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Research Article
A taxonomic treatment of Synopeas Förster (Platygastridae, Platygastrinae) from the island of New Guinea
expand article infoJessica Awad, Jonathan S. Bremer§, Philip T. Butterill|, Matthew R. Moore§, Elijah J. Talamas§
‡ State Museum of Natural History Stuttgart, Stuttgart, Germany
§ FDACS-DPI, Gainesville, United States of America
| University of South Bohemia, České Budějovice, Czech Republic
¶ New Guinea Binatang Research Centre, Madang, Papua New Guinea
Open Access

Abstract

Synopeas from New Guinea is revised, including 16 new species and four previously named species. The concepts for these species were developed in conjunction with a CO1 analysis that includes 16 New Guinean species and 3 Synopeas species from other regions. The molecular analysis determined that the New Guinea fauna does not form a clade, indicating multiple migrations. The following species are described and keyed: S. amandae Awad, sp. nov.; S. anunu Awad, sp. nov.; S. butterilli Buhl; S. codex Awad, sp. nov.; S. csoszi Buhl; S. kalubia Awad, sp. nov.; S. kiki Awad, sp. nov.; S. kira Awad, sp. nov.; S. klingunculum Awad, sp. nov.; S. luli Awad, sp. nov.; S. occultum Awad, sp. nov.; S. pattiae Awad, sp. nov.; S. psychotriae Buhl; S. pterocarpi Buhl; S. pulupulu Awad, sp. nov.; S. roncavei Awad, sp. nov.; S. sanga Awad, sp. nov.; S. toto Awad, sp. nov.; S. valavala Awad, sp. nov.; S. zhangi Awad, sp. nov. Leptacis pleuralis (Buhl), comb. nov. is transferred from Synopeas. Images of 56 holotypes of Synopeas are made publicly available online.

Keywords

Cecidomyiidae, CO1 barcoding, galls, Papua New Guinea, parasitoid wasps, Platygastroidea, taxonomy

Introduction

Synopeas Förster, 1856, is one of the largest genera in Platygastridae (sensu Masner and Huggert 1989; Talamas et al. 2019) with 378 described species (Hymenoptera Online 2019). It is found worldwide and is commonly collected in all habitats where platygastrids are present. However, Synopeas has never been revised and the existing keys to species are few in number and limited in scope. Past taxonomic work was often based upon simple measurements of one or few specimens, and most descriptions lack thorough illustrations. Thus, there is a need for imaging and redescription of type specimens for the worldwide fauna. Additionally, molecular data is unavailable for most Synopeas species. The BOLD database has records for only four identified species of Synopeas. Ecological data is also rare, as most species have been described from adult specimens caught by net or trap and host or plant associations are available for only 42 species. Species of Synopeas, like other Platygastrinae, are known to be obligate primary endoparasitoids of gall midges (Diptera: Cecidomyiidae). Female wasps oviposit in eggs or early first instars of the host. The parasitoid development is delayed until later host instars, at which point it proceeds rapidly (Abram et al. 2012).

The previously described Synopeas species of Papua New Guinea can be traced to three collecting events: the voyage of the Noona Dan in 1961–1962 (Wolff 1963; Buhl 1997); a visit to Mt. Wilhelm by the Hungarian acarologist János Balogh in 1968 (Dózsa-Farkas 2003; Buhl 2004); and the extensive rearing work of Philip Butterill in 2010–2012. Of these, only Balogh and Butterill sampled the island of New Guinea.

The Noona Dan expedition was sponsored by the University of Copenhagen with a mission to collect natural history specimens and data from several Indo-Australian island groups. The Papuan Synopeas specimens described from the voyage were collected from the Bismarck Archipelago, off the island of New Britain. The Noona Dan expedition also collected Synopeas specimens from the Philippines and the Solomon Islands, which we examined and photographed.

Synopeas reared from galls collected in Papua New Guinea provided the basis for our study, which seeks to elevate the standard of taxonomy for Platygastridae through the integration of morphological, molecular, and ecological data. We examined primary types of Synopeas described from the Philippines, Australia, New Zealand, Papua New Guinea, and Indonesia to be as thorough as possible and to avoid redescription of species that may be widespread. Images of these types are made publicly available to facilitate future work in the Australasian fauna of the genus.

The present work describes 20 species of Synopeas from the island of New Guinea. These include 16 new species of Synopeas from the Butterill collection and four previously described species from the Butterill and Balogh collections: S. butterilli Buhl, S. csoszi Buhl, S. psychotriae Buhl, and S. pterocarpi Buhl. An illustrated key to the Synopeas species of New Guinea is provided.

Materials and methods

Collections

This work is based on specimens deposited in the following repositories with abbreviations used in the text:

FSCA Florida State Collection of Arthropods, Gainesville, USA

HNHM Hungarian National History Museum, Budapest, Hungary

NHMUK Natural History Museum, London, UK

NMINH National Museum of Ireland – Natural History, Dublin, Ireland

RMNH Naturalis Biodiversity Center, Leiden, Netherlands

SAMA South Australian Museum, Adelaide, Australia

SMNS State Museum of Natural History, Stuttgart, Germany

USNM National Museum of Natural History, Smithsonian Institution, Washington, DC, USA

ZMUC Natural History Museum of Denmark, Copenhagen, Denmark

Specimens

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 (Butterill and Novotny 2015) and in the montane rainforests of the Finisterre Mountains, Morobe Province (Butterill, unpublished data). All parasitoid specimens were reared from galls formed by Cecidomyiidae and stored in 95% ethanol. In total, we examined 249 Synopeas specimens (Table 1) reared from 21 plant species in 14 genera.

Table 1.

Summary of Synopeas specimens by gall host plant identity.

Host plant family Host plant species Number of specimens
Actinidiaceae Saurauia conferta Warb. 23
Saurauia schumanniana Diels 2
Elaeocarpaceae Elaeocarpus dolichostylus Schltr. 1
Euphorbiaceae Homalanthus nervosus J.J.Sm. 1
Macaranga strigosa Pax & K.Hoffm. 9
Loganiaceae Neuburgia corynocarpa (A.Gray) Leenh. 22
Monimiaceae Palmeria sp. 2
Moraceae Ficus adenosperma Miq. 2
Myristicaceae Paramyristica cf. sepicana (Foreman) W.J. de Wilde 2
Myrtaceae Syzygium decipiens (Koord. and Valeton) Merr. and L.M.Perry 4
Piperaceae Piper amboinense (Miq.) C.DC. 14
Piper celtidiforme Opiz 3
Piper macropiper Pennant 5
Piper sp. 1
Rubiaceae Nauclea tenuiflora (Havil.) Merr. 22
Nauclea sp. 1 99
Psychotria ramuensis Sohmer 2
Urticaceae Cypholophus friesianus (K.Schum.) H.J.P.Winkl. 12
Cypholophus macrocephalus (Blume) Wedd. 3
Cypholophus sp. 1 18
Debregeasia longifolia (Burm.f.) Wedd. 2
Total 14 genera 249

Holotypes of new species are deposited in USNM. Paratypes for species described here are deposited in USNM, FSCA, and SMNS (details in Suppl. material 1).

