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First record of an encyrtid wasp (hymenoptera: chalcidoidea) as a true primary parasitoid of ants (hymenoptera: formicidae).

Records of associations with ants involving hymenopteran wasp parasitoids include more than 500 wasp species, but only a fraction unambiguously pertains to true parasitoids (Lachaud & Perez-Lachaud 2012). Parasitoid wasps known to attack adult ants or their brood belong to 8 families: Diapriidae (Diaprioidea); Chalcididae, Eucharitidae, Eulophidae, Eurytomidae and Peril ampidae (Chalcidoidea); Braconidae and Ichneumonidae (Ichneumonoidea) (Wilson 1971; Kistner 1982; Holldobler & Wilson 1990; Schmid-Hempel 1998; Lachaud & Perez-Lachaud 2012). The Eucharitidae sensu stricto is the only group whose known hosts are exclusively larval ants (Heraty 2002).

Here we describe the first known case of true primary parasitism of a species of ant by an encyrtid wasp (Chalcidoidea: Encyrtidae). Our results are based on ant specimens collected in 2002 in the course of investigations on the eucharitid fauna associated with formicids in French Guiana. An exhaustive recent review of all the material preserved in alcohol, focusing on eucharitid wasps associated with poneromorph ants (Lachaud et al. 2012), fortuitously allowed us to detect new examples of hymenopteran parasitoids of ants, other than eucharitids. Moreover, a survey of the literature over the last 100 years allowed us to list both all the recorded cases of associations between encyrtid wasps and ants and the exact nature of their known relationships.

MATERIALS AND METHODS

Two nests of the arboreal ant-garden species, Pachycondyla goeldii (Forel) (Formicidae: Ponerinae), were collected on 12 Nov 2002 in French Guiana, at Km 13 along the road leading to the Hydroelectric complex at Petit Saut, Sinnamary (N 05[degrees] 07' 25.3" W 52[degrees] 57' 16.7"). Pachycondyla goeldii is a monogynous, polydomous ponerine species (Denis et al. 2006) that colonizes pioneer vegetal formations where plants are characterized by a rapid, continuous growth and a high production of leaves and soft wood (Corbara & Dejean 1996; Dejean et al. 2000b). Founding queens and first generation workers initiate their own ant garden by building a cardboard-like structure into which epiphyte seeds are integrated. Following the growth of the epiphyte, the colony establishes its nest within the root system (Corbara & Dejean 1996; Orivel et al. 1998). Ant gardens were transported to the laboratory, and their content preserved in alcohol for later examination.

The presence and number of ant dealate females, alate females, males, workers, cocoons, and larvae, as well as the presence of any adult myrmecophile (especially eucharitid parasitoids), were recorded. At first, the contents of cocoons preserved in alcohol were superficially examined while backlit to check for the presence of any ectoparasitoid. In particular, we looked for wasp remains (exuvia) within empty ant cocoons denoting previous parasitoid emergence and for the presence of scars upon ant larvae resulting from unsuccessful attacks of eucharitid first instar larvae. During June 2011, preserved material was revised again: all of the cocoons were dissected under a stereomicroscope and ant pupae were this time thoroughly checked for the presence of both ecto- and endoparasitoids or for any evidence of parasitism. We discovered some encyrtids, some of these were slide-mounted and identified by direct comparison with type material of the relevant species. Voucher specimens were deposited in the Natural History Museum, London, England and in the Arthropod collection of El Colegio de la Frontera Sur-Chetumal, Quintana Roo, Mexico.

RESULTS

The first nest collected was composed of 1 queen, 77 workers, 4 pupae, 2 larvae, and several eggs. The second nest contained no queen, 5 alate females, 208 workers, 67 pupae (of which 16 large pupae, presumably winged sexuals), and neither larvae nor eggs. Initial cursory inspection of the brood, in 2002, showed no signs of external attack.

During the later examination, in 2011, it was noted that the color of the gaster of some ant pupae looked slightly different and, in one instance, very small dark points were visible through the cuticle. In this last case, dissection showed the presence of already pigmented developing individuals of a gregarious, very minute (body length < 1 mm) endoparasitoid wasp. Five ant pupae from nest no. 2 were found to be parasitized. One worker pupa contained 15 almost fully developed and pigmented adult wasps (8 females, 7 males). A second worker pupa contained 11 wasp pupae (at that developmental stage sex could not he determined, see Fig. 1), and a female (queen) pupa contained 4 white wasp larvae-prepupae. Two other queen pupae had a hole in their gaster, from which parasitoids had previously emerged (Fig. 1).

The wasp was identified as Blanchardiscus pollux Noyes (Chalcidoidea: Encyrtidae: Encyrtinae). Only 2 species of the genus Blanchardiscus de Santis are currently known: B. scutellaris De Santis and B. pollux. Blanchardiscus scutellaris was described from a female collected with sweep net upon vegetation in Tucuman, Argentina (De Santis 1964), and has been reported from Brazil (Noyes 1980), whereas B. pollux is known from both the male and the female and was collected from Costa Rica and Belize (Noyes 2004). The biology of both species was heretofore unknown and no host data were available for any of the two species.

