Eucharitid Wasps

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All but one species of Eucharitid wasps attack ant larvae.

  • The life cycle of a typical orasemine wasp. On the left are shown behaviours occurring outside the ant nest: mating, oviposition and planidial acquisition by the ants (EFN, extrafloral nectary). On the right are shown behaviours occurring within the ant nest: attachment to the ant larva, development on the ant pupa, pupation and eclosion. (Baker et al., 2019)
Parasitic eucharitid wasp.

Peeters et al (2015) - Among 9 families of wasps parasitic on ants, only Eucharitidae (56 genera) specialize in attacking the immature stages of ants. Unlike most parasitoid wasps, eucharitid females deposit their eggs away from the host, in or on plant tissue (leaves and flower buds). The very active, minute, strongly sclerotized first-instar larvae (‘planidia’) are transported by foraging ants to their nest where they attach to ant larvae and development proceeds once the latter have spun a cocoon (Clausen, 1940; Heraty and Murray, 2013). The eucharitid parasitoids attacking Ponerinae, Ectatomminae, Myrmeciinae and Formicinae belong to a monophyletic derived lineage of Eucharitidae (Murray et al., 2013).

Heraty et al. (2015) - All known Eucharitidae are parasitoids of the immature stages of ants. Larval stages were first described by Wheeler (1907) for Orasema Cameron. The late instar larvae and pupae of Eucharitinae and Oraseminae were then described in a series of early papers summarized by Clausen (1940). The first-instar larvae are highly conservative in their morphology, but late instars can be strikingly ornate with numerous odd protuberances and swellings over the body.

Another oddity of Eucharitidae is the inverse evolutionary association with their ant hosts. Phylogenetic results based on molecular data suggest that the more ancestral eucharitid lineages are associated with more derived groups of ants (e.g., Formicinae and Myrmicinae), and the more derived eucharitid lineages attack the more primitive groups (e.g., Ponerinae) (Murray et al. 2013).

Planidia on adult Camponotus sp.

Schwitzke et al. (2015) found an unspecialized Eucharitid laid its eggs on stipules of the plant Leea manillensis (Leeaceae). The widespread forest plant, occurring throughout the Philippines and parts of Southeast Asia, is a large erect shrub up to 6 m tall. The plant also produces extrafloral nectaries (EFN) that attract a broad array of ant species.

In Leea manillensis, the stipule that encloses the next generation of shoot is the spatial focus of investment into defensive mechanisms via EFN and FB. Obviously not general herbivores are the most damaging threats for Leea manillensis in the study area, but the specialized ant-parasitoid.

The hatching planidia had a massive impact on the whole ant–plant–herbivore system. In our study, about 53 % of the plants’ shoots were observed to be infested with planidia. All of the common ant visitors at the extrafloral nectaries strictly avoided the shoots after the eclosion of the wasps’ larvae. That might explain why we noticed planidia to leave the shoot and disperse. The

Planidia moving along strands.

avoidance of shoots and the observed aggressive behaviour of ants towards adult wasps indicate that the ants were able to recognize this parasitoid as a threat. On 81 % of the planidia-infested shoots, ants were never observed to visit the extrafloral nectaries. Our observations suggest that this avoidance finally led to a cessation of EFN secretion in most of the shoots. Hence, the avoidance of planidia directly resulted in a lasting lack of ants on the shoots, which then caused an almost 10-fold increase in complete shoot losses compared to the unaffected shoots.