Morphological techniques

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.

Molecular techniques

DNA extraction was performed using a non-destructive protocol adapted for microhymenoptera (Sabbatini Peverieri et al. 2018). DNA extracts were quantified with a NanoDrop 2000 spectrophotometer (Thermo Fisher Scientific, Waltham, MA) and PCRs had a target input of at least 20 ng of genomic DNA. PCRs were performed using a KAPA HiFi HotStart PCR Kit (Kapa Biosystems, Wilmington, MA). PCRs were 25 µL in total volume. The standard 5’-CO1 barcode region was targeted for each species using the primer pairs LCO1490/HCO2198 (Folmer et al. 1994) or LEPF1/LEPR1 (Hebert et al. 2004). Thermocycles were as follows: 1) initial denaturing at 95 °C for 2 min; 2) 98 °C for 30 sec; 3) 50 °C for 30 sec; 4) 72 °C for 40 sec [32 cycles of steps 2–4]; and 5) a final extension at 72 °C for 7 min. PCR amplicons were visualized on 1.5% agarose gels. Positive PCR amplicons were purified using the QIAGEN QIAquick PCR Purification Kit (QIAGEN Group, Hilden, Germany) and prepared for sequencing with BigDye Terminator v3.1 chemistry. Sequence chromatograms were trimmed and assembled in Sequencher 5.4.6 (2016). New CO1 barcodes were submitted to Genbank (Suppl. material 2).

CO1 gene sequences were aligned using the default settings of MUSCLE (Edgar 2004) as implemented in MEGA7 (Kumar et al. 2016). Specimens of Inostemma Haliday and Platygaster Latreille were selected as outgroups based on an unpublished phylogeny of the family. For geographic reference, the analysis included Synopeas specimens from Myanmar, the United Kingdom, and the United States. The resulting alignment included 75 terminal taxa (70 Synopeas) and was 655 bp in length. Neighbor-joining trees were constructed using the K2P model (Kimura 1980) with partial deletion of missing data and a site coverage cutoff of 95%. Node support was assessed using 10,000 bootstrap replicates. The resulting tree topology was manipulated in FigTree v1.4.3 (Rambaut 2012) to aesthetically arrange nodes and collapse terminal clusters. The tree was further edited in Adobe Illustrator. CO1 barcodes were queried to the Barcode of Life Database (BOLD) (Ratnasingham and Hebert 2007) in search of high-similarity matches.

Aligned sequences were also evaluated in IQ-TREE 2.1.1 (Nguyen et al. 2015). ModelFinder (Kalyaanamoorthy et al. 2017) was used to select the best substitution model, which was TIM+F+I+G4 based on the Bayesian Information Criterion. Maximum likelihood tree reconstruction was inferred based on 1000 ultrafast bootstrap replicates (Minh et al. 2013). The resulting tree was edited in Adobe Illustrator.

Descriptions

Character matrices of morphological data were constructed using the online program vSysLab (https://vsyslab.osu.edu/). Names were assigned to specimens by comparison to holotypes of species from Australasia, Indonesia, and the Philippines (Table 2). If no matching types were found, species were described as new.

Table 2.

Species of Synopeas described from Australasia, Indonesia, and the Philippines.

Species Year Type locality Holotype location Host plant genus Images of primary type (DOI)
S. isus (Walker) 1839 Tasmania NHMUK
S. leda (Walker) 1839 Tasmania NHMUK
S. idarniforme (Dodd) 1916 Australia SAMA https://doi.org/10.5281/zenodo.4503382
S. saccharale (Dodd) 1916 Australia SAMA Saccharum https://doi.org/10.5281/zenodo.4503406
S. decumbens Buhl 1997 Papua New Guinea ZMUC https://doi.org/10.5281/zenodo.4501922
S. lemkaminensis Buhl 1997 Papua New Guinea ZMUC https://doi.org/10.5281/zenodo.4501952
S. saintexuperyi Buhl 1997 Papua New Guinea ZMUC https://doi.org/10.5281/zenodo.4501968
S. ventricosum Buhl 1997 Papua New Guinea ZMUC https://doi.org/10.5281/zenodo.4501974
S. acutiventris Buhl 1997 Philippines ZMUC https://doi.org/10.5281/zenodo.4503063
S. balabacensis Buhl 1997 Philippines ZMUC https://doi.org/10.5281/zenodo.4503179
S. crassiceps Buhl 1997 Philippines ZMUC https://doi.org/10.5281/zenodo.4503402
S. luteolipes Buhl 1997 Philippines ZMUC https://doi.org/10.5281/zenodo.4503726
S. montanum Buhl 1997 Philippines ZMUC https://doi.org/10.5281/zenodo.4503804
S. mukerjeei Buhl 1997 Philippines ZMUC https://doi.org/10.5281/zenodo.4503979
S. palawanense Buhl 1997 Philippines ZMUC https://doi.org/10.5281/zenodo.4504012
S. pallescens Buhl 1997 Philippines ZMUC https://doi.org/10.5281/zenodo.4504064
S. csoszi Buhl 2004 Papua New Guinea HNHM https://doi.org/10.5281/zenodo.4502781
S. pleuralis Buhl 2004 Australia HNHM https://doi.org/10.5281/zenodo.4502881
S. queenslandicus Buhl 2004 Australia HNHM https://doi.org/10.5281/zenodo.4503338
S. achterbergi Buhl 2008 Indonesia RMNH https://doi.org/10.5281/zenodo.4503326
S. ciliaris Buhl 2008 Indonesia RMNH https://doi.org/10.5281/zenodo.4503318
S. flavispinum Buhl 2008 Indonesia RMNH https://doi.org/10.5281/zenodo.4503314
S. halmaherense Buhl 2008 Indonesia RMNH https://doi.org/10.5281/zenodo.4503235
S. infuscatus Buhl 2008 Indonesia RMNH https://doi.org/10.5281/zenodo.4503181
S. lombokensis Buhl 2008 Indonesia RMNH https://doi.org/10.5281/zenodo.4503152
S. longulum Buhl 2008 Indonesia RMNH https://doi.org/10.5281/zenodo.4503105
S. nigricoxum Buhl 2008 Indonesia RMNH https://doi.org/10.5281/zenodo.4503038
S. praemorsum Buhl 2008 Indonesia RMNH https://doi.org/10.5281/zenodo.4503000
S. quasimodo Buhl 2008 Indonesia RMNH https://doi.org/10.5281/zenodo.4502953
S. dumogabonense 2009 Indonesia NMINH https://doi.org/10.5281/zenodo.4502004
S. masneri Buhl 2009 Indonesia NMINH https://doi.org/10.5281/zenodo.4563030
S. mineoi Buhl 2009 Indonesia NMINH https://doi.org/10.5281/zenodo.4539460
S. butterilli Buhl 2013 Papua New Guinea USNM Paramyristica https://doi.org/10.5281/zenodo.4501840
S. psychotriae Buhl 2013 Papua New Guinea USNM Psychotria https://doi.org/10.5281/zenodo.4501681
S. pterocarpi Buhl 2013 Papua New Guinea USNM Pterocarpus https://doi.org/10.5281/zenodo.4501868
S. alternatum Buhl 2014 Australia ZMUC https://doi.org/10.5281/zenodo.4501898
S. bangmadseni Buhl 2014 Australia ZMUC https://doi.org/10.5281/zenodo.4501912
S. carlseni Buhl 2014 Australia ZMUC https://doi.org/10.5281/zenodo.4503302
S. eucalypti Buhl 2014 Australia ZMUC https://doi.org/10.5281/zenodo.4503566
S. salicorniae Buhl 2014 Australia ZMUC https://doi.org/10.5281/zenodo.4504104
S. striolagaster Buhl 2014 Australia ZMUC https://doi.org/10.5281/zenodo.4504394
S. triangulatum Buhl 2014 Australia ZMUC https://doi.org/10.5281/zenodo.4563047