According to our observations, B. pollux is a gregarious endoparasitoid of P. goeldii pupae. Wasp larvae pupate inside the ant host pupae, which are protected by a cocoon. Developing wasps were found in the dorsal anterior portion of the gaster of the ant host, near the petiole. They were grouped in a compact mass, and for the 15 nearly adult specimens whose sexual gender could be assessed, the wings were somewhat distorted. Only a small portion of the ant's gaster was occupied by the parasitoids: B. pollux adults measure less than 1 mm, whereas P. goeldii workers measure 9-12 mm. At emergence, adult wasps chew a hole (1 mm in diam), in the host cuticle and through the host cocoon. A single emergence hole was observed in the apical, ventral portion of the gaster, near the meconium (Fig. 1), in the 2 queen pupae from which wasps had already emerged. Both worker and queen pupae of P. goeldii were parasitized and both male and female wasps emerged from the same host. It is unknown whether more than a single generation occurs inside the ant nest or whether adults leave the host nest to mate and disperse, but the latter seems more likely because both males and females were collected outside ant nests (Noyes 20041. However, even if no free adult wasp was found within both P. goeldii nests, they might have passed unnoticed when nests were collected.

DISCUSSION

Encyrtid wasps are widespread and common throughout the world. The family currently includes more than 480 genera and 4500 species and is one of the most important chalcidoid families for biological control of insect pests (Noyes & Hayat 1994; Noyes 2000, 2012; Trjapitzin 2008). Though a few species are egg predators (e.g. in Microterys), most encyrtids are endoparasitoids of insects and arachnids, including other hymenopteran parasitoids. About half of all encyrtids whose host is known are parasitic upon Coccoidea (Noyes 2012). Two subfamilies are recognized: the Tetracneminae with 111 genera and 848 species, and the Encyrtinae with 370 genera and 3700 species (Trjapitzin 1973a, 1973b; Noyes 2012).

Numerous encyrtids have been recorded associated with ants belonging to 3 subfamilies (Dolichoderinae, Formicinae, and Myrmicinae). However, almost all these records referred to indirect associations with ants (i.e. interference, see Table 1) through primary parasitism of the trophohionts that ants tend (e.g. Bartlett 1961; Cudjoe et al. 1993; Hubner & Volkl 1996; Gonzalez-Hernandez et al. 1999; Barzman & Daane 2001; Mgocheki & Addison 2009). These associations are more numerous in Encyrtinae, with 27 species from 17 genera involved in 45 associations with ants, than in Tetracneminae, with 15 species from 8 genera involved in 27 associations. However, such a difference in the number of reported associations likely reflects species richness within each subfamily, because there are more encyrtine than tetracmine species.

Various encyrtid species reported in interference association with ants (Table 1) have a very wide primary host range, and some are even hyperparasitoids: e.g. Prionomitus mitratus is parasitic on a fairly wide range of psyllids, Prochiloneurus pulchellus is a hyperparasitoid of many species of mealybugs, and Syrphophagus aphidivorus is a hyperparasitoid of virtually any aphid that feeds on grasses or herbaceous vegetation. Likewise, in numerous genera (Aenasius, Anagyrus, Anicetus, Coccidoxenoides, Comperiella, Leptomastix, Ooencyritis, Prochiloneurus, Syrphophagus, see Table 1), the indirect association with the ant is not specific and can even involve species from different ant subfamilies. However, in some genera, e.g. Ananusia, Encyrtus, Holcencyrtus, Metaphycus, Microterys, Prionomitus, and Tetracnemoidea (Table 1), more specific interferences with ants--at species or, at least, at subfamily level--can occur and could suggest some level of selection of the ant associates by the parasitoids.

A few cases of assumed direct associations involving encyrtids and ants have been reported (Table 1), but true primary parasitism has never been demonstrated before. An unidentified species of encyrtid was recorded from a refuse heap of Eciton burchellii (Westwood) (Rettenmeyer et al. 2011), but direct interaction with this ant host has not been observed and the encyrtid may have only been a prey. The 2 females of Taftia prodeniae Ashmead found by Roepke (1919) clanging to the antennae of Dolichoderus thoracicus (Fr. Smith) (referred to as D. bituberculatus Mayr) may have been phoretic rather than parasitic. Only Holcencyrtus wheeleri (Ashmead) (referred to as Pheidaloxenus wheeleri), found in nests of the myrmicine ants Pheidole tepicana Pergande (referred to as P. instabilis) (Wheeler 1907) and P. ceres Wheeler (referred to as P ceres var. tepaneca Wheeler) (Mann 1914), seems to have symphilic relationships with its hosts and has been suspected of being "probably also entoparasitic on these ants or their progeny during its larval stages" (Wheeler 1910). Wheeler (1907) stated that this "exquisite little Chalcidid ... runs about in the dense throng of Pheidole workers like one of their number. It is not easily detected, as it resembles the workers in its small size (1 mm) and in being subapterous or practically wingless". In some occasions as many as 6 or 8 H. wheeleri have been observed in a single nest of P. tepicana (Wheeler 1907). It is worth noting that 2 related encyrtid species, Holcencyrtus osborni Timberlake and H. myrmicoides (Compere & Zinna), have been reared from mealybug hosts (e.g. Dysmicoccus boninsis (Kuwana), D. brevipes (Cockerell), Planococcus citri (Risso), Pseudococcus gahani (Green), Saccharicoccus sacchari (Cockerell), see Noyes & Hayat 1994; Noyes 2012) that are often associated with ants, and therefore association with ants might be fortuitous. Nevertheless, in the case of H. wheeleri, considering that it seems to be a regular myrmecophile in P. tepicana nests, that adults present subapterism, and that ant hosts are not associated with Aphididae or Coccidae (Wheeler 1907), such an association is unlikely to be only circumstancial; however, a primary parasitic relationship was never proved and the wasp developmental stages remained unknown. Therefore, our record of B. pollux from French Guiana reared from pupae of the neotropical ant P. goeldii constitutes both the first record of primary parasitism of ants for the Encyrtidae and the first case associating an encyrtid species with a ponerine ant. Our finding increases to 9 the number of parasitic wasp families that attack ants. As hypothesized by Huggert and Masner (1983) and Hanson et al. (1995), a possible evolutionary path to the parasitism of ants by hymenopterous parasitoids could have been through the occurence of a shift from the initial primary host--an ant symbiont--to the ant host through a gradual process of association and integration with the ant hosts. Such a hypothesis seems plausible for numerous families (Lachaud and Perez-Lachaud 2012), and a supporting example has recently been proposed among eulophids for 2 Horismenus species associated with the weaver ant Camponotus sp. near textor (Hansson et al. 2011). Phylogenetically, Blanchardiscus is very probably near genera that include species known to parasitize scale-insects. Our record of B. pollux parasitizing ants may also support this hypothesis.