We found no evidence that the parasitoid wasp on Leea manillensis had its focus on a specific ant species, genus, or subfamily. In the contrary, we observed a broad range of ant species to get attacked and killed by the planidia, but have no information about the effect of the parasitoid on the ant colony level. As far as it is known, hatching planidia are carried into the colony via foraging ants (Pe´rez-Lachaud et al. 2006; Lachaud and Perez-Lachaud 2012). On Leea, the number of larvae on the shoots declined over time, and we observed planidia to spread over the whole plant along strands. We assume but do not know whether they were carried away by ant foragers. So far, the attack of adult ants has not been reported for the family Eucharitidae. In our experiments with planidia and different ant species, all of the ants died quite fast. On Leea, we found foraging ants killed and attached to leaves and stems. Ants tend to keep foraging trails free of dead nestmates to avoid disease transmission (e.g. Wilson et al. 1958). Besides lurking on the shoot and other plant parts, lurking on corpses might be another way to ensure a contact with foraging ants. Access to ant colonies might also occur via alternate hosts (Heraty et al. 2004). As we observed the planidia to attack all kinds of insects in the vial experiments, they might indeed use a number of different taxa of insect prey as a transport vehicle. However, the killing of potential host ants foraging on the plant does not fit to what is known so far about phoretic behaviour of the Eucharitidae. At this point, we could only speculate what strategies planidia might use to gain access to the ant host colony.

