Corresponding author: Jessica Awad (
Academic editor: Z. Lahey
Awad J, Bremer JS, Butterill PT, Moore MR, Talamas EJ (2021) A taxonomic treatment of
The previously described
The
The present work describes 20 species of
This work is based on specimens deposited in the following repositories with abbreviations used in the text:
Wasps were reared as a part of two community ecology projects focused on gall-forming insects in Papua New Guinea. Sampling occurred from 2010 to 2012 in the lowland rainforests of Madang (
Summary of
Host plant family | Host plant species | Number of specimens |
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23 | |
2 | ||
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1 | |
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1 | |
9 | ||
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22 | |
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2 | |
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2 | |
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2 | |
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4 | |
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14 | |
3 | ||
5 | ||
1 | ||
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22 | |
99 | ||
2 | ||
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12 | |
3 | ||
18 | ||
2 | ||
Total |
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Holotypes of new species are deposited in
Specimens were examined with Leica M205C, Zeiss Discovery V8, and Wild M5A microscopes. Images were produced with a Macropod imaging system with a Canon EOS 6D Mark II camera, EF 70–200mm lens, and 10× and 20× M Plan APO Mitutoyo objective lenses. Microphotography software included EOS 6D Mark II camera utility and Helicon Focus Pro 6.8.0 for image stacking. Scanning electron micrographs were produced with a Phenom XL Desktop SEM using a eucentric sample holder and Phenom ProSuite software. Specimens were coated with gold-palladium using a Denton Vacuum Desk V sputter coater. Adobe Photoshop was used for addition of scale bars and post processing of both brightfield images and electron micrographs.
DNA extraction was performed using a non-destructive protocol adapted for microhymenoptera (
CO1 gene sequences were aligned using the default settings of MUSCLE (
Aligned sequences were also evaluated in IQ-TREE 2.1.1 (
Character matrices of morphological data were constructed using the online program vSysLab (
Species of
Species | Year | Type locality | Holotype location | Host plant genus | Images of primary type (DOI) |
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1839 | Tasmania |
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1839 | Tasmania |
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1916 | Australia |
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1916 | Australia |
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1997 | Papua New Guinea |
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1997 | Papua New Guinea |
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1997 | Papua New Guinea |
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1997 | Papua New Guinea |
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1997 | Philippines |
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1997 | Philippines |
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1997 | Philippines |
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1997 | Philippines |
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1997 | Philippines |
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1997 | Philippines |
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1997 | Philippines |
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1997 | Philippines |
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2004 | Papua New Guinea |
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2004 | Australia |
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2004 | Australia |
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2008 | Indonesia |
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2008 | Indonesia |
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2008 | Indonesia |
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2008 | Indonesia |
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2008 | Indonesia |
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2008 | Indonesia |
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2008 | Indonesia |
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2008 | Indonesia |
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2008 | Indonesia |
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2008 | Indonesia |
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2009 | Indonesia |
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2009 | Indonesia |
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2009 | Indonesia |
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2013 | Papua New Guinea |
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2013 | Papua New Guinea |
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2013 | Papua New Guinea |
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2014 | Australia |
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2014 | Australia |
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2014 | Australia |
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2014 | Australia |
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2014 | Australia |
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2014 | Australia |
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2014 | Australia |
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All specimens of
Arrangement of clypeal setae in
Some species of
In the
Epomial carina in
The shape of the mesoscutellar spine has historically been used as a species-level diagnostic character for
Morphology of the mesoscutellar spine in
In
Margin of lateral propodeal carinae
The mesoscutal lamella is a posteromedial extension of the mesoscutum (Fig.
Mesoscutal lamella in
In most species, the presence of sculpture is absent or restricted to a narrow band on the posterior margins of S2 (Fig.
Sculpture of metasomal sternite 2
Sculpture of metasomal tergite 2
The setae on the forewing disc vary in density and arrangement (Fig.