Little is known about the diversity of parasitoids of ants in general, though knowledge on this topic has significantly increased in the last two decades (Lachaud & Perez-Lachaud 2012). Likewise, the hosts of most parasitic hymenopteran species already described are still unknown, especially those of rare or rarely collected species. Several recent reports (Hansson et al. 2011; Lachaud et al. 2012; Lachaud & Perez-Lachaud 2012; Gates & Perez-Lachaud 2012) have called attention to the diversity of parasitoids that attack ants, particularly in the case of arboreal ant species. For example, hymenopterous myrmecophiles associated with the neotropical weaver ant Camponotus sp. near textor, another arboreal ant, include 2 species of Eucharitidae, 2 of Eulophidae, and 1 of Eurytomidae (Hansson et al. 2011; Gates & Perez-Lachaud 2012; Perez-Lachaud & Lachaud, unpublished data). Those findings and the present record of a new family of parasitic wasps attacking ants inhabiting ant-gardens strongly suggest that arboreal ant nests may constitute a hot spot of diversity that has little been studied. As highlighted by Schmid-Hempel (1998) and Lachaud & Perez-Lachaud (2012), parasitic wasps associated with ants as the primary host are diverse, but most associations still await discovery.

ACKNOWLEDGMENTS

We thank all the team of the Laboratoire Environnement at Petit Saut (HYDRECO) for both logistic help and assistance with local accommodation. We are also grateful to Jerome Orivel (ECOFOG, French Guiana) for collecting the ants, and to Enrique Ruiz Cancino (Universidad Autonoma de Tamaulipas, Mexico) for providing bibliographic information. This research was partially supported by a grant from the French Ministere de l'Ecologie et du Developpement Durable (Program "Recherche de procedes limitant l'activite de fourmis tropicales d'importance ecologique et economique").

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GABRIELA PEREZ-LACHAUD (1), JOHN NOYES (2) AND JEAN-PAUL LACHAUD (1,3) *

(1) El Colegio de la Frontera Sur, Avenida Centenario Km 5.5, Chetumal 77014, Quintana Roo, Mexico

(2) Natural History Museum, Department of Entomology, Cromwell Road, South Kensington, London SW7 5BD, UK

(3) Centre de Recherches sur la Cognition Animale, CNRS-UMR 5169, Universite de Toulouse UPS, US route de Narbonne, 31062 Toulouse Cedex 09, France

* Corresponding author; E-mail: jean-paul.lachaud@univ-tlse3.fr; jlaehaud@ecosur.rnx

TABLE 1. LIST OP ENCYRTID WASPS KNOWN TO BE ASSOCIATED WITH
ANTS, OF THEIR ANTS ASSOCIATES, AND OF THEIR PRIMARY HOSTS,
AND NATURE OF THE ASSOCIATION.

Encyrtid wasp           Ant associate            Primary host
species

Encyrtinae

Acerophagus          Linepithema            Pseudococcus
flavidulus           humile (Mayr)          viburni
(Brethes)            [D]                    (Signoret)
(= Pseudaphycus
flavidulus)

Ageniaspis           Solenapsis             Phyllocnistis
citricola            invicta                citrella Stainton
Logvinovskaya        Buren [M]

Anicetos beneficas   Lasius japonicas       Ceroplastes rubens
Ishii & Yasumatsu    Santschi (= L.         Maskell
                     niger (L.)) [F]
                     Tapinoma sp. [D]       Ceroplastes rubens
                                            Maskell

Arketypon vaderi     Camponotus sp.         Myzolecanium sp.
Guerrieri & Noyes    [F]

Asty machas          Crematogaster          Aclerdid species
phainae Sugonjaev    sp. [M]

Blanchardiscus       Pachycondyla           Pachycondyla
pollux Noyes         goeldii (Forel) [P]    goeldii (Forel)

Bothriothorax        Lasius niger           Xanthogramma sp.
intermedius          (Linnaeus) [F]
Claridge

Compendia            Formica aerata         Aonidiella aurantii
bifasciata Howard    (Francoeur) [F]        (Maskell)
                     Iridomyrmex            Aonidiella aurantii
                     rufoniger (Lowne)      (Maskell) and Coccus
                     gr. spp. [D]           hesperidum L.
                     Linepithema humile     Aonidiella aurantii
                     (Mayr) [D]             (Maskell)
                     Linepithema humile     Aonidiella citrina
                     (Mayr) [D]             Coquillet
                     Solenopsis xyloni      Aonidiella aurantii
                     McCook [M]             (Maskell)

Encyrtus ingae       Crematogaster          Coccus formicarii
Sugonjaev            dohrni Mayr [M]        (Green) and Coccus
                                            hesperidum L.
                     Crematogaster          Coccus formicarii
                     dohrni Mayr [M]        (Green)
                     Crematogaster sp.      Coccus formicarii
                     [M]                    (Green) and Coccus
                                            hesperidum L.

