
coelius dufourii, Paravespa rex, Gymnomerus
laevipes, Syneuodynerus egregius, Euodynerus dan-
tici, Eu. posticus, Ancistrocerus auctus, A. gazella,
A. nigricornis, A. parietinus, Eumenes coarctatus
lunulatus, and Eu. coronatus, revealed only one spe-
cies of cuckoo wasps. At the same time, it was noted
that two species could develop at the expense of
Katamenes flavigularis and probably Tropidodynerus
interruptus, and three species, at the expense of
Euodynerus disconotatus and Ancistrocerus antilope.
No cuckoo wasps were found in the nests of the re-
maining 20 species of Eumeninae, possibly due to the
insufficient material examined. As can be seen from
the data in the table, the species of Eumeninae in
whose nests cuckoo wasps were found were repre-
sented by a greater number of nests than those which
revealed no cuckoo wasps.
Analysis of data on the host-parasite associations
revealed some specific traits of cuckoo wasp speciali-
zation to eumenine hosts. First, cuckoo wasp larvae
can be conditionally subdivided into oligophages,
which develop at the expense of one, less frequently
two host species, and polyphages, whose host range
may include many species in the given natural zone.
The oligophagous species in our material were
Ch. sexdentata and Ch. splendidula which infested
nests of the same host species even in different land-
scape zones of Crimea. In contrast, the polyphagous
Ch. rutilans developed in nests of non-related species
of Eumeninae which also had different modes of nest-
ing.
The modern system of Chrysididae is based on the
morphological criterion, whereas biological data are
used only as additional characters; therefore, the cor-
rectness of boundaries of taxa of polyphagous species
raises doubts in some cases. In taxonomically difficult
species groups, such as ignita and splendidula, the use
of the biological criterion would obviously be an ad-
vantage. Such problems can be solved only by pur-
poseful studies utilizing a systemic approach.
Second, cuckoo wasps are morphologically special-
ized to the habitats of their hosts (Kilimnik, 1993),
which is reflected in their life forms. For example, the
geobionts Ch. cylindrica, Ch. graelsii, and Ch. vale-
siana infest the nests of potter wasps positioned in the
soil, whereas such species as Ch. ignita, Ch. pseudo-
brevitarsis, C
h. brevitarsis, and Ch. longula prefer the
nests built in the preexisting cavities above the soil
surface. In both cases, host species with a similar nest
structure are usually quite close taxonomically. Stil-
bum calens should be regarded as a special life form
associated with the hosts that build their own nests,
rather than use existing cavities.
Third, it is obvious that all the reared species of
cuckoo wasps, with the exception of S. calens (and
also Ch. cylindrica, which was captured close to the
nests), develop as inquilines in the potter wasp nests.
Not a single cuckoo wasp cocoon was found inside the
host cocoon. Some observations of developing larvae
have shown that at preimaginal stages the cuckoo
wasps are difficult to distinguish from their hosts,
since both feed on provisions; therefore, the cells in-
fested with cuckoo wasps can be identified only after
pupation of their larvae. For this reason, indication of
bees as hosts for such species as Ch. graelsii,
Ch. rutilans, Ch. ignita, Ch. longula, Ch. ruddii, and
Ch. sexdentata raises doubts. The cuckoo wasps that
consume provisions in the potter wasp nests evidently
cannot do the same in the bee nests which contain
provisions of plant origin. On the other hand, it would
be hard to assume that cuckoo wasps were so ecologi-
cally flexible that they could change their development
strategy depending on the host. No such problems
arise in the case of S. calens, since its orthoparasitic
larvae feed on the cocooned prepupae and pupae of the
host and thus do not depend on the nature of the provi-
sions stored. Consequently, the host range of such
species can include both wasps and bees.
Thus, rearing of cuckoo wasps directly from host
nests makes it possible not only to reliably character-
ize the host-parasite associations, but also to solve a
number of special taxonomic and biological problems.
ACKNOWLEDGMENTS
The authors are grateful to S.P. Ivanov (Vernadsky
Taurida National University, Simferopol, Crimea) and
V.Yu. Zhidkov (Charivna Gavan National Nature
Park, Chernomorskoe, Crimea) for help in material
collection, and also to A.N. Kilimnik (Odessa State
Ecological University, Ukraine) and P. Rosa (Ber-
nareggio, Italy) for help in material identification,
finding some literature, and consultations.

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