Results

Character discussion

Clypeal setae

All specimens of Synopeas examined had four clypeal setae, which may be a stable character across the genus (Figure 1). Occasionally two setae may arise from a common base, bringing the total number of setae to five. The relative lengths of the medial two setae to the lateral pair of setae can be helpful in identification, as can the relative distance between the medial and lateral setae. In many species, the medial pair of setae are distinctly longer than the lateral setae (Figure 1A). Some species have the medial pair of setae closer to each other than to the lateral setae (Figure 1B), while others have the four setae evenly spaced (Figure 1C). In S. zhangi, the medial setae are closer to the lateral setae than to each other.

Figure 1. 

Arrangement of clypeal setae in Synopeas A S. anunu (FSCA 00090255) B S. amandae (FSCA 00000323) C S. luli (FSCA 00090412). Scale bars: 0.2 mm.

Lateral ocellar depression

Some species of Synopeas bear a distinct furrow or pit between the lateral ocellus and the compound eye (Figure 16D).

Epomial carina

In the Synopeas species examined in this study, the epomial carina originates next to a pit at the anterodorsal end of the pronotal cervical sulcus (Figure 2). When complete, it extends to the dorsal margin of the pronotum. Occasionally, there is a pit posterior to the dorsal terminus of the carina, as in S. klingunculum (Figure 2A). The carina may be entirely absent (Figure 2B), dorsally weakened or incomplete.

Figure 2. 

Epomial carina in Synopeas A S. klingunculum (FSCA 00000237) B S. codex (FSCA 00000310) C S. toto (FSCA 00000258) D S. toto (FSCA 00000258). Scale bars in mm.

Mesoscutellar spine

The shape of the mesoscutellar spine has historically been used as a species-level diagnostic character for Synopeas. However, there is some intraspecific variation, so it should not be solely relied upon for species diagnosis. The three categories used in this revision are: absent or inconspicuous (Fig. 3A–C), short and pointed (Figure 3D–F), and long and blunt (Figure 3G, H).

Figure 3. 

Morphology of the mesoscutellar spine in Synopeas A S. codex (FSCA 00034030) B S. amandae (FSCA 00000403) C Synopeas sp. (FSCA 00000413) D S. occultum (FSCA 00090403) E S. luli (FSCA 00000334) F S butterilli (FSCA 00090264) G S. klingunculum (FSCA 00000376) H S. anunu (FSCA 00090255) I S. kira (FSCA 00000398).

Margin of lateral propodeal carina in lateral view

In Synopeas, as in most Platygastridae, the dorsal propodeum bears a pair of “keels” or well-developed carinae (Fig. 4), which are relatively close together. The margin of these carinae may be evenly rounded (Fig. 4B), or pointed dorsally (Fig. 4A, C), ventrally, or both. In some species not treated here, the carinae are relatively elongate and posterodorsally flattened. This character can be difficult to interpret and the states require further refinement. The ventral margin of the carinae may be obscured by setae (Fig. 4D).

Figure 4. 

Margin of lateral propodeal carinae A S. klingunculum (FSCA 00000237) B S. toto (FSCA 00000258) C S. codex (FSCA 00000310) D S. valavala (FSCA 00000259). Scale bars in mm.

Mesoscutal lamella

The mesoscutal lamella is a posteromedial extension of the mesoscutum (Fig. 5). Typically, it has a line of setae on either side along the posterior margin of the mesoscutum. The mesoscutal lamella may be rounded (Fig. 5B), square (Fig. 5D), truncate (Fig. 5F), or roughly triangular (Fig. 5H). A broad mesoscutal lamella does not extend further than its own width beyond the posterior mesoscutal margin (Fig. 5B). A narrow mesoscutal lamella extends further than its own width beyond the posterior mesoscutal margin (Fig. 5A).

Figure 5. 

Mesoscutal lamella in Synopeas A S. sanga (FSCA 00000361) B S. luli (FSCA 00000334) C S. valavala (FSCA 00000350) D S. codex (FSCA 00034030) E S. zhangi (FSCA 00000383) F S. klingunculum (FSCA 00000411) G S. kalubia (FSCA 00000344) H S. kira (FSCA 00090416) I undescribed species (FSCA 00034033). Scale bars: 0.2 mm.

Microsculpture of S2 and T2

In most species, the presence of sculpture is absent or restricted to a narrow band on the posterior margins of S2 (Fig. 6) and T2 (Fig. 7). However, in some species, the sculpture is more extensive (Figs 6A–D, 7A, C) and the length and shape of the sculpture can be diagnostic. Here, a “long” band of sculpture on S2 or T2 is defined as an extensive area as long as or longer than S3 or T3, respectively. The length of the band is measured in an anterior-posterior direction.

Figure 6. 

Sculpture of metasomal sternite 2 A S. sanga (FSCA 00000399) B S. occultum (FSCA 00090403) C S. roncavei (FSCA 00090249) D S. kira (FSCA 00000398) E S. codex (FSCA 00034030) F S. luli (FSCA 00090412). Scale bars: 0.2 mm.

Figure 7. 

Sculpture of metasomal tergite 2 A S. sanga (FSCA 00094615) B S. occultum (FSCA 00094465) C S. kira (FSCA 00090258) D S. klingunculum (FSCA 00000411). Scale bar: 0.2 mm.

Linea setosa

The setae on the forewing disc vary in density and arrangement (Fig. 8). The wings of some species are uniformly setose (Fig. 8B). In others, the wing is proximally glabrous, with or without a linea setosa. The linea setosa is a novel term that we use for a curved line of setae at the base of the forewing against a glabrous or sparsely setose background (Fig. 8C). It is thus the inverse of a linea calva, which is a glabrous line against a setose background. The linea setosa cannot be observed on a wing that is uniformly and densely setose.

Figure 8. 

Forewing of Synopeas A S. butterilli (FSCA 00090264) B S. kiki (FSCA 00090410) C S. roncavei (FSCA 00094606) D S. luli (FSCA 00094549). Scale bar: 0.2 mm.