Encyrtus ludmilae    Crematogaster          Megalocryptes
Sugonjaev            sp. [M]                bambusicola (Green)

Homalotylus          Azteca instabilis      Azya orbigera
shuvakhinae          (F. Smith) [D]         Mulsant
Trjapitzin &
Triapitsyn

Metaphycus           Linepithema humile     Saissetia oleae
anneckei Guerrieri   (Mayr) [D]             (Olivier)
& Noyes

Metaphycus hageni    Linepithema humile     Saissetia oleae
Daane &              (Mayr) [D]             (Olivier)
Caltagirone

Metaphycus           Iridomyrmex            Aonidiella aurantii
helvolus (Compere)   rufoniger (Lowne)      (Maskell) and Coccus
                     gr. spp, [D]           hesperidum. L.
                     Linepithema humile     Saissetia oleae
                     (Mayr) (=              (Olivier)
                     Iridomyrmex
                     humilis) [D]

Metaphycus           Iridomyrmex            Aonidiella aurantii
lounsburyi           rufoniger (Lowne)      (Maskell) and Coccus
(Howard)             gr. spp, [D]           hesperidum L.
                     Linepithema humile     Saissetia oleae
                     (Mayr) [D]             (Olivier)

Metaphycus           Linepithema humile     Coccus hesperidum
luteolus             (Mayr) (=              Linnaeus
(Timberlake)         Iridomyrmex
                     humilis) [D]

Metaphycus           Crematogaster          undet. Coccidae
monastyrskii         sp. [M]
Sugonjaev

Metaphycus           Linepithema humile     Saissetia oleae
stanleyi Compere     (Mayr) (=              (Olivier)
                     Iridomyrmex
                     humilis) [D]
                     Linepithema humile     Coccus hesperidum L.
                     (Mayr) (=
                     Iridomyrmex
                     humilis) [D]

Microterys           Linepithema humile     Saissetia oleae
nietneri             (Mayr) (=              (Olivier), Coccus
(Motschulsky)        Iridomyrmex            hesperidum L.
(= Microterys        humilis) [D]
flavas Howard)

Microterys roseni    Dolichoderus           Eucalymnatus
Sugonjaev            thoracicus (F.         tessellatus (Signoret)
                     Smith) (= D.
                     bituberculatus
                     Mayr) [D]
                     Dolichoderus sp.       Eucalymnatus
                     [D]                    tessellatus (Signoret)

Neodusmetia          Solenopsis invicta     Antoninagraminis
sangwani (Subba      Buren [M]              (Maskell)
Rao) Ooencyrtus      Camponotus             Euphyonarthex
sp.                  acvapimensis           phyllostoma Schmidt
                     Mayr [F]
                     Camponotus brutus      Euphyonarthex
                     Forel [F]              phyllostoma Schmidt
                     Camponotus brutus      Caternaultiella rugosa
                     Forel [F]              Schouteden
                     Myrmicaria             Caternaultiella rugosa
                     opaciventris Emery     Schouteden
                     [M]

Prionomitus          Formica pratensis      Cacopsylla crataegi
mitratus (Dalman)    Retzius [F]            (Schrank), C. pyrisuga
                                            (Forster)
                     Lasius niger           Cacopsylla crataegi
                     (Linnaeus) [F]         (Schrank)

Prionomitus          Lasius niger           Cacopsylla crataegi
tiliaris (Dolman)    (Linnaeus) [F]         (Schrank)

Prochiloneurus       Camponotus             Anagyrus lopezi
pulchellus           acvapimensis           (De Santis)
Silvestri (= P.      Mayr [F]
insolitus (Alam))    Camponotus             Anagyrus lopezi
                     flavomargin atus       (De Santis)
                     Mayr [F]
                     Crematogaster          Anagyrus lopezi
                     luctans Forel or       (De Santis)
                     C. kneri Mayr [M]
                     Pheidole               Anagyrus lopezi
                     megacephala            (De Santis)
                     (Fabricius) [M]

Syrphophagus         Lasius niger           Lysiphlebus cardui
aphidivorus (Mayr)   (Linnaeus) [F]         (Marshall)
(= Aphidencyrtus     Myrmica rubra          Lysipklebus cardui
aphidivorus)         (Linnaeus) (= M.       (Marshall)
                     laevinodis) [M]

Syrphophagus         Lasius niger           Lysiphlebus fabarum
mamitus (Walker)     (Linnaeus) [F]         (Marshall)

Tetracneminae

Aenasius             Azteca xanthochroa     Cataenococcus sp.
tachigaliae          (Roger) [D]
(Brues) (= A.        Crematogaster          Pseudococcus sp. nr.
brasiliensis         curvispinosa           neobrevipes Beardsley
Mercet)              Mayr [M]
                     Crematogaster          Dysmicoccus brevipes
                     sp. [M]                (Cockerell)
                     Paratrechina           Dysmicoccus brevipes
                     sp. [F]                (Cockerel1)

