Research Article |
Corresponding author: Ryan D. Ridenbaugh ( r.ridenbaugh@knights.ucf.edu ) Academic editor: Jose Fernandez-Triana
© 2018 Ryan D. Ridenbaugh, Erin Barbeau, Barbara J. Sharanowski.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Ridenbaugh RD, Barbeau E, Sharanowski BJ (2018) Description of four new species of Eadya (Hymenoptera, Braconidae), parasitoids of the Eucalyptus Tortoise Beetle (Paropsis charybdis) and other Eucalyptus defoliating leaf beetles. Journal of Hymenoptera Research 64: 141-175. https://doi.org/10.3897/jhr.64.24282
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Eucalyptus L’Héritier, 1789 (Myrtales: Myrtaceae) plantations are a global economic resource with a wide array of uses. As this forestry crop grows in popularity around the world, the exotic introduction of pests such as the leaf beetles belonging to the genera Paropsis Oliver, 1807 and Paropsisterna Motschulsky, 1860 increases in frequency. These pest introductions have spurred a need to understand the natural enemies of these pests for use in classical biological control programs. One such enemy, Eadya paropsidis Huddleston & Short, 1978 (Hymenoptera: Braconidae), has shown potential as a biological control agent against Paropsis charybdis, an exotic pest of New Zealand Eucalyptus plantations. However, observations made by biocontrol researchers have raised concerns that E. paropsidis is a complex of cryptic species. A comprehensive large-scale phylogenetic study utilizing both host and molecular data (
Parasitoid wasps, Taxonomy, DNA Barcoding, Morphometrics, Biological Control, Tasmania, New South Wales, Victoria, Euphorinae , Eucalyptus
Although native to Australia, the cultivation of production and trade of goods derived from Eucalyptus L’Héritier, 1789 (Myrtales: Myrtaceae) is a massive global industry. The largest subdivision of this industry is the Eucalyptus oil market (
Species of Paropsis Oliver, 1807 and Paropsisterna Motschulsky, 1860 are endemic Australian leaf-beetles (Coleoptera: Chrysomelidae: Chrysomelinae) that feed upon the leaves and shoots of Eucalyptus. These beetles have been known to cause serious damage to Eucalyptus plantations both within (
Larval endoparasitic wasps in the genus Eadya Huddleston & Short, 1978 (Hymenoptera: Braconidae) have great potential as biocontrol agents for invasive paropsines. Classical biological control studies have begun for Eadya from Tasmania to control the Eucalyptus Tortoise Beetle, Paropsis charybdis Stål, 1860 (
In this paper, we formally describe these three new species discovered from
We utilized material collected from
A molecular diagnostic key was created using the barcoding region (
Photographs were taken using a Canon 7D Mark II with the following lenses: MP-E 65mm 1–5× Canon macro lens, and a M Plan Apo 10× Mitutoyo objective mounted onto the EF Telephoto 70–200mm Canon zoom lens. For lighting, the MT-24EX Macro Twin Lite Canon Flash was used in conjunction with a custom made diffuser. Multiple images were taken of each specimen and compiled into a single image using Zerene Stacker 1.04 (Zerene Systems LLC.). Scale bars were added using ImageJ 1.51 (
Of the four species supported by the molecular data presented in
The characters evaluated in this study were as follows: Lateral ocellar line (LOL), ocular ocellar line (OOL), posterior ocellar line (POL), occipital ocellar line (oci.l), genal space (gsp.l), malar space (mlr.l), head breadth (hea.b), and metasomal tergite 1 breadth (mt1.b). The definition of these characters and how they were measured can be found in Table
The host, Paropsisterna variicollis* (Chapuis, 1877) is listed with an asterisk within descriptions due to the uncertainty surrounding the taxonomic validity of this species with respect to Pst. obovata (Chapuis, 1877) and Pst. cloelia (Stål, 1860). For a detailed discussion on the taxonomic uncertainty of this species, see
Abbreviations and definitions of the 8 morphological characters used for the morphometrics analysis of Eadya paropsidis and Eadya daenerys.