Forewing of
Most
Among the
Generic characters for identification of
We here follow Jackson’s concept of
1 | T2 longer than the head and mesosoma combined (Fig. |
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– | T2 equal to or less than length of head and mesosoma combined (Fig. |
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2 | Notaulus distinctly grooved (Fig. |
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– | Notaulus unmarked or faintly suggested (Fig. |
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3 | Length of mesoscutellar spine only slightly greater than its thickness in lateral view; lateral pronotum with microsculpture dorsally, smooth ventrally, with smooth band anterior to tegula (Fig. |
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– | Length of mesoscutellar spine more than twice its thickness at its midpoint; lateral pronotum with microsculpture dorsally, smooth ventrally (Fig. |
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4 | Epomial carina absent or reduced, extending less than halfway up pronotum (Fig. |
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– | Epomial carina distinct and extending more than halfway up pronotum (Fig. |
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5 | Mesoscutellar spine absent or inconspicuous (Fig. |
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– | Mesoscutellar spine present, with variable form, often long or pointed (Fig. |
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6 | Head large and triangular in anterior view (Fig. |
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– | Head circular or ovoid in anterior view (Fig. |
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7 | Hyperoccipital carina absent or very weak (Fig. |
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– | Hyperoccipital carina present and complete (Fig. |
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8 | Frons with parallel, arched rugae above torulus; clypeal setae evenly spaced; lateral pronotum without microsculpture in ventral third (Fig. |
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– | Frons without parallel, arched rugae (but may have minute, irregular epiclypeal rugulae); median pair of clypeal setae closer to each other than to lateral setae; lateral pronotum with microsculpture throughout or with narrow smooth area at ventral apex (Fig. |
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9 | Mesoscutellar spine long and blunt (Fig. |
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– | Mesoscutellar spine short and pointed (Fig. |
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10 | Hyperoccipital carina weakly developed and interrupted (Fig. |
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– | Hyperoccipital carina robust and complete (Fig. |
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11 | Central keel well developed and extending to median ocellus (Fig. |
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– | Central keel absent, weakly developed, or not extending to median ocellus (Fig. |
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12 | Patch of microsculpture on posterior S2 extensive, longer than S3 (Fig. |
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– | S2 with no microsculpture or with very narrow band at posterior margin (Fig. |
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13 | T2 with extensive band of microsculpture at posterior margin that is longer than T3 (Fig. |
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– | T2 without microsculpture, or with narrow band at posterior margin (Fig. |
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14 | Parapsidal lines well impressed (Fig. |
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– | Parapsidal lines unmarked or faintly indicated (Fig. |
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15 | Distance between lateral ocellus and hyperoccipital carina at least 1.5 times ocellar diameter (Fig. |
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– | Distance between lateral ocellus and hyperoccipital carina less than 1.5 times ocellar diameter (Fig. |
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16 | Hyperoccipital carina robust and complete |
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– | Hyperoccipital carina weak or incomplete |
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17 | Mesoscutum evenly setose; mesoscutellum evenly setose (Fig. |
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– | Mesoscutum medially glabrous or with few scattered setae; mesoscutellum medially glabrous (Fig. |
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18 | Forewing disc proximally glabrous with linea setosa (Fig. |
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– | Forewing disc without linea setosa (Fig. |
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19 | Central keel impressed or partially present only in ventral part of frons (Fig. |
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– | Central keel absent (Fig. |
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Lateral habitus
Dorsal mesoscutum and mesoscutellum
Mesoscutellar spine
Lateral pronotum
Frontal head
Lateral and dorsal head
Frons and lateral pronotum
Lateral and dorsal head
Central keel
Dorsal mesosoma showing parapsidal lines
Head and lateral pronotum
Setation of dorsal mesoscutum and mesoscutellum
Setation of forewing disc including linea setosa
Head, mesoscutellum, and lateral propodeal carina
Body length 0.9–1.25 mm. Body color: black. Color of legs: coxae brown, otherwise yellow. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The central keel is variable, often present only in the ventral part of the frons, but sometimes complete or nearly so, extending to the median ocellus. The length of the forewing marginal setae is only slightly longer on the posterior margin than on the anterior margin.
This species is named in honor of the entomologist Amanda Hodges.
Reared from variously shaped galls on
Body length 1.3–1.9 mm. Body color: black. Color of legs: coxae brown, otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The head of this species is mostly covered in rugose macrosculpture and the notauli are distinctly grooved, characters which it shares with
The central keel is variable and difficult to discern due to the sculpture of the frons. The keel is present ventrally, and occasionally also present dorsally, just ventral to the median ocellus.
The epithet “
Reared from a vein swelling gall on
Body length 1.3 mm. Body color: black. Color of legs: coxae brown, otherwise yellow. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The lateral ocellus and the hyperoccipital carina are widely separated in
The female holotype and paratype of
Reared from galls on
Body length 1.1–1.5 mm. Body color: black; dark brown. Color of legs: coxae brown, otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The shape of the head in
The epithet “
The distance between the dorsal margin of the eye and the vertex is very small, as in
Reared from vein swelling, globoid, pimple, and blister galls on
Holotype:
Holotype of
Body length 1.7–2.3 mm. Body color: brown. Color of legs: coxae brown, legs otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The forewing disc is setose distally and glabrous proximally, with a broad linea setosa.