Anagyrus             Technomyrmex albipes   Nipaecoccus vastator
agraensis Saraswat   F. Smith [D]           (Maskell)
(= A, indicus
Shafee, Alam &
Agarwal)

Anagyrus ananatis    Pheidole               Dysmicoccus brevipes
Gahan                megacephala            (Cockerel1)
                     (Fabricius) [M]

Anagyrus lopezi      Camponotus             Phenacoccus manihoti
(De Santis) (as      acvapimensis
Epidinocarsis        Mayr [F]
lopezi)              Camponotus             Matile-Ferrero
                     flavomarginatus        Phenacoccus manihoti
                     Mayr [F]
                     Crematogaster          Matile-Ferrero
                     luctans Forel or C.    Phenacoccus manihoti
                     kneri Mayr [M]
                     Pheidole               Matile-Ferrero
                     megacephala            Phenacoccus manihoti
                     (Fabricius) [M]

Anagyrus             Lasius niger           Matile-Ferrero
pseudoeocci          (Linnaeus) (= L.       Planococcus citri
(Girault)            niger (Latreille)      (Risso)
                     [F]
                     Formica perpilosa      Planococcus ficus
                     Wheeler [F]            (Signoret)

Anagyrus sp.         Anoplolepis            Planococcus ficus
near pseudoeocci     steingroeveri          (Signoret)
                     (Forel) [F]
                     Crematogaster          Planococcus ficus
                     peringueyi Emery       (Signoret)
                     [M]
                     Linepithema humile     Planococcus ficus
                     (Mayr) [D]             (Signoret)

Ananusia australis   Ochetellus glaber      ?
(Gordh &             (Mayr) (=
Trjapitzin) (=       Iridomyrmex
Mynttencyrtus        glaber) [D]
australis)

Ananusia             Iridomyrmex            ?
longiscapus          rufoniger
(Girault)            domesticas Forel
                     (= domestica) [D]

Coccidoxenoides      Anoplolepis            Planococcus ficus
perminutus           steingroeveri          (Signoret)
Girault (= C.        (Forel) [F]
perminutus           Crematogaster          Planococcus ficus
(Timberlake))        peringueyi Emery [M]   (Signoret)
                     Linepithema humile     Planococcus ficus
                     (Mayr) [D]             (Signoret)

Holcencyrtus         Pheidole tepicana      ?
wheeleri (Ashmead)   Pergande (= P.
(= Pheidaloxenus     instabilis, = P.
wheeleri)            kingi) [M]
                     Pheidole ceres         ?
                     Wheeler (= Pheidole
                     ceres var.
                     tepaneca) [M]

Leptomastix          Lasius niger           Planococcus citri
dactylopii Howard    (Linnaeus) (= L.       (Risso)
                     niger (Latreille)
                     [F]

Leptomastix epona    Linepithema humile     Pseudococcus viburni
(Walker)             (Mayr) [D]             (Signoret)

Taftia prodeniae     Dolichoderus           Dolichoderus thoracicus
Ashmead              thoracicus (F.         (F. Smith) (= D.
                     Smith) (= D.           bituberculatus Mayr)
                     bituberculatus
                     Mayr) [D]

Tetracnemoidea       Linepithema humile     Pseudococcus gahani
peregrina            (Mayr) {=              Green
(Compere) (=         Iridomyrmex
Tetraenemus          humilis) [D]
peregrinus)

Tetracnemoidea       Linepithema humile     Pseudococcus gahani
brevicornis          (Mayr) (=              Green
(Girault) (=         Iridomyrmex
Tetraenemus          humilis) [D]
pretiosus
Timberlake)

Encyrtid wasp           Ant associate         Relationship
species

Encyrtinae

Acerophagus          Linepithema            Interference
flavidulus           humile (Mayr)
(Brethes)            [D]
(= Pseudaphycus
flavidulus)

Ageniaspis           Solenapsis             Interference
citricola            invicta
Logvinovskaya        Buren [M]

Anicetus beneficas   Lasius japonicas       Interference
Ishii & Yasumatsu    Santschi (= L.
                     niger (L.)) [F]
                     Tapinoma sp. [D]       Interference

Arketypon vaderi     Camponotus sp.         Found in the
Guerrieri & Noyes    [F]                    nest

Astymachas           Crematogaster          Found in shelters
phainae Sugonjaev    sp. [M]

Blanchardiscus       Pachycondyla           True parasitoid
pollux Noyes         goeldii (Forel) [P]

Bothriothorax        Lasius niger           Interference
intermedius          (Linnaeus) [F]
Claridge

Compendia            Formica aerata         Interference
bifasciata Howard    (Francoeur) [F]
                     Iridomyrmex            Interference
                     rufoniger (Lowne)
                     gr. spp. [D]
                     Linepithema humile     Interference
                     (Mayr) [D]
                     Linepithema humile     Interference
                     (Mayr) [D]
                     Solenopsis xyloni      Interference
                     McCook [M]

Encyrtus ingae       Crematogaster          Found in the nest
Sugonjaev            dohrni Mayr [M]

                     Crematogaster          Found in shelters
                     dohrni Mayr [M]
                     Crematogaster sp.      Found in shelters
                     [M]

Encyrtus ludmilae    Crematogaster          Found in the nest
Sugonjaev            sp. [M]

Homalotylus          Azteca instabilis      Interference
shuvakhinae          (F. Smith) [D]
Trjapitzin &
Triapitsyn