Known Ant Hosts

Ant Wasp Species Locality Source Notes
Anoplolepis Athairocharis vannoorti Universal Chalcidoidea Database primary host
Calomyrmex purpureus Stilbuloida calomyrmecis Universal Chalcidoidea Database primary host
Camponotus Ancylotropus manipurensis Universal Chalcidoidea Database primary host
Camponotus Lophyrocera variabilis Universal Chalcidoidea Database primary host
Camponotus Mateucharis rugulosa Universal Chalcidoidea Database primary host
Camponotus Pseudochalcura gibbosa Universal Chalcidoidea Database primary host
Camponotus Pseudochalcura nigrocyanea Universal Chalcidoidea Database associate
Camponotus Rhipipalloidea madangensis Universal Chalcidoidea Database primary host
Camponotus Rhipipalloidea sp. Universal Chalcidoidea Database primary host
Camponotus Stilbula cyniformis Universal Chalcidoidea Database primary host
Camponotus Stilbuloida doddi Universal Chalcidoidea Database primary host
Camponotus Zulucharis campbelli Universal Chalcidoidea Database primary host
Camponotus aethiops Stilbula cyniformis Universal Chalcidoidea Database primary host
Camponotus atriceps Obeza floridana Universal Chalcidoidea Database primary host
Camponotus herculeanus Stilbula cyniformis Universal Chalcidoidea Database primary host
Camponotus maculatus Stilbula cyniformis Universal Chalcidoidea Database primary host
Camponotus novaeboracensis Pseudochalcura gibbosa Universal Chalcidoidea Database primary host
Camponotus planatus Pseudochalcura sculpturata Universal Chalcidoidea Database primary host
Cataglyphis bicolor Eucharis bedeli Universal Chalcidoidea Database primary host
Cataglyphis bicolor Eucharis bytinskisalzi Universal Chalcidoidea Database associate and primary host
Cataglyphis nodus Eucharis microcephala Universal Chalcidoidea Database primary host
Diacamma scalpratum Schizaspidia diacammae Universal Chalcidoidea Database primary host
Diacamma scalpratum Schizaspidia diacammae Thailand Peeters et al. 2015
Dinoponera lucida Kapala sp. Universal Chalcidoidea Database primary host
Eciton burchellii Isomerala azteca Universal Chalcidoidea Database associate
Eciton quadriglume Orasema rapo Heraty, 1990; Baker et al., 2019; Universal Chalcidoidea Database primary host; questionable, needs confirmation
Ectatomma Tricoryna ectatommae Universal Chalcidoidea Database associate, primary host
Ectatomma brunneum Dicoelothorax platycerus Universal Chalcidoidea Database primary host
Ectatomma ruidum Isomerala azteca Universal Chalcidoidea Database primary host
Ectatomma ruidum Isomerala coronata Universal Chalcidoidea Database primary host
Ectatomma ruidum Kapala sulcifacies Universal Chalcidoidea Database primary host
Ectatomma tuberculatum Dilocantha lachaudii Universal Chalcidoidea Database associate
Ectatomma tuberculatum Isomerala azteca Universal Chalcidoidea Database associate
Ectatomma tuberculatum Isomerala coronata Universal Chalcidoidea Database associate
Formica Eucharis adscendens Universal Chalcidoidea Database primary host
Formica Eucharis esakii Universal Chalcidoidea Database primary host
Formica fusca Eucharis esakii Universal Chalcidoidea Database primary host
Formica oreas Orasema coloradensis Universal Chalcidoidea Database associate
Formica rufa Eucharis adscendens Universal Chalcidoidea Database primary host
Formica rufa Eucharis bedeli Universal Chalcidoidea Database primary host
Formica subnitens Orasema coloradensis Wheeler, 1907; Johnson et al., 1986; Baker et al., 2019; Universal Chalcidoidea Database questionable, needs confirmation
Gnamptogenys menadensis Pogonocharis browni Universal Chalcidoidea Database associate
Hypoponera Neolosbanus gemma Universal Chalcidoidea Database primary host
Hypoponera Neolosbanus palgravei Universal Chalcidoidea Database primary host
Lasius Pseudometagea schwarzii Universal Chalcidoidea Database primary host
Lasius fuliginosus Pseudometagea sp. Universal Chalcidoidea Database primary host
Lasius neoniger Pseudometagea schwarzii Universal Chalcidoidea Database primary host
Messor Eucharis adscendens Universal Chalcidoidea Database primary host
Messor Eucharis punctata Universal Chalcidoidea Database primary host
Messor barbarus Eucharis adscendens Universal Chalcidoidea Database primary host
Messor barbarus Eucharis punctata Universal Chalcidoidea Database primary host
Messor concolor Eucharis punctata Universal Chalcidoidea Database primary host
Monomorium Zuparka monomoria Heraty, 2000; Baker et al., 2019; Universal Chalcidoidea Database primary host
Myrmecia Austeucharis sp. Universal Chalcidoidea Database primary host