Forewing marginal setae

Most Synopeas lack a long fringe of marginal setae at the posterior margin of the forewing, as is often seen in the closely related Leptacis. However, some Synopeas species have relatively short marginal setae on the anterior and posterior margins of the forewing, and the relative lengths of these setae are a useful character. In the species treated here, the setae on the posterior margin may be equal to (Fig. 8A) or longer than those on the anterior margin of the forewing (Fig. 8B–D).

Limits of Synopeas

Among the Platygastrinae, Leptacis and Synopeas (Synopeadini sensu Kozlov 1970) are characterized by having the lateral propodeal carinae close together, often with an elongate mesoscutellum forming a spine, and sparse to dense setae around the junction of T1 and T2. Masner (1960) differentiated the two genera by the shape and setation of T1 and the length of forewing marginal setae. Jackson (1969) found the fusion of T1 and T2 (Fig. 9A) to be a principal synapomorphy of Synopeas, separating it from Leptacis. He also described a deep depression in the anteroventral corner of the pronotum, or ventral pronotal pit, which is present in Synopeas (Fig. 9B) and absent in Leptacis.

Figure 9. 

Generic characters for identification of Synopeas A Fusion of T1 and T2 in S. klingunculum (FSCA 00000237) B Ventral pronotal pit in S. codex (FSCA 00000310).

We here follow Jackson’s concept of Synopeas. The length of the forewing marginal setae is not always a reliable character for generic separation. On this basis, Leptacis pleuralis (Buhl, 2004), comb. nov., (Fig. 10) is transferred from Synopeas. Although L. pleuralis has very short forewing marginal setae like many Synopeas species, the structure of the metasoma and lack of a ventral pronotal pit lead us to place it in Leptacis.

Figure 10. 

Leptacis pleuralis (Buhl), comb. nov. Inset: anterior metasoma. Scale bar: 0.5 mm.

Key to the species of Synopeas of New Guinea (females)

1 T2 longer than the head and mesosoma combined (Fig. 11A) Synopeas csoszi Buhl
T2 equal to or less than length of head and mesosoma combined (Fig. 11B) 2
2 Notaulus distinctly grooved (Fig. 12A); mesoscutellar spine distinct and conspicuous; lateral portion of frons with rugose macrosculpture 3
Notaulus unmarked or faintly suggested (Fig. 12B); mesoscutellar spine variable; lateral portion of frons without rugose macrosculpture 4
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. 13A); pit present posterior to dorsal terminus of epomial carina Synopeas klingunculum sp. nov.
Length of mesoscutellar spine more than twice its thickness at its midpoint; lateral pronotum with microsculpture dorsally, smooth ventrally (Fig. 13B); without pit near dorsal terminus of epomial carina Synopeas anunu sp. nov.
4 Epomial carina absent or reduced, extending less than halfway up pronotum (Fig. 14A) Synopeas codex sp. nov.
Epomial carina distinct and extending more than halfway up pronotum (Fig. 14B) 5
5 Mesoscutellar spine absent or inconspicuous (Fig. 3A–C) 6
Mesoscutellar spine present, with variable form, often long or pointed (Fig. 3D–I) 9
6 Head large and triangular in anterior view (Fig. 15A); dorsal margin of compound eyes nearly colinear with vertex; length of malar space about equal to height of compound eye Synopeas valavala sp. nov.
Head circular or ovoid in anterior view (Fig. 15B, C); dorsal margin of compound eyes not colinear with vertex; length of malar space less than height of compound eye 7
7 Hyperoccipital carina absent or very weak (Fig. 16A, B); head in frontal view ovoid, distinctly wider than high Synopeas zhangi sp. nov.
Hyperoccipital carina present and complete (Fig. 16C–F); head in frontal view circular to ovoid 8
8 Frons with parallel, arched rugae above torulus; clypeal setae evenly spaced; lateral pronotum without microsculpture in ventral third (Fig. 17A, B) Synopeas pterocarpi Buhl
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. 17C, D) Synopeas amandae sp. nov.
9 Mesoscutellar spine long and blunt (Fig. 3G–I) 10
Mesoscutellar spine short and pointed (Fig. 3D–F) 11
10 Hyperoccipital carina weakly developed and interrupted (Fig. 18A, B) Synopeas kalubia sp. nov.
Hyperoccipital carina robust and complete (Fig. 18C, D) Synopeas kira sp. nov.
11 Central keel well developed and extending to median ocellus (Fig. 19A) Synopeas toto sp. nov.
Central keel absent, weakly developed, or not extending to median ocellus (Fig. 19B–D) 12
12 Patch of microsculpture on posterior S2 extensive, longer than S3 (Fig. 6A, B, D, E) 13
S2 with no microsculpture or with very narrow band at posterior margin (Fig. 6C, F) 14
13 T2 with extensive band of microsculpture at posterior margin that is longer than T3 (Fig. 7A) Synopeas sanga sp. nov.
T2 without microsculpture, or with narrow band at posterior margin (Fig. 7B) Synopeas occultum sp. nov.
14 Parapsidal lines well impressed (Fig. 20A); ocular-ocellar length (OOL) less than 1 ocellar diameter Synopeas psychotriae Buhl
Parapsidal lines unmarked or faintly indicated (Fig. 20B); OOL variable 15
15 Distance between lateral ocellus and hyperoccipital carina at least 1.5 times ocellar diameter (Fig. 21A); dorsal margin of lateral pronotum smooth (Fig. 21C) Synopeas butterilli Buhl
Distance between lateral ocellus and hyperoccipital carina less than 1.5 times ocellar diameter (Fig. 21B); dorsal margin of lateral pronotum with microsculpture (Fig. 21D) 16
16 Hyperoccipital carina robust and complete 17
Hyperoccipital carina weak or incomplete 19
17 Mesoscutum evenly setose; mesoscutellum evenly setose (Fig. 22A) Synopeas pulupulu sp. nov.
Mesoscutum medially glabrous or with few scattered setae; mesoscutellum medially glabrous (Fig. 22B) 18
18 Forewing disc proximally glabrous with linea setosa (Fig. 23A) Synopeas roncavei sp. nov.
Forewing disc without linea setosa (Fig. 23B) Synopeas pattiae sp. nov.
19 Central keel impressed or partially present only in ventral part of frons (Fig. 24A); mesoscutellar spine angled posterodorsally (Fig. 24C); posterior margin of lateral propodeal carina pointed ventrally in lateral view (Fig. 24C) Synopeas luli sp. nov.
Central keel absent (Fig. 24B); mesoscutellar spine posteriorly pointed (Fig. 24D); posterior margin of lateral propodeal carina rounded ventrally in lateral view (Fig. 24D) Synopeas kiki sp. nov.
Figure 11. 

Lateral habitus A S. csoszi B S. occultum (FSCA 00090403).

Figure 12. 

Dorsal mesoscutum and mesoscutellum A S. anunu (FSCA 00090255) B S. occultum (FSCA 00090403). Scale bars: 0.2 mm.

Figure 13. 

Mesoscutellar spine A S. klingunculum (FSCA 00000376) B S. anunu (FSCA 00090255). Scale bars: 0.2 mm.

Figure 14. 

Lateral pronotum A S. codex (FSCA 00034030) B S. amandae (FSCA 00000403). Scale bars: 0.2 mm.

Figure 15. 

Frontal head A S. valavala (FSCA 00033890) B S. codex (FSCA 00034030) C S. amandae (FSCA 00000323). Scale bars 0.2 mm.

Figure 16. 

Lateral and dorsal head A S. zhangi (FSCA 00000291) B S. zhangi (FSCA 00000383) C S. amandae (FSCA 00000403) D S. amandae (FSCA 00000403) E S. pterocarpi (USNMENT01335974) F S. pterocarpi (USNMENT01335974). Scale bars: 0.2 mm.

Figure 17. 

Frons and lateral pronotum A S. pterocarpi (USNMENT01335974) B S. pterocarpi (USNMENT01335974) C S. amandae (FSCA 00000323) D S. amandae (FSCA 00000403). Scale bars: 0.2 mm.

Figure 18. 

Lateral and dorsal head A S. kalubia (FSCA 00000344) B S. kalubia (FSCA 00000344) C S. kira (FSCA 00000398) D S. kira (FSCA 00090258). Scale bars: 0.2 mm.

Figure 19. 

Central keel A S. toto (FSCA 00090421) B S. sanga (FSCA 00000399) C S. luli (FSCA 00090412) D S. psychotriae (FSCA 00090261). Scale bars: 0.2 mm.

Figure 20. 

Dorsal mesosoma showing parapsidal lines A S. psychotriae (FSCA 00090261) B S. butterilli (FSCA 00090264). Scale bars: 0.2 mm.

Figure 21. 

Head and lateral pronotum A S. butterilli (FSCA 00090264) B S. kiki (FSCA 00090410) C S. butterilli (FSCA 00090264) D S. luli (FSCA 00000334). Scale bars: 0.2 mm.

Figure 22. 

Setation of dorsal mesoscutum and mesoscutellum A S. pulupulu (FSCA 00090430) B S. roncavei (FSCA 00090249). Scale bars: 0.2 mm.

Figure 23. 

Setation of forewing disc including linea setosa A S. roncavei (FSCA 00094606) B S. pattiae (FSCA 00090423). Scale bar: 0.2 mm.

Figure 24. 

Head, mesoscutellum, and lateral propodeal carina A S. luli (FSCA 00090412) B S. kiki (FSCA 00090410) C S. luli (FSCA 00000334) D S. kiki (FSCA 00090410). Scale bars: 0.2 mm.

Descriptions

Synopeas amandae Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: slightly ovoid. Central keel: complete; partial. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: minute rugulae; reticulate microsculpture. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Lateral ocellar depression: present laterally. Hyperoccipital carina: complete. Hyperoccipital carina strength: laterally weakened. Distance between lateral ocellus and hyperoccipital carina: approximately 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present throughout. Lateral pronotal sculpture coverage: more than 3/4. Mesoscutellar spine: absent or inconspicuous. Posterior margin of lateral propodeal carina in lateral view: rounded. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: moderately dense. Mesoscutal lamella: broad and square. Setation of mesoscutellum: laterally dense, medially sparse.

Metasoma. Microsculpture of S2: narrow band at posterior margin. Sculpture of T2: narrow transverse band of microsculpture at posterior margin. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: longer than distance between setal bases. Density of setae on disc of forewing: dense. Arrangement of setae on disc of forewing: uniformly setose distally, proximally glabrous with linea setosa. Forewing marginal setae: setae on posterior margin distinctly longer than setae on anterior margin; approximately uniform in length on anterior and posterior margins.

Diagnosis

Synopeas amandae has the pronotum entirely covered with reticulate microsculpture, which is not typical among the species treated here. The absent or inconspicuous mesoscutellar spine also helps to identify it. It can be distinguished from the similar S. codex by the shape of the head, which is blocky in S. codex, and by the epomial carina, which is complete or nearly so in S. amandae and absent or reduced in S. codex.

Comments

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.

Figure 25. 

Synopeas amandae A lateral habitus (female holotype; FSCA 00000403) B dorsal habitus (female holotype; FSCA 00000403) C ventral habitus (female paratype; FSCA 00000323).

Etymology

This species is named in honor of the entomologist Amanda Hodges.

Plant associations

Reared from variously shaped galls on Nauclea sp. (Rubiaceae) [GALL218, GALL219, GALL223].

Material examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 16.XI.2010, Philip Butterill leg., Nauclea sp. 1, (FSCA 00000403). Paratypes: Papua New Guinea: 7♀, Morobe, Yawan, 15.XI.2010, Nauclea sp. 1, (FSCA 00000037, 00000313–00000318); 2♀, Morobe, Yawan, 16.XI.2010, Nauclea sp. 1, (FSCA 00094594–00094595); 1♀, Morobe, Yawan, 25.XI.2010, Nauclea sp. 1, (FSCA 00000323); 1♀, Morobe, Yawan, 26.I.2011, Nauclea sp. 1, (FSCA 00000331); 6♀, Morobe, Yawan, 04.X.2012, Nauclea sp. 1, (FSCA 00090253, 00094584–00094588).

Figure 26. 

Synopeas anunu (female holotype; FSCA 00090255) A lateral habitus B dorsal habitus C ventral habitus.

Synopeas anunu Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: distinctly ovoid. Central keel: partial. Sculpture on frons: rugose macrosculpture. Epitorular sculpture: rugose. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): approximately 1 ocellar diameter; greater than 1 ocellar diameter. Hyperoccipital carina: absent.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally. Lateral pronotal sculpture coverage: 1/4 to 1/2. Mesoscutellar spine: long and thick. Mesoscutellar spine in lateral view: pointing posteriorly. Posterior margin of lateral propodeal carina in lateral view: pointed dorsally and ventrally. Mesosomal dorsum in lateral view: convex. Notauli: distinctly grooved. Parapsidal lines: marked by absence of sculpture. Setation of mesoscutum: sparse. Mesoscutal lamella: truncate, with striate sculpture. Setation of mesoscutellum: present lateral to striate sculpture.

Metasoma. Microsculpture of S2: narrow band at posterior margin. Sculpture of T2: microsculpture present in patch at posterior margin, not extending to posterolateral corners. Length of T2: shorter than mesosoma.

Wing. Length of setae on disc of forewing: shorter than distance between setal bases. Density of setae on disc of forewing: moderately dense. Arrangement of setae on disc of forewing: setose distally, glabrous proximally. Forewing marginal setae: setae on posterior margin distinctly longer than setae on anterior margin.

Diagnosis

The head of this species is mostly covered in rugose macrosculpture and the notauli are distinctly grooved, characters which it shares with S. klingunculum. The mesoscutellar spine of S. anunu is long and thick, with its length more than twice the thickness at its midpoint (Fig. 13B). Synopeas klingunculum has a mesoscutellar spine that is more wart-shaped, with the length only slightly greater than its thickness in lateral view (Fig. 13A). The sculpture of the lateral pronotum in S. klingunculum is present dorsally and absent ventrally, with a smooth band anterior to the tegula. This smooth band is lacking in S. anunu. Additionally, the epomial carina of S. klingunculum terminates posteriorly in a pit, while that of S. anunu does not.

Comments

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.

Etymology

The epithet “anunu” is Manam for “shadow” (Osmond and Ross 2016). It is applied to this species because it resembles S. klingunculum, being distinguishable based on the silhouette or shadow of the mesoscutellar spine. The name is to be treated as a noun in apposition.

Plant associations

Reared from a vein swelling gall on Nauclea sp. (Rubiaceae) [GALL217].

Material examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 30.XI.2010, Philip Butterill leg., Nauclea sp. 1 (FSCA 00090255). Paratypes: Papua New Guinea: 1♀1♂, Morobe, Yawan, 6.14141°S, 146.87415°E, 30.XI.2010, Philip Butterill leg., Nauclea sp. 1 (FSCA 00094589–00094590).

Synopeas butterilli Buhl, 2013

Description

Body length 1.3 mm. Body color: black. Color of legs: coxae brown, otherwise yellow. Color of mesoscutellar spine: concolorous with mesoscutellar disc.

Head. Shape of head in anterior view: circular. Central keel: absent. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: minute rugulae. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: evenly spaced; medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Hyperoccipital carina: complete. Hyperoccipital carina strength: uniformly robust. Distance between lateral ocellus and hyperoccipital carina: greater than 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present medially, absent ventrally and dorsally. Lateral pronotal sculpture coverage: 1/4 to 1/2. Mesoscutellar spine: short and pointed. Mesoscutellar spine in lateral view: pointing posteriorly; angled posterodorsally. Posterior margin of lateral propodeal carina in lateral view: rounded. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: sparse. Mesoscutal lamella: short, projecting less than its width beyond margin; broad and rounded. Setation of mesoscutellum: laterally dense, medially sparse.

Figure 27. 

Synopeas butterilli (male specimen; FSCA 00090264) A lateral habitus B dorsal habitus C anteroventral habitus.

Metasoma. Microsculpture of S2: narrow band at posterior margin. Sculpture of T2: narrow transverse band of microsculpture at posterior margin. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: shorter than distance between setal bases. Density of setae on disc of forewing: sparse. Arrangement of setae on disc of forewing: uniformly setose distally, proximally glabrous with linea setosa. Forewing marginal setae: approximately uniform in length on anterior and posterior margins.

Diagnosis

The lateral ocellus and the hyperoccipital carina are widely separated in S. butterilli, by approximately 1.5 ocellar diameters. The sculpture of the lateral pronotum is restricted to a narrow line medially, and the lateral pronotum is smooth dorsally and ventrally. These two characters set it apart from other species with a short, pointed mesoscutellar spine: S. luli, S. occultum, S. roncavei, S. sanga, and S. toto.

Comments

The female holotype and paratype of S. butterilli have a posterodorsally pointed mesoscutellar spine. The male specimens examined have a posteriorly pointed mesoscutellar spine. The angle of the mesoscutellar spine may be sexually dimorphic in this species, but more specimens are needed to confirm this. The central keel is present and complete, but very weak and may be difficult to see.

Plant associations

Reared from galls on Paramyristica cf. sepicana (Myristicaceae). Occasionally found in the same gall as Inostemma sp.

Material examined

Holotype: Papua New Guinea: ♀, Madang, Mis 5.183°S, 145.758°E; 15.IX.2010–17.II.2011, Philip Butterill leg. Paramyristica sp. (USNMENT01335986). Paratype: Papua New Guinea: ♀, Madang, Mis 5.183°S, 145.758°E; 15.IX.2010–17.II.2011; Philip Butterill leg.; Paramyristica sp. (USNMENT01335984). Other material: ♂, Madang, Mis, 5.183°S, 145.758°E; 16.VII.2010; Philip Butterill leg. Paramyristica cf sepicana. (FSCA 00090264); ♂, Madang, Mis, 17.II.2011, Paramyristica cf sepicana. (FSCA 00090428).

Synopeas codex Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: blocky. Central keel: partial. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: minute rugulae; reticulate microsculpture. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Lateral ocellar depression: present laterally. Hyperoccipital carina: present in medial third. Hyperoccipital carina strength: uniformly weak. Distance between lateral ocellus and hyperoccipital carina: greater than 1 ocellar diameter.

Mesosoma. Epomial carina: absent; present only in ventral half. Microsculpture of lateral pronotum: present throughout. Lateral pronotal sculpture coverage: more than 3/4. Mesoscutellar spine: absent or inconspicuous. Posterior margin of lateral propodeal carina in lateral view: rounded; pointed dorsally. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: unmarked. Setation of mesoscutum: absent medially, present laterally. Mesoscutal lamella: broad and rounded; broad and square. Setation of mesoscutellum: absent medially.

Metasoma. Microsculpture of S2: present on posterior half. Sculpture of T2: long transverse band of microsculpture at posterior margin. Length of T2: shorter than mesosoma.

Wing. Length of setae on disc of forewing: longer than distance between setal bases. Density of setae on disc of forewing: dense. Arrangement of setae on disc of forewing: uniformly setose distally, proximally glabrous with linea setosa. Forewing marginal setae: setae on posterior margin distinctly longer than setae on anterior margin.

Diagnosis

The shape of the head in S. codex is distinctly blocky in anterior view. Among the species treated here, Synopeas codex may be recognized by the absent or reduced epomial carina. When present, the epomial carina is short, extending less than halfway up the pronotum. S2 is covered with reticulate microsculpture on the posterior half (Fig. 6E), and there is a wide band of microsculpture at the posterior margin of dorsal T2.

Figure 28. 

Synopeas codex (female paratype; FSCA 00034030) A lateral habitus B dorsal habitus C ventral habitus.

Etymology

The epithet “codex” is Latin for a book or block of wood. We here used it to refer to the shape of the head in this species. The name is to be treated as a noun in apposition.

Comments

The distance between the dorsal margin of the eye and the vertex is very small, as in S. valavala. It thus may represent an intermediate form between the typical head shape in Synopeas and the extreme found in S. valavala. The bidentate mandibles are rather short, just meeting in the middle rather than overlapping.

Plant associations

Reared from vein swelling, globoid, pimple, and blister galls on Nauclea (Rubiaceae) [GALL217, GALL218, GALL222, GALL219]. One record each from Homalanthus nervosus (Euphorbiaceae) [GALL354] and Ficus adenosperma (Moraceae) [GALL318].

Material examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 26.XI.2010, Philip Butterill leg., Nauclea sp. 1 (FSCA 00000385). Paratypes: Papua New Guinea: 1♀, Morobe, Yawan, 24.VII.2010, Nauclea sp. 1 (FSCA 00000352); ♂, Morobe, Yawan, 16.XI.2010, Nauclea sp. 1 (FSCA 00000374); ♀, Morobe, Yawan, 17.XI.2010, Homalanthus nervosus, (FSCA 00000377); ♀ 2♂, Morobe, Yawan, 25.XI.2010, Nauclea sp. 1 (FSCA 00034030, 00094609–00094610); ♂, Morobe, Yawan, 26.XI.2010, Nauclea sp. 1 (FSCA 00000386) 4♀ 4♂, Morobe, Yawan, 30.XI.2010, Nauclea sp. 1 (FSCA 00000380–00000382, 00000388–00000392); ♂, Morobe, Yawan, 25.I.2011, Ficus adenosperma, (FSCA 00090425); 3♀, Morobe, Yawan, 08.X.2012, Nauclea tenuiflora, (FSCA 00000310, 00034029 00094583).

Figure 29. 

Holotype of Synopeas csoszi.

Synopeas csoszi Buhl, 2004

Description

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.

Head. Shape of head in anterior view: distinctly ovoid. Central keel: absent. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: minute rugulae. Number of clypeal setae: unknown. Length of median pair of clypeal setae: unknown. Arrangement of clypeal setae: unknown. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): approximately 1 ocellar diameter. Hyperoccipital carina: complete. Hyperoccipital carina strength: uniformly robust. Distance between lateral ocellus and hyperoccipital carina: approximately 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present anterodorsally, absent posteroventrally. Lateral pronotal sculpture coverage: unknown. Mesoscutellar spine: long and thin. Mesoscutellar spine in lateral view: pointing posteriorly. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: uncertain, absent medially, present laterally. Mesoscutal lamella: broad and rounded; roughly triangular. Setation of mesoscutellum: laterally dense, medially sparse.

Metasoma. Microsculpture of S2: narrow band at posterior margin. Sculpture of T2: narrow transverse band of microsculpture at posterior margin. Length of T2: longer than head and mesosoma combined.

Wing. Length of setae on disc of forewing: shorter than distance between setal bases. Density of setae on disc of forewing: dense. Arrangement of setae on disc of forewing: uniformly setose distally, proximally glabrous with linea setosa. Forewing marginal setae: approximately uniform in length on anterior and posterior margins.

Diagnosis

Synopeas csoszi is easily recognized by the extraordinary length of the second metasomal segment, which is longer than the head and mesosoma combined. The metasoma is narrow anteriorly, expanding posteriorly to become wider than the rest of the body at the junction between metasomal segments 2 and 3. The shape of the antennal scape is unusually expanded distally forming a club, then contracting into a narrowly curved apex (Fig. 29).

Comments

The forewing disc is setose distally and glabrous proximally, with a broad linea setosa.

Plant associations

Unknown.

Material examined

Holotype: Papua New Guinea: ♀, Mt. Wilhelm, alt. 3900 m, 1324.IX.1968, Janos Balogh leg. Paratypes: Papua New Guinea: ♀ ♂, Mt. Wilhelm, alt. 3900 m, 1324.IX.1968, Janos Balogh leg. (deposited in HNHM).

Synopeas kalubia Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: slightly ovoid. Central keel: absent. Sculpture on frons: rugulose macrosculpture. Epitorular sculpture: minute rugulae. Number of clypeal setae: 4. Length of median pair of clypeal setae: uncertain, longer than lateral pair. Arrangement of clypeal setae: unknown. Shape of mandible: unknown. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Hyperoccipital carina: complete. Hyperoccipital carina strength: medially weakened. Distance between lateral ocellus and hyperoccipital carina: approximately 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present throughout. Lateral pronotal sculpture coverage: more than 3/4. Mesoscutellar spine: long and thick. Mesoscutellar spine in lateral view: pointing posteriorly. Posterior margin of lateral propodeal carina in lateral view: pointed dorsally. Mesosomal dorsum in lateral view: weakly convex. Notauli: unmarked or faintly suggested. Parapsidal lines: weakly impressed. Setation of mesoscutum: moderately dense. Mesoscutal lamella: short and narrow. Setation of mesoscutellum: laterally dense, medially sparse.

Figure 30. 

Synopeas kalubia (female holotype; FSCA 00000344) A lateral habitus B dorsal habitus C ventral habitus.

Metasoma. Microsculpture of S2: narrow band at posterior margin. Sculpture of T2: narrow transverse band of microsculpture at posterior margin. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: longer than distance between setal bases. Density of setae on disc of forewing: dense. Arrangement of setae on disc of forewing: uniformly setose distally, proximally glabrous with linea setosa. Forewing marginal setae: approximately uniform in length on anterior and posterior margins.

Diagnosis

Important characters for identifying S. kalubia include the weak hyperoccipital carina and the microsculpture of the lateral pronotum, which is absent only in the ventralmost portion. It can be distinguished from S. kira by the pattern of microsculpture on S2. In S. kalubia, the microsculpture is limited to a narrow band along the posterior margin that is half as long as that of S. kira (Fig. 6D), and S. kira also has a well-developed hyperoccipital carina. The notauli of S. kalubia are not indicated, which allows it to be easily separated from S. klingunculum and S. anunu.

Etymology

The epithet “kalubia” is Bariai for “barracuda” (Osmond, 2011) and refers to the streamlined appearance of this species. The name is to be treated as a noun in apposition.

Plant associations

Reared from a blister gall on Nauclea sp. (Rubiaceae) [GALL219].

Material examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 30.XI.2010, Philip Butterill leg., Nauclea sp. 1 (FSCA 00000344).

Synopeas kiki Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: slightly ovoid. Central keel: absent. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: minute rugulae. Number of clypeal setae: unknown. Length of median pair of clypeal setae: unknown. Arrangement of clypeal setae: uncertain, medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): approximately 1 ocellar diameter. Hyperoccipital carina: complete. Hyperoccipital carina strength: medially weakened; uniformly weak. Distance between lateral ocellus and hyperoccipital carina: approximately 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present anterodorsally, absent posteroventrally. Lateral pronotal sculpture coverage: 1/4 to 1/2. Mesoscutellar spine: short and pointed. Mesoscutellar spine in lateral view: pointing posteriorly. Posterior margin of lateral propodeal carina in lateral view: rounded. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: sparse. Mesoscutal lamella: long and narrow. Setation of mesoscutellum: laterally dense, medially sparse.

Metasoma. Microsculpture of S2: absent. Sculpture of T2: absent; narrow transverse band of microsculpture at posterior margin. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: longer than distance between setal bases. Density of setae on disc of forewing: moderately dense. Arrangement of setae on disc of forewing: unknown. Forewing marginal setae: setae on posterior margin distinctly longer than setae on anterior margin.

Diagnosis

Identification of S. kiki requires the use of numerous characters because it does not have any obviously distinct features. It is similar to S. roncavei, from which it can be separated by the development of the hyperoccipital carina, which is medially weakened in S. kiki and uniformly robust in S. roncavei. It is also similar to S. luli, but S. kiki has a posteriorly angled mesoscutellar spine and no central keel, while S. luli has a posterodorsally angled mesoscutellar spine and a partial central keel. The forewing of S. luli has setae of approximately equal length around the wing margin, while in S. kiki, the setae are distinctly longer on the posterior margin (Fig. 8).

Figure 31. 