Metaphycus           Linepithema humile     Interference
anneckei Guerrieri   (Mayr) [D]
& Noyes

Metaphycus hageni    Linepithema humile     Interference
Daane &              (Mayr) [D]
Caltagirone

Metaphycus           Iridomyrmex            Interference
helvolus (Compere)   rufoniger (Lowne)
                     gr. spp, [D]
                     Linepithema humile     Interference
                     (Mayr) (=
                     Iridomyrmex
                     humilis) [D]

Metaphycus           Iridomyrmex            Interference
lounsburyi           rufoniger (Lowne)
(Howard)             gr. spp, [D]
                     Linepithema humile     Interference
                     (Mayr) [D]

Metaphycus           Linepithema humile     Interference
luteolus             (Mayr) (=
(Timberlake)         Iridomyrmex
                     humilis) [D]

Metaphycus           Crematogaster          Found in shelters
monastyrskii         sp. [M]
Sugonjaev

Metaphycus           Linepithema humile     Interference
stanleyi Compere     (Mayr) (=
                     Iridomyrmex
                     humilis) [D]
                     Linepithema humile     Interference
                     (Mayr) (=
                     Iridomyrmex
                     humilis) [D]

Microterys           Linepithema humile     Interference
nietneri             (Mayr) (=
(Motschulsky)        Iridomyrmex
(= Microterys        humilis) [D]
flavas Howard)

Microterys roseni    Dolichoderus           Found in the nest
Sugonjaev            thoracicus (F.
                     Smith) (= D.
                     bituberculatus
                     Mayr) [D]
                     Dolichoderus sp.       Found in the nest
                     [D]

Neodusmetia          Solenopsis invicta     Interference
sangwani (Subba      Buren [M]
Rao) Ooencyrtus      Camponotus             Interference
sp.                  acvapimensis
                     Mayr [F]
                     Camponotus brutus      Interference
                     Forel [F]
                     Camponotus brutus      Interference
                     Forel [F]
                     Myrmicaria             Interference
                     opaciventris Emery
                     [M]

Prionomitus          Formica pratensis      Interference
mitratus (Dalman)    Retzius [F]

                     Lasius niger           Interference
                     (Linnaeus) [F]

Prionomitus          Lasius niger           Interference
tiliaris (Dolman)    (Linnaeus) [F]

Prochiloneurus       Camponotus             Interference
pulchellus           acvapimensis
Silvestri (= P.      Mayr [F]
insolitus (Alam))    Camponotus             Interference
                     flavomargin atus
                     Mayr [F]
                     Crematogaster          Interference
                     luctans Forel or
                     C. kneri Mayr [M]
                     Pheidole               Interference
                     megacephala
                     (Fabricius) [M]

Syrphophagus         Lasius niger           Interference
aphidivorus (Mayr)   (Linnaeus) [F]
(= Aphidencyrtus     Myrmica rubra          Interference
aphidivorus)         (Linnaeus) (= M.
                     laevinodis) [M]

Syrphophagus         Lasius niger           Interference
mamitus (Walker)     (Linnaeus) [F]

Tetracneminae

Aenasius             Azteca xanthochroa     Interference
tachigaliae          (Roger) [D]
(Brues) (= A.        Crematogaster          Found in the nest
brasiliensis         curvispinosa           Found in the nest
Mercet)              Mayr [M]
                     Crematogaster
                     sp. [M]
                     Paratrechina           Found in the nest
                     sp. [F]

Anagyrus             Technomyrmex albipes   Interference
agraensis Saraswat   F. Smith [D]
(= A, indicus
Shafee, Alam &
Agarwal)

Anagyrus ananatis    Pheidole               Interference
Gahan                megacephala
                     (Fabricius) [M]

Anagyrus lopezi      Camponotus             Interference
(De Santis) (as      acvapimensis
Epidinocarsis        Mayr [F]
lopezi)              Camponotus             Interference
                     flavomarginatus
                     Mayr [F]
                     Crematogaster          Interference
                     luctans Forel or C.
                     kneri Mayr [M]
                     Pheidole               Interference
                     megacephala
                     (Fabricius) [M]

Anagyrus             Lasius niger           Interference
pseudoeocci          (Linnaeus) (= L.
(Girault)            niger (Latreille)
                     [F]
                     Formica perpilosa      Interference
                     Wheeler [F]

Anagyrus sp.         Anoplolepis            Interference
near pseudoeocci     steingroeveri
                     (Forel) [F]
                     Crematogaster          Interference
                     peringueyi Emery
                     [M]
                     Linepithema humile     Interference
                     (Mayr) [D]

Ananusia australis   Ochetellus glaber      Interference
(Gordh &             (Mayr) (=
Trjapitzin) (=       Iridomyrmex
Mynttencyrtus        glaber) [D]
australis)

Ananusia             Iridomyrmex            Interference
longiscapus          rufoniger
(Girault)            domesticas Forel
                     (= domestica) [D]

Coccidoxenoides      Anoplolepis            Interference
perminutus           steingroeveri
Girault (= C.        (Forel) [F]
perminutus           Crematogaster          Interference
(Timberlake))        peringueyi Emery [M]
                     Linepithema humile     Interference
                     (Mayr) [D]

Holcencyrtus         Pheidole tepicana      Symphilic
wheeleri (Ashmead)   Pergande (= P.
(= Pheidaloxenus     instabilis, = P.
wheeleri)            kingi) [M]
                     Pheidole ceres         Symphilic
                     Wheeler (= Pheidole
                     ceres var.
                     tepaneca) [M]