Abbreviation | Character name | Definition | Magnification (E. paropsidis) | Magnification (E. daenerys) |
---|---|---|---|---|
LOL | Lateral Ocellar Line | The shortest distance between the median and lateral ocellus, dorsal view (Fig. |
100× | 100× |
OOL | Ocular Ocellar Line | The shortest distance between the lateral ocellus and the eye, dorsal view (Fig. |
100× | 100× |
POL | Posterior Ocellar Line | The shortest distance between the lateral ocelli, dorsal view (Fig. |
100x | 100× |
oci.l | Occipital Ocellar Line | The shortest distance from the posterior edge of the lateral ocellus at a 90° angle to the occipital carinae, dorsal view (Fig. |
100× | 100× |
gsp.l | Genal Space | Length of the genal space taken midway between the dorsal and ventral margins of the eye from the posterior edge at a 90° angle to the occipital carinae, lateral view (see Fig. |
100× | 100× |
mlr.l | Malar Space | Length of the malar space taken from the posterior margin of the eye to the base of the mandible, anterior view (Fig. |
100× | 100× |
hea.b | Head breadth | Greatest breadth of head, dorsal view (see Fig. |
50× | 50× |
mt1.b | Metasomal tergite 1 breadth | Greatest breadth of metasomal tergite 1 at the posterior margin, dorsal view (see Fig. |
50× | 100× |
Separating most species of Eadya was relatively straightforward using morphological characters (see Key to Species of Eadya below). However, E. paropsidis and E. daenerys sp. n. presented only size differences morphologically, with the latter species being smaller, even though they were well supported phylogenetic species based on molecular data (
A PCA and allometry ratio spectrum were generated to determine which characters were the best for delimiting the two cryptic species. The most discerning ratios according to the first principal component were LOL:mlr.l, LOL:mt1.b, and LOL:gsp.l (Fig.
Multivariate morphometric ratio analysis of female specimens of Eadya paropsidis, and Eadya daenerys Ridenbaugh, sp. n. A Scatterplot of the first shape principal component plotted against the second shape principal component. Black - Eadya paropsidis, Green - Eadya daenerys sp. n. B Scatterplot of isosize plotted against the first shape principal component. Black - Eadya paropsidis, Green - Eadya daenerys sp. n. C Ratio spectrum for the first principal component with horizontal bars representing 68% confidence based on 1000 bootstrap replicates D Allometry ratio spectrum with horizontal bars representing 68% confidence based on 1000 bootstrap replicates.
When applied to E. paropsidis and E. daenerys sp. n., the morphometrical analysis only supported one species, contrasting with the results of
Eadya can be recognized from other braconid genera by the following combination of characteristics: head large, subcubic and as wide as thorax, clypeus flat, labrum flat, interantennal carina present; forewing with r-m crossvein present, 3RSb curved and meeting R1a before apex of wing, and 2cu-a absent; metasoma petiolate.
Eadya annleckieae sp. n. can be distinguished from all other members of Eadya by the following combination of characters: Clypeus flanged across ventral margin, without medial tubercles (Fig.
Female. Body length 6.46mm. Ovipositor length 1.72mm.
Color. Head black except for mandible orange with base black and apex ferruginous, maxillary and labial palp orange, and antenna dark brown (Figs
Head. Clypeus simple, punctate and pubescent, flanged across ventral margin, without medial tubercles (Fig.
Mesosoma. Pronotum exposed in dorsal view, pronope and subpronobe present, covered in rugulose sculpturing (Fig.
Forewing. r-m sinuous (Fig.
Hindwing. R1a with three hamuli.
Metasoma. Metasomal tergite 1 petiolate, spiracle protruding as a tubercle at about the middle of the segment, dorsal surface smooth, lateral surface punctate with associated setae; ovipositor straight (Fig.
Male. Same as female.
Paropsisterna nobilitata (Erichson, 1842), Paropsisterna variicollis* , Paropsisterna selmani Reid & de Little, 2013, Paropsis charybdis.
Paratype with propleuron black except for lateral posterior margin orange; mesoscutum orange except for the median mesonotal lobe black with the anterior margin and lateral mesonotal lobes ferrunginous (Fig.
Amino acid positions (22–27) MWAGIL; (32–34) SII; (41–46) SRGSLL; (54) R; (67–73) MVMPVIM; (81) I; (90) I; (95–98) MNNM; (104–109) LPSLFI; (112–118) LRRLTNI; (126) I; (133–139) GGRHSGV; (143–144) VA; (150) I; (157) [I or K]; (167–169) FNM; (172–191) NGIAVDRVTLFRWSVKITAF (Fig.