Myrmecia Chalcura affinis Universal Chalcidoidea Database primary host
Myrmecia forficata Austeucharis fasciiventris Universal Chalcidoidea Database primary host
Myrmecia forficata Austeucharis myrmeciae Universal Chalcidoidea Database primary host
Myrmecia gulosa Austeucharis fasciiventris Universal Chalcidoidea Database primary host
Myrmecia pilosula Austeucharis implexa Universal Chalcidoidea Database primary host
Odontomachus Kapala sp. Universal Chalcidoidea Database associate
Odontomachus chelifer Latina rugosa Universal Chalcidoidea Database associate, primary host
Odontomachus haematodus Chalcura deprivata Universal Chalcidoidea Database primary host
Odontomachus haematodus Kapala terminalis Universal Chalcidoidea Database primary host
Odontomachus haematodus Schizaspidia convergens Universal Chalcidoidea Database primary host
Odontomachus insularis Kapala terminalis Universal Chalcidoidea Database primary host
Odontomachus rixosus Schizaspidia nasua Heraty et al., 2015; Universal Chalcidoidea Database primary host
Odontomachus ruficeps Chalcura affinis Universal Chalcidoidea Database primary host
Pachycondyla crassinoda Kapala cuprea Myers, 1931:276
Pachycondyla crassinoda Kapala cuprea Universal Chalcidoidea Database primary host
Pheidole Australosema valgius Girault, 1913; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole Hayatosema assectator Das, 1963; Kerrich, 1963; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole Losbanus "sp. nr. rugulosa" Heraty, 1990; Baker et al., 2019 primary host
Pheidole Orasema "sp. 2 nr. bakeri" Heraty, 1990; Baker et al., 2019
Pheidole Orasema "sp. 2 nr. bakeri" Heraty, 1990; Baker et al., 2019 ''Pheidole'' nr. ''californica''
Pheidole Orasema "sp. 2 nr. bakeri" Heraty, 1990; Baker et al., 2019 ''Pheidole'' nr. ''clementensis''
Pheidole Orasema minuta Heraty, 1990; Baker et al., 2019; Universal Chalcidoidea Database primary host; ''Pheidole'' nr. ''tetra''
Pheidole Orasema minutissima Mann, 1918; Heraty, 1994; Wetterer & Porter, 2003; Burks et al. 2018; Baker et al., 2019
Pheidole Orasema sp. Universal Chalcidoidea Database primary host
Pheidole Orasema susanae Universal Chalcidoidea Database primary host; ''Pheidole'' nr. ''tetra''
Pheidole Orasema tolteca Universal Chalcidoidea Database primary host
Pheidole Orasema wayqecha Herreid, 2015; Baker et al., 2019
Pheidole Orasemorpha eribotes Boucek, 1988; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole Orasemorpha myrmicae Heraty, 1994; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole alfaroi Orasema wayqecha Universal Chalcidoidea Database primary host
Pheidole bicarinata Orasema coloradensis Wheeler, 1907; Johnson et al., 1986; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole bicarinata Oresema sp. Arizona Stefan Cover (pers. obs.)
Pheidole capensis Timioderus acuminatus Heraty, 2000; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole ceres Orasema wheeleri Wheeler, 1907; Gahan, 1940; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole dentata Orasema achkola Heraty, 1994; Baker et al., 2019
Pheidole dentata Orasema evansi Burks et al., 2018; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole dentata Orasema robertsoni Van Pelt, 1950; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole desertorum Orasema simulatrix Carey et al., 2012; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole flavens Orasema costaricensis Wheeler & Wheeler, 1937; Kempf, 1972; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole gibba Orasema chunpi Brazil Silva & Casadei-Ferreira, 2020
Pheidole hirtula Orasema tolteca Mann, 1914; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole hyatti Orasema tolteca Universal Chalcidoidea Database primary host
Pheidole megacephala Ivieosema fraudulenta Reichensperger, 1913; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole nitidula Orasema argentina Universal Chalcidoidea Database primary host
Pheidole nitidula Orasema worcesteri Gemignani, 1933; Heraty et al., 1993; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole pilifera Orasema occidentalis Wheeler & Wheeler, 1986; Baker et al., 2019
Pheidole proxima Orasemorpha tridentata Brues, 1934; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole radoszkowskii Orasema worcesteri Gemignani, 1933; Heraty et al., 1993; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole sciophila Orasema wheeleri Universal Chalcidoidea Database primary host
Pheidole tasmaniensis Orasemorpha xeniades Heraty, 1994; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole tepicana Orasema viridis Wheeler, 1907; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole tepicana Orasema wheeleri Wheeler, 1907; Gahan, 1940; Baker et al., 2019; Universal Chalcidoidea Database primary host
Pheidole vallifica Orasema costaricensis Wheeler & Wheeler, 1937; Kempf, 1972; Baker et al., 2019
Pheidole veteratrix Ibitya commnunis Burks et al., 2017; Baker et al., 2019
Pogonomyrmex badius Kapala floridana Universal Chalcidoidea Database primary host
Pogonomyrmex serpens Galearia coleopteroides Universal Chalcidoidea Database primary host
Polyrhachis Rhipipalloidea sp. Universal Chalcidoidea Database primary host
Polyrhachis Stilbula arenae Universal Chalcidoidea Database primary host; ''Polyrhachis'' sp., ''Cyrtomyrma'' sp.
Polyrhachis dives Stilbula polyrhachicida Universal Chalcidoidea Database primary host
Polyrhachis femorata Rhipipalloidea mira Universal Chalcidoidea Database primary host
Rhytidoponera Tricoryna ectatommae Universal Chalcidoidea Database primary host
Rhytidoponera Tricoryna iello Universal Chalcidoidea Database primary host
Rhytidoponera metallica Chalcura nigricyanea Universal Chalcidoidea Database primary host
Rhytidoponera metallica Tricoryna chalcoponerae Universal Chalcidoidea Database primary host
Rhytidoponera metallica Tricoryna minor Universal Chalcidoidea Database primary host
Rhytidoponera victoriae Tricoryna minor Universal Chalcidoidea Database primary host
Rhytidoponera violacea Stilbula cyniformis Universal Chalcidoidea Database primary host
Rhytidoponera violacea Tricoryna sp. Universal Chalcidoidea Database primary host
Solenopsis Orasema pireta Varone et al., 2010; Baker et al., 2019 primary host
Solenopsis geminata Orasema taii Heraty, 1990; Chien & Heraty, 2018; Baker et al., 2019; Universal Chalcidoidea Database primary host; ''Solenopsis geminata'' X ''xyloni'' hybrid
Solenopsis invicta Orasema salebrosa Heraty et al., 1993; Varone et al., 2010; Baker et al., 2019; Universal Chalcidoidea Database primary host
Solenopsis invicta Orasema simplex Heraty et al., 1993; Varone et al., 2010; Baker et al., 2019; Universal Chalcidoidea Database primary host
Solenopsis invicta Orasema sp. Universal Chalcidoidea Database associate, primary host
Solenopsis invicta Orasema xanthopus Silveira-Guido et al., 1964; Heraty et al., 1993; Varone et al., 2010; Baker et al., 2019; Universal Chalcidoidea Database primary host
Solenopsis macdonaghi Orasema salebrosa Universal Chalcidoidea Database primary host
Solenopsis macdonaghi Orasema simplex Heraty et al., 1993; Varone et al., 2010; Baker et al., 2019
Solenopsis molesta Orasema coloradensis Wheeler, 1907; Johnson et al., 1986; Baker et al., 2019; Universal Chalcidoidea Database primary host
Solenopsis quinquecuspis Orasema aenea Universal Chalcidoidea Database primary host
Solenopsis quinquecuspis Orasema freychei Varone et al., 2010; Baker et al., 2019;
Solenopsis quinquecuspis Orasema simplex Heraty et al., 1993; Varone et al., 2010; Baker et al., 2019;
Solenopsis quinquecuspis Orasema xanthopus Silveira-Guido et al., 1964; Heraty et al., 1993; Varone et al., 2010; Baker et al., 2019 primary host
Solenopsis richteri Orasema salebrosa Heraty et al., 1993; Varone et al., 2010; Baker et al., 2019; Universal Chalcidoidea Database primary host
Solenopsis richteri Orasema simplex Heraty et al., 1993; Varone et al., 2010; Baker et al., 2019; Universal Chalcidoidea Database primary host
Solenopsis richteri Orasema xanthopus Silveira-Guido et al., 1964; Heraty et al., 1993; Varone et al. 2010; Baker et al., 2019; Universal Chalcidoidea Database primary host
Solenopsis saevissima Orasema xanthopus Universal Chalcidoidea Database associate, primary host
Solenopsis tenuis Orasema sixaolae Wheeler & Wheeler, 1937; Baker et al., 2019; Universal Chalcidoidea Database primary host
Solenopsis xyloni Orasema taii Heraty, 1990; Chien & Heraty, 2018; Baker et al., 2019; Universal Chalcidoidea Database primary host; ''Solenopsis geminata'' X ''xyloni'' hybrid
Temnothorax allardycei Orasema minuta Heraty, 1990; Baker et al., 2019; Universal Chalcidoidea Database primary host
Tetramorium Orasema "sp. 2 nr. bakeri" Heraty 1990; Baker et al., 2019 primary host
Wasmannia auropunctata Orasema minutissima Mann, 1918; Heraty, 1994; Wetterer & Porter, 2003; Burks et al., 2018; Baker et al., 2019; Universal Chalcidoidea Database primary host
Wasmannia sigmoidea Orasema minutissima Mann, 1918; Heraty, 1994; Wetterer & Porter, 2003; Burks et al., 2018; Baker et al., 2019; Universal Chalcidoidea Database primary host

References