Leptomastix          Lasius niger           Interference
dactylopii Howard    (Linnaeus) (= L.
                     niger (Latreille)
                     [F]

Leptomastix epona    Linepithema humile     Interference
(Walker)             (Mayr) [D]

Taftia prodeniae     Dolichoderus           Phoresis
Ashmead              thoracicus (F.
                     Smith) (= D.
                     bituberculatus
                     Mayr) [D]

Tetracnemoidea       Linepithema humile     Interference
peregrina            (Mayr) {=
(Compere) (=         Iridomyrmex
Tetraenemus          humilis) [D]
peregrinus)

Tetracnemoidea       Linepithema humile     Interference
brevicornis          (Mayr) (=
(Girault) (=         Iridomyrmex
Tetraenemus          humilis) [D]
pretiosus
Timberlake)

Encyrtid wasp           Ant associate             References
species

Encyrtinae

Acerophagus          Linepithema            Daane et al, 2007
flavidulus           humile (Mayr)
(Brethes)            [D]
(= Pseudaphycus
flavidulus)

Ageniaspis           Solenapsis             Zappala et al. 2007
citricola            invicta
Logvinovskaya        Buren [M]

Anicetos beneficas   Lasius japonicas       Itioka & Inoue 1996a,
Ishii & Yasumatsu    Santschi (= L.         1996b
                     niger (L.)) [F]
                     Tapinoma sp. [D]       Krull & Basedow 2005

Arketypon vaderi     Camponotus sp.         Guerrieri & Noyes 2002
Guerrieri & Noyes    [F]

Asty machas          Crematogaster          Sugonyaev 1996a
phainae Sugonjaev    sp. [M]                this work

Blanchardiscus       Pachycondyla
pollux Noyes         goeldii (Forel) [P]

Bothriothorax        Lasius niger           Trjapitzin 1978
intermedius          (Linnaeus) [F]
Claridge

Compendia            Formica aerata         Martinez-Ferrer
bifasciata Howard    (Francoeur) [F]        et al. 2003
                     Iridomyrmex            James et al. 1999
                     rufoniger (Lowne)
                     gr. spp. [D]
                     Linepithema humile     Martinez-Ferrer
                     (Mayr) [D]             et al. 2003
                     Linepithema humile     Flanders 1945
                     (Mayr) [D]
                     Solenopsis xyloni      Martinez-Ferrer
                     McCook [M]             et al. 2003

Encyrtus ingae       Crematogaster          Sugonyaev 1998
Sugonjaev            dohrni Mayr [M]

                     Crematogaster          Sugonyaev 1998
                     dohrni Mayr [M]
                     Crematogaster sp.      Sugonyaev 1998
                     [M]

Encyrtus ludmilae    Crematogaster          Sugonyaev 1999
Sugonjaev            sp. [M]

Homalotylus          Azteca instabilis      Liere &
shuvakhinae          (F. Smith) [D]         Perfecto 2008
Trjapitzin &
Triapitsyn

Metaphycus           Linepithema humile     Bartlett 1961 *,
anneckei Guerrieri   (Mayr) [D]             Barzman & Daane 2001
& Noyes

Metaphycus hageni    Linepithema humile     Barzman & Daane 2001
Daane &              (Mayr) [D]
Caltagirone

Metaphycus           Iridomyrmex            James et al. 1999
helvolus (Compere)   rufoniger (Lowne)
                     gr. spp, [D]
                     Linepithema humile     Bartlett 1961,
                     (Mayr) (=              Barzman & Daane 2001
                     Iridomyrmex
                     humilis) [D]

Metaphycus           Iridomyrmex            James et al. 1999
lounsburyi           rufoniger (Lowne)
(Howard)             gr. spp, [D]
                     Linepithema humile     Barzman & Daane 2001
                     (Mayr) [D]

Metaphycus           Linepithema humile     Bartlett 1961
luteolus             (Mayr) (=
(Timberlake)         Iridomyrmex
                     humilis) [D]

Metaphycus           Crematogaster          Sugonyaev 1996b
monastyrskii         sp. [M]
Sugonjaev

Metaphycus           Linepithema humile     Bartlett 1961
stanleyi Compere     (Mayr) (=
                     Iridomyrmex
                     humilis) [D]
                     Linepithema humile     Bartlett 1961
                     (Mayr) (=
                     Iridomyrmex
                     humilis) [D]

Microterys           Linepithema humile     Bartlett 1961
nietneri             (Mayr) (=
(Motschulsky)        Iridomyrmex
(= Microterys        humilis) [D]
flavas Howard)

Microterys roseni    Dolichoderus           Sugonyaev 1996b
Sugonjaev            thoracicus (F.
                     Smith) (= D.
                     bituberculatus
                     Mayr) [D]
                     Dolichoderus sp.       Sugonyaev 1996b
                     [D]

Neodusmetia          Solenopsis invicta     Chantos 2007
sangwani (Subba      Buren [M]
Rao) Ooencyrtus      Camponotus             Dejean et al. 2000a
sp.                  acvapimensis
                     Mayr [F]
                     Camponotus brutus      Dejean et al. 2000a
                     Forel [F]
                     Camponotus brutus      Gibernau & Dejean 2001
                     Forel [F]
                     Myrmicaria             Gibernau & Dejean 2001
                     opaciventris Emery
                     [M]