Unknown.
Body length 1.6 mm. Body color: black. Color of legs: coxae brown, otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
Important characters for identifying
The epithet “
Reared from a blister gall on
Body length 0.9 mm. Body color: dark brown. Color of legs: coxae brown, otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
Identification of
The epithet “
Reared from a pimple gall on
Body length 1.2–1.6 mm. Body color: black. Color of legs: coxae brown, otherwise yellow. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The arrangement of the clypeal setae is variable, with the median pair of setae either closer to each other than to the lateral setae, or with all four setae evenly spaced.
The epithet “
Reared from galls on
Body length 1.2–1.8 mm. Body color: black. Color of legs: coxae brown, otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The head of
The central keel is variable and difficult to discern due to the sculpture of the frons. The keel is present ventrally, and occasionally also present dorsally, just ventral to the median ocellus. Similarly, the hyperoccipital carina is often obscured by head sculpture, but when visible, it is weak and incomplete. The epomial carina terminates posteriorly in a pit.
The epithet “
Reared from vein swelling and globoid galls on
Body length 1.1–1.3 mm. Body color: black; dark brown. Color of legs: coxae brown, otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The central keel is variable in this species. It can be absent, or present ventrally within a shallow longitudinal depression.
The epithet “
Reared from galls on
Body length 1.1–1.5mm. Body color: black. Color of legs: coxae brown, otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The epithet “
Reared from nodule-like galls on
Body length 1.5–1.7 mm. Body color: black. Color of legs: coxae brown, otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The central keel may be ventrally present or entirely absent. The forewing disc is proximally glabrous or with a few scattered setae.
This species is named in honor of botanist Patti J. Anderson.
Reared from globoid galls on
Body length 1.3–1.4 mm. Body color: dark brown. Color of legs: coxae brown, otherwise yellow. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The parapsidal lines of
The female types of
Reared from galls on
Holotype:
Body length 1.4–1.5 mm. Body color: black. Color of legs: coxae brown, otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The frons of
Reared from galls on
Holotype of
Body length 1.6–1.8 mm. Body color: black. Color of legs: coxae brown, otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The epithet “
Reared from galls on
Body length 1.2–1.5 mm. Body color: black. Color of legs: coxae brown, otherwise yellow; coxae brown, otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
This species is named in honor of the biological control researcher and beetle taxonomist Ronald D. Cave.
Primarily reared from blister galls on
Body length 1.1–1.4mm. Body color: black. Color of legs: coxae brown, otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The epithet “
Reared from blister galls on
Body length 1.1–1.5 mm. Body color: black. Color of legs: coxae brown, otherwise yellow. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
The epithet “
Reared from galls on
Body length 1.3–1.5 mm. Body color: black. Color of legs: coxae brown, otherwise yellow. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
A similar head shape is found in
The epithet “
Reared from round, fuzzy galls on
Body length 1.1–1.3 mm. Body color: black. Color of legs: coxae brown, otherwise yellow to brown. Color of mesoscutellar spine: concolorous with mesoscutellar disc.
This species is named in honor of the hymenopterist Yuanmeng Miles Zhang.
Reared from galls on
Neighbor-joining and maximum likelihood analyses recovered cohesive clusters of morphospecies, within which all individuals were more similar to each other than to any other species (Figs
The New Guinea material did not all cluster together.
The CO1 sequences of three species,
We recognize the possibility that these species are widespread and may have already been described. However,
Some morphological characters formed discernible patterns. The cluster including
Other small clusters have more subtle morphological similarities.
Neighbor-joining tree of
Maximum likelihood tree of
Of the 19
For example, the cluster of five gall types formed on
The number of reared parasitoids is directly related to the number of host galls reared, and the
More data are needed to confirm host associations and, to that end, further work is underway in order to establish the relationships among the host cecidomyiid gall formers (Kolesik and Butterill, unpublished data). Future workers are encouraged to acquire specimens through rearing of host material, due to the value of ecological data in understanding
We thank the technical staff of the Florida State Collection of Arthropods and molecular diagnostics laboratory at the Florida Department of Agriculture and Consumer Services, Division of Plant Industry (FDACS-DPI) for their assistance with this project. Specimen loans were provided by Lars Vilhelmsen (
List of specimens and their depositories
material examined
List of specimens examined, depositories, type status, and image links. Link to holotype image repository:
CO1 data in Genbank
COI data
List of barcode vouchers, internal identifiers, and Genbank accession numbers. Link to PNG image repository: