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Annals of the Entomological Society of America
Published by: Entomological Society of America
Annals of the Entomological Society of America 93(3):374-390. 2000
doi: 10.1603/0013-8746(2000)093[0374:PROOHE]2.0.CO;2
Phylogenetic Relationships of Oraseminae (Hymenoptera: Eucharitidae)
Department of Entomology, University of California, Riverside, CA 92521
Abstract
Old World Oraseminae were revised recently and a phylogeny proposed; however, nothing was known at that time about the biology or immature stages for genera other than Orasema. The immature stages, habits, and host associations of Timioderus and Orasemorpha are herein described, and a new species of Orasema, O. monomoria, is described based on material reared from a species of Monomorium (Formicidae: Myrmicinae) in Madagascar. Using this new information, the relationships of Old World Oraseminae were reanalyzed. After successive weighting of 48 trees, a set of six trees (length 191, retention index 0.80) was found that matched the earlier hypothesis. To better evaluate behavioral traits of Oraseminae, nine species groups of Orasema from the New World were characterized and included in a separate analysis, resulting in 54 successively weighted trees (length 204 steps, retention index 0.81). Results suggest that a host shift to Pheidole (Myrmicinae) and internal parasitism of the host larva by the first-instar larva, with a later shift to external parasitism of the pupa, are derived for Oraseminae. All Oraseminae deposit eggs into punctures formed in the plant tissue by the ovipositor, but the habit of laying single eggs is plesiomorphic for Orasema and laying multiple eggs is derived for Orasemorpha and Timioderus. The use of an intermediate host for gaining access to the ant host appears to have been derived twice within Oraseminae.
Received: June 7, 1999
Keywords: cladistics, taxonomy, behavior, parasitoid, immatures
References
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Character matrix for adult features of O. monomoria and the New World species groups added to the matrix presented in Heraty (1994a). Numbers correspond to character states as listed in Heraty (1994a). Question marks denote missing values. Paired values in brackets are polymorphic within a species group.
Appendix 2. Character matrix for immature stages and behavioral characteristics.Species group names capitalized with abbreviations as outlined inHeraty (1994a). Numbers correspond to character stateslisted in Table 1. Question marks denote missing taxa. Paired values inbrackets are polymorphic within the genus or speciesgroup.
Figs. 1–7. O. monomoria n. sp. (1) Head and mesosoma in dorsal view. (2) Antenna. (3) Head and mesosoma. (4) Head. (5) Pupa, arrows point to nodules over petiole. (6) Hypopygium and ovipositor in lateral view. (7) Forewing. ss, Spiracular swelling
Figs. 8–11. O. valgius (Walker). (8) Unfed first-instar. (9) Exocarpus plant host, arrows point to oviposition punctures. (10) Second instar. (11) Third instar. (12–13) Orasema species (uichancoi group). (12) Pheidole host with imbedded first instar (enlarged). (13) Pupa, arrows point to nodules over petiole
Figs. 14–24. T. acuminatus Heraty (14, 17–24) and Timioderus peridentatus Waterston (15, 16). (14) Unfed first-instar (planidium): (a) habitus, dorsal (left) and ventral (right), (b) head in lateral view, (c) apical tergites in lateral view. (15) Oviposition puncture and distribution of planidia in plant spicule. (16) egg extracted from oviposition puncture. (17–18) Mature ant larva with embedded planidium. (19) Expanding planidium under shedding cuticle of ant pupa, margin of cuticle marked by arrows. (20) First-instar feeding on host pupa. (21) Second-instar feeding on host pupa. (22) Third-instar larva feeding on host pupa. (23) Third instar. (24) Pupa, arrows point to nodules over petiole
Figs. 25–26. (25) T. peridentatus Heraty, planidia on immature Tenothrips. (26) Timioderus sp. in colony of host, Pheidole capensis Mayr (photo courtesy of Hamish Roberston, South African Museum, Cape Town)
Figs. 27 -29. O. didentata Girault. (27) Unfed first-instar (planidium): habitus, dorsal (left) and ventral (right). (28) Egg with mature first instar. (29) Egg punctures. cr, Cercus; me, mature egg within scar; sc, oviposition scar; tp, tergopleural line
Fig. 30. Strict consensus cladogram for complete data set (characters 1–62) with only Old World species group of Orasema and based on successive approximations character weighting. Length = 191, 48 trees unweighted, six trees weighted, consistency index 0.52, retention index 0.80. Dark lines identify relationships supported on both unweighted and weighted trees. Dark bars represent synapomorphies; white bars reversals, and shaded bars homoplasies. Superscripts refer to multistate characters. Abbreviations after genera refer to species groups
Fig. 31. Strict consensus cladogram for Oraseminae, including representative New World species groups; pruned from the cladogram including all taxa as presented in Fig. 30. For the complete data set (weighted), length = 205, 53 trees, consistency index 0.48, retention index 0.81. Dark lines identify relationships supported on both unweighted and weighted trees. On the cladogram, dark bars represent unique synapomorphic changes, shaded bars indicate two transitions to use of an intermediate host using an ACCTRAN model of change. On the chart, black cells are positive host associations, and shaded are definite for no intermediate host association. Question marks for ant hosts indicate hosts have been associated but not verified; for the intermediate host record, it refers to an implied association with thrips on bananas by O. costaricensis.
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