Prionomitus          Formica pratensis      Novak 1994
mitratus (Dalman)    Retzius [F]

                     Lasius niger           Novak 1994
                     (Linnaeus) [F]

Prionomitus          Lasius niger           Novak 1994
tiliaris (Dolman)    (Linnaeus) [F]

Prochiloneurus       Camponotus             Cudjoe et al. 1993
pulchellus           acvapimensis
Silvestri (= P.      Mayr [F]
insolitus (Alam))    Camponotus             Cudjoe et al. 1993
                     flavomargin atus
                     Mayr [F]
                     Crematogaster          Cudjoe et al. 1993
                     luctans Forel or
                     C. kneri Mayr [M]
                     Pheidole               Cudjoe et al. 1993
                     megacephala
                     (Fabricius) [M]

Syrphophagus         Lasius niger           Hubner & Volkl 1996
aphidivorus (Mayr)   (Linnaeus) [F]
(= Aphidencyrtus     Myrmica rubra          Hubner & Volkl 1996
aphidivorus)         (Linnaeus) (= M.
                     laevinodis) [M]

Syrphophagus         Lasius niger           Sanders & Van Veen 2010
mamitus (Walker)     (Linnaeus) [F]

Tetracneminae

Aenasius             Azteca xanthochroa     Noyes & Ren 1995
tachigaliae          (Roger) [D]
(Brues) (= A.        Crematogaster          Noyes & Ren 1995
brasiliensis         curvispinosa           Noyes 2000
Mercet)              Mayr [M]
                     Crematogaster
                     sp. [M]
                     Paratrechina           Noyes 2000
                     sp. [F]

Anagyrus             Technomyrmex albipes   Nechols & Seibert 1985
agraensis Saraswat   F. Smith [D]
(= A, indicus
Shafee, Alam &
Agarwal)

Anagyrus ananatis    Pheidole               Gonzalez-Hernandez et
Gahan                megacephala            al. 1999
                     (Fabricius) [M]

Anagyrus lopezi      Camponotus             Cudjoe et al. 1993
(De Santis) (as      acvapimensis
Epidinocarsis        Mayr [F]
lopezi)              Camponotus             Cudjoe et al. 1993
                     flavomarginatus
                     Mayr [F]
                     Crematogaster          Cudjoe et al. 1993
                     luctans Forel or C.
                     kneri Mayr [M]
                     Pheidole               Cudjoe et al. 1993
                     megacephala
                     (Fabricius) [M]

Anagyrus             Lasius niger           Campos et al. 2006
pseudoeocci          (Linnaeus) (= L.
(Girault)            niger (Latreille)
                     [F]
                     Formica perpilosa      Tollerup 2007
                     Wheeler [F]

Anagyrus sp.         Anoplolepis            Mgocheki & Addison
near pseudoeocci     steingroeveri          2009
                     (Forel) [F]
                     Crematogaster          Mgocheki & Addison 2009
                     peringueyi Emery
                     [M]
                     Linepithema humile     Mgocheki & Addison 2009
                     (Mayr) [D]

Ananusia australis   Ochetellus glaber      Gordh & Trjapitzin 1979
(Gordh &             (Mayr) (=
Trjapitzin) (=       Iridomyrmex
Mynttencyrtus        glaber) [D]
australis)

Ananusia             Iridomyrmex            Dahms & Gordh 1997
longiscapus          rufoniger
(Girault)            domesticas Forel
                     (= domestica) [D]

Coccidoxenoides      Anoplolepis            Mgocheki & Addison 2009
perminutus           steingroeveri
Girault (= C.        (Forel) [F]
perminutus           Crematogaster          Mgocheki & Addison 2009
(Timberlake))        peringueyi Emery [M]
                     Linepithema humile     Mgocheki & Addison 2009
                     (Mayr) [D]

Holcencyrtus         Pheidole tepicana      Wheeler 1907, Peck 1963
wheeleri (Ashmead)   Pergande (= P.
(= Pheidaloxenus     instabilis, = P.
wheeleri)            kingi) [M]
                     Pheidole ceres         Mann 1914
                     Wheeler (= Pheidole
                     ceres var.
                     tepaneca) [M]

Leptomastix          Lasius niger           Campos et al. 2006
dactylopii Howard    (Linnaeus) (= L.
                     niger (Latreille)
                     [F]

Leptomastix epona    Linepithema humile     Daane et al 2007
(Walker)             (Mayr) [D]

Taftia prodeniae     Dolichoderus           Roepke 1919
Ashmead              thoracicus (F.
                     Smith) (= D.
                     bituberculatus
                     Mayr) [D]

Tetracnemoidea       Linepithema humile     Bartlett 1961
peregrina            (Mayr) {=
(Compere) (=         Iridomyrmex
Tetraenemus          humilis) [D]
peregrinus)

Tetracnemoidea       Linepithema humile     Bartlett 1961
brevicornis          (Mayr) (=
(Girault) (=         Iridomyrmex
Tetraenemus          humilis) [D]
pretiosus
Timberlake)

* Referred to as M. lounsburyi

[D] Dolichoderinae; [F] Formicinae;
[M] Myrmicinae; (P] Ponerinae.


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Author:Perez-Lachaud, Gabriela; Noyes, John; Lachaud, Jean-Paul
Publication:Florida Entomologist
Article Type:Report
Geographic Code:3FREN
Date:Dec 1, 2012
Words:6178
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