Tasmania.
This species is named in honor of the science fiction author, Ann Leckie by the second author (EB).
This species is referred to as Eadya sp.1 in
Holotype, Female (
Eadya daenerys sp. n. can be distinguished from all other members of Eadya by the following combination of characters: Clypeus flanged along ventral margin, with two medial tubercles projecting outward (Fig.
Female. Body Length 5.77mm. Ovipositor Length 0.82mm.
Color. Head orange except for antenna, apex of mandible, and ocellar triangle black (Figs
Head. Clypeus simple, smooth with scattered setae, flanged at ventral margin, with two medial tubercles projecting outward (Fig.
Mesosoma. Pronotum exposed in dorsal view, pronope and subpronope absent, smooth except for a crenulate line extending laterally and rugulose sculpturing along the lateral posterior margin (Fig.
Forewing. r-m curved slightly towards stigma before reaching the junction of 3RSa and 3RSb (Fig.
Hindwing. R1a with three hamuli.
Metasoma. Metasomal tergite 1 petiolate, spiracle protruding as a tubercle at about the middle of the segment, dorsal and lateral surface punctate with associated setae (Fig.
Male. Same as female.
Eadya daenerys Ridenbaugh, sp. n. paratype. A Head, frontal view B Head, dorsal view, arrow indicating simple occipital carinae C Head and mesoscutum, dorsal view, paratype D Mesopleuron, lateral view, paratype E Propodeum, dorsal view F Propodeum, posterio-dorsal view. All scale bars are 1mm in length.
Paropsisterna agricola (Chapuis, 1877), Paropsisterna bimaculata (Olivier, 1807), Paropsisterna nobilitata, Paropsis charybdis.
Amino acid positions (22–27) [M or R]WSGII; (32–34) RVL; (41–46) ILGRLL; (54) S; (67–73) IVIPIII; (81) I; (90) I; (95–98) INNI; (104–109) PPSL[I or V]L; (112–118) IRNFIGA; (126) I; (133–139) NLSHRGV; (143–144) [V or I]S; (150) L; (157) I; (167–169) INI; (172–191) LGLSYDNISLLVWSVNITAI (Fig.
Australian Capital Territory, New South Wales, Tasmania.
This species is named for Daenerys Stormborn of House Targaryen, the First of Her Name, Queen of the Andals and the First Men, Protector of the Seven Kingdoms, the Mother of Dragons, Khaleesi of the Great Grass Sea, the Unburnt, the Breaker of Chains, from the literary series A Song of Ice and Fire by George R.R. Martin, as well as the television series Game of Thrones on Home Box Office (HBO). This is a noun in apposition to the generic name in order to retain integrity of the fictional character name Daenerys.
This species is referred to as Eadya sp.3 in
Holotype, Female (
See Suppl. material
Eadya duncan sp. n. can be distinguished from all other members of Eadya by the following combination of characters: Clypeus flanged at ventral margin, with two medial tubercles projecting outward (Fig.
Male. Body length 6.37mm.
Color. Head orange except for antenna, apex of mandible, and ocellar triangle black (Figs
Head. Clypeus simple, smooth with scattered setae, flanged at ventral margin, with two medial tubercles projecting outward (Fig.
Mesosoma. Pronotum exposed in dorsal view, pronope and subpronope absent, smooth except for a faint crenulate line extending laterally and rugulose sculpturing along the lateral posterior margin (Fig.
Forewing. r-m curved slightly towards stigma before reaching the junction of 3RSa and 3RSb (Fig.
Hindwing. R1a with three hamuli.
Metasoma. Metasomal tergite 1 petiolate, spiracle protruding as a tubercle at about the middle of the segment, dorsal and lateral surface punctate with associated setae (Fig.
Female. Unknown.
Unknown.
New South Wales, Victoria (see discussion).
This epithet is named in honor of the senior author’s (BJS) sister in law, Julie Brant nee Duncan, who is an Australian-born beauty. This is a noun in apposition to the generic name in order to retain integrity of the surname Duncan.
The holotype for this species was identified as a species of Eadya by Huddleston in 1977 and deposited at
Holotype, Male (
Eadya falcata can be distinguished from all other members of Eadya by the following combination of characters: Clypeus flanged at ventral margin, with two medial tubercles projecting outward (Fig.
Female. Body Length 5.26mm. Ovipositor Length 1.80mm.
Color. Head orange except for antenna, apex of mandible, and ocellar triangle black; pronotum orange expect for lateral posterior margins black (Fig.
Head. Clypeus simple, smooth with scattered setae, flanged at ventral margin, with two medial tubercles projecting outward (Fig.
Mesosoma. Pronotum exposed in dorsal view, pronope and subpronope absent, smooth (Fig.
Forewing. r-m curved slightly towards stigma before reaching the junction of 3Rsa and 3RSb.
Hindwing. R1a with three hamuli.
Metasoma. Metasomal tergite 1 petiolate, spiracle protruding as a tubercle at about the middle of the segment, dorsal and lateral surface punctate with associated setae; ovipositor curved downward (Fig.
Male. Same as female.
Unknown.
Paratype with foreleg coxa orange and trochanter, trochantellus, femur, tibia, and tarsus black.
Western Australia.
The crenulation at the apex of the notaulus is difficult to see in the holotype due to damage caused by pinning (Fig.
Holotype, Female (
Eadya paropsidis can be distinguished from all other members of Eadya by the following combination of characters: Clypeus flanged at ventral margin, with two medial tubercles projecting outward (Fig.
Female. Body Length 6.29mm. Ovipositor Length 1.08mm.
Color. Head orange except for antenna, apex of mandible, and ocellar triangle black (Figs
Head. Clypeus simple, smooth with scattered setae, flanged at ventral margin, with two medial tubercles projecting outward (Fig.
Mesosoma. Pronotum exposed in dorsal view, pronope and subpronope absent, smooth except for a faint crenulate line extending laterally and rugulose sculpturing along the lateral posterior margin (Fig.
Forewing. r-m curved slightly towards stigma before reaching the junction of 3RSa and 3RSb (Fig.
Hindwing. R1a with three hamuli.
Metasoma. Metasomal tergite 1 petiolate, spiracle protruding as a tubercle at about the middle of the segment, dorsal and lateral surface punctate with associated setae (Fig.
Male. Same as female.
Eadya paropsidis. A Head, frontal view B Head, dorsal view, arrow pointing to emarginate occipital carinae C Head and mesoscutum, dorsal view D Mesopleuron, lateral view E Propodeum, dorsal view F Propodeum, dorsal view, with arrows indicating transverse carinae. All scale bars are 1mm in length.
Paropsis atomaria Olivier, 1807, Paropsis tasmanica Baly, 1866, Paropsis charybdis.
(22–27) MWSGII; (32–34) SVL; (41–46) ILGRLI; (54) S; (67–73) IVIPIII; (81) V; (90) M; (95–98) INNI; (104–109) PPSLIL; (112–118) TRNFIGI; (126) I; (133–139) NLRHRGI; (143–144) IS; (150) L; (157) M; (167–169) INI; (172–191) LGLNYDNISLLVWSVNITAI (Fig.
Australian Capital Territory, Victoria, New South Wales, Tasmania.
Holotype, Female (
See Suppl. material
Eadya spitzer sp. n. can be distinguished from all other members of Eadya by the following combination of characters: Clypeus flanged at ventral margin, with two medial tubercles projecting outward (Fig.
Female. Body length without abdomen 3.30mm. Abdomen 2.86mm. Ovipositor 1.17mm.
Color. Head orange except for antenna, apex of mandible, and ocellar triangle black (Figs
Head. Clypeus simple, smooth with scattered setae, flanged as ventral margin, with two medial tubercles projecting outward (Fig.
Mesosoma. Pronotum exposed in dorsal view (Fig.
Forewing. r-m curved slightly towards stigma before reaching the junction of 3RSa and 3RSb (Fig.
Hindwing. R1a with three hamuli.
Metasoma. Metasomal tergite 1 petiolate, spiracle protruding as a tubercle at about the middle of the segment, dorsal and lateral surface punctate with associated setae (Fig.
Male. Unknown.
Paropsis charybdis, Paropsis aegrota elliotti Selman, 1983.
Paratype with clypeus orange (Fig.
(22–27) IWSGII; (32–34) SVL; (41–46) [M or K]LGRLL; (54) S; (67–73) IVIPIII; (81) I; (90) MM; (95–98) INNI; (104–109) PPSLIL; (112–118) IRNFIGM; (126) M; (133–139) NLRHRGI; (143–144) MS; (150) L; (157) I; (167–169) INI; (172–191) LGLNYDNISLLVWSVNITAI (Fig.
Tasmania.
This species is named in honor of Edwin Spitzer, the first author’s (RDR) late grandfather. This is a noun in apposition to the generic name in order to retain integrity of the surname Spitzer.
The paratype is for this series is badly damaged, missing both antennae, all six legs, and the abdomen excluding metasomal tergite 1. However, the specimen was photographed before destruction and can be seen in Figures
Holotype, Female (
Characters used in the morphometric analysis. A Frontal view of the head illustrating the morphometric character malar space (mlr.l), Eadya annleckieae Ridenbaugh, sp. n. paratype B Dorsal view of the head illustrating the morphometric characters lateral ocellar line (LOL), ocular ocellar line (OOL), posterior ocellar line (POL), and occipital ocellar line (oci.l), Eadya annleckieae Ridenbaugh, sp. n. paratype. All scale bars are 1mm in length.
1 | Propodeum with transverse carinae (See arrows, Fig. |
2 |
– | Propodeum without transverse carinae (Fig. |
3 |
2 | Occipital carinae simple (See arrow, Fig. |
E. duncan Ridenbaugh, sp. n. |
– | Occipital carinae emarginate (See arrow, Fig. |
E. paropsidis Huddleston & Short, 1978 |
3 | Notaulus impressed towards anterior margin of mesoscutum, crenulate at apex (Fig. |
E. falcata Huddleston & Short, 1978 |
– | Notaulus rugulose (Fig. |
4 |
4 | Head black (Fig. |
E. annleckieae Ridenbaugh, sp. n. |
– | Head orange except for antenna, apex of mandible, and ocellar triangle black (Fig. |
5 |
5 | Pronotum orange (Fig. |
E. spitzer Ridenbaugh, sp. n. |
– | Pronotum black except for anterior dorsal margin orange (Figs |
E. daenerys Ridenbaugh, sp. n. |
With the description of the four new species described here, the distribution of Eadya has expanded to include Tasmania, the Australian Capital Territory, New South Wales, Victoria, and Western Australia. As
Interestingly, knowledge on Eadya distribution has grown from a citizen science observation. Citizen science initiatives are a valuable, yet underutilized, resource for biodiversity research which can survey large geographical areas over extended periods of time (
“A small (7mm) wasp with an orange head, thorax and first pair of legs. The rest of the wasp was black. The larvae being parasitized were those of the eucalyptus leaf beetle (Paropsis atomaria), probably the 2nd instar”.
Although the image quality and detail was not sufficient to positively identify the beetle larvae, the images of the wasp coupled with the contributor’s description matches that of E. duncan sp. n., and represents a new distribution record. With this observation, the distribution of E. duncan sp. n. is expanded to include Victoria, AUS in addition to New South Wales, AUS. Thus, citizen science observations can be invaluable for expanding knowledge on species and provides additional collecting localities for future research into this relatively unknown species.
Host records for Eadya outside of Tasmania are incomplete as well, with only E. paropsidis recorded from Paropsis atomaria (synonym P. reticulata) in the Australian Capital Territory and New South Wales (
Much is still unknown about the species of Eadya, but as the popularity of Eucalyptus grows internationally as an ornamental landscape and forestry product (
Finally, it is prudent to discuss the subfamily placement of Eadya. In the original description,
Three new species from the genus Eadya are described (Eadya annleckieae Ridenbaugh, sp. n., Eadya daenerys Ridenbaugh, sp. n., Eadya spitzer Ridenbaugh, sp. n.) based upon the results of
We would like to thank Geoff Allen, Anthony Rice, and Dean Satchell for collecting the material used in the description, Toni Withers and her team at Scion for their feedback on the key to the species of Eadya, and Joshua Hogan for his assistance with photographing specimens. Further we gratefully thank the Hymenopteran team at
Table S1