Synopeas kiki (female holotype; FSCA 00090410) A lateral habitus B dorsal habitus C ventral habitus.

Etymology

The epithet “kiki” is Proto-Oceanic for “small” (Ross and Osmond 2016) and refers to the relatively diminutive size of this species. The name is to be treated as a noun in apposition.

Plant associations

Reared from a pimple gall on Nauclea sp. (Rubiaceae) [GALL222].

Material examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 25.XI.2010, Philip Butterill leg., Nauclea sp. 1 (FSCA 00090410).

Synopeas kira Awad, sp. nov.

Description

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.

Figure 32. 

Synopeas kira A lateral habitus (female paratype; FSCA 00000398) B dorsal habitus (female paratype; FSCA 00090258) C ventral habitus (female paratype; FSCA 00000398).

Head. Shape of head in anterior view: slightly ovoid. Central keel: absent. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: minute rugulae; reticulate microsculpture. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: evenly spaced; medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Hyperoccipital carina: complete. Hyperoccipital carina strength: uniformly robust. Distance between lateral ocellus and hyperoccipital carina: less than 1 ocellar diameter; approximately 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally. Lateral pronotal sculpture coverage: 1/2 to 2/3; 2/3 to 3/4. Mesoscutellar spine: long and thick. Mesoscutellar spine in lateral view: pointing posteriorly. Posterior margin of lateral propodeal carina in lateral view: pointed dorsally and ventrally. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: sparse. Mesoscutal lamella: long and narrow; roughly triangular. Setation of mesoscutellum: laterally dense, medially sparse.

Metasoma. Microsculpture of S2: long band at posterior margin. Sculpture of T2: long transverse band of microsculpture at posterior margin; microsculpture present in patch at posterior margin, not extending to posterolateral corners. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: longer than distance between setal bases. Density of setae on disc of forewing: dense. Arrangement of setae on disc of forewing: uniformly setose distally, proximally glabrous with linea setosa. Forewing marginal setae: setae on posterior margin distinctly longer than setae on anterior margin.

Diagnosis

Synopeas kira is similar to S. kalubia, from which it can be separated by the pattern of microsculpture at the posterior margin of S2. In S. kira, the sculpture forms a wide band, longer than S3 (Fig. 6D), and in S. kalubia it is limited to a very narrow band. The absent or faint notauli of S. kira enable it to be easily separated from S. klingunculum and S. anunu. Synopeas kira has a dorsally convex and blunt mesoscutellar spine, distinguishing it from S. sanga and S. occultum, both of which have a short, pointed mesoscutellar spine.

Comments

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.

Etymology

The epithet “kira” is Manam for “stone axe” (Osmond and Ross 1998) and refers to the shape of the mesoscutellar spine. The name is to be treated as a noun in apposition.

Plant associations

Reared from galls on Cypholophus (Urticaceae) [GALL266, GALL428, GALL429]. One record from Debregeasia longifolia (Urticaceae) [GALL268].

Specimens examined. Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 17.VII.2012, Philip Butterill leg., Cypholophus macrocephalus (FSCA 00090258). Paratypes: Papua New Guinea: ♀, Morobe, Yawan, 25.XI.2010, Debregeasia longifolia, (FSCA 00090431); ♂, Morobe, Yawan, 25.I.2011, Cypholophus friesianus (FSCA 00090434); 12♀ 2♂, Morobe, Yawan, 28.III.2012, Cypholophus sp. 1 (FSCA 00090416, 00094498–00094501, 00094516–00094523, 00094616); 2♂, Morobe, Yawan, 29.III.2012, Cypholophus sp. 1 (FSCA 00000416, 00094503); 2♂, Morobe, Yawan, 10.VII.2012, Cypholophus sp. 1 (FSCA 00000398, 00094546); ♀ ♂, Morobe, Yawan, 17.VII.2012, Cypholophus macrocephalus (FSCA 00094566–00094567).

Synopeas klingunculum Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: distinctly ovoid. Central keel: partial. Sculpture on frons: rugose macrosculpture. Epitorular sculpture: rugose. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): approximately 1 ocellar diameter. Hyperoccipital carina: interrupted medially; present in medial third; absent; obscured by sculpture. Hyperoccipital carina strength: uniformly weak. Distance between lateral ocellus and hyperoccipital carina: approximately 1 ocellar diameter; greater than 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally, smooth band anterior to tegula. Lateral pronotal sculpture coverage: 1/4 to 1/2. Mesoscutellar spine: long and thick. Mesoscutellar spine in lateral view: pointing posteriorly. Posterior margin of lateral propodeal carina in lateral view: rounded; pointed dorsally. Mesosomal dorsum in lateral view: convex. Notauli: distinctly grooved. Parapsidal lines: marked by absence of sculpture. Setation of mesoscutum: sparse. Mesoscutal lamella: truncate, with striate sculpture. Setation of mesoscutellum: present lateral to striate sculpture.

Figure 33. 

Synopeas klingunculum A lateral habitus (female paratype; FSCA 00000376) B dorsal habitus (male paratype; FSCA 00000411) C ventral habitus (male paratype; FSCA 00000411).

Metasoma. Microsculpture of S2: narrow band at posterior margin. Sculpture of T2: narrow transverse band of microsculpture at posterior margin. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: shorter than distance between setal bases. Density of setae on disc of forewing: sparse. Arrangement of setae on disc of forewing: densely setose distally, sparsely setose proximally. Forewing marginal setae: setae on posterior margin distinctly longer than setae on anterior margin.

Diagnosis

The head of S. klingunculum is mostly covered in rugose macrosculpture and the notauli are distinctly grooved, characters which it shares with S. anunu. The mesoscutellar spine of S. klingunculum is papilliform or wart-like, with the length of the spine only slightly greater than its thickness in lateral view. Synopeas anunu has a mesoscutellar spine that is similar in form, but its length is more than twice the thickness at its midpoint (Fig. 13). The sculpture of the lateral pronotum in S. klingunculum is present dorsally and absent ventrally, with a smooth band anterior to the tegula. This smooth band is lacking in S. anunu. Additionally, the epomial carina of S. klingunculum terminates posteriorly in a pit, while that of S. anunu does not.

Comments

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.

Etymology

The epithet “klingunculum” means “little Klingon” and refers to the rugose head sculpture, which resembles that of the fictional alien race from “Star Trek”.

Plant associations

Reared from vein swelling and globoid galls on Nauclea spp. (Rubiaceae) [GALL217, GALL218].

Material examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 17.X.2012, Philip Butterill leg., Nauclea tenuiflora (FSCA 00034032). Paratypes: Papua New Guinea: ♀, Morobe, Yawan, 01.VII.2010, Nauclea sp. 1 (FSCA 00000365); ♀, Morobe, Yawan, 23.VII.2010, Nauclea sp. 1 (FSCA 00000368); ♀, Morobe, Yawan, 24.VIII.2010, Nauclea sp. 1, (FSCA 00000309); 2♀ 3♂, Morobe, Yawan, 22.IX.2010, Nauclea tenuiflora (FSCA 00000362–00000364, 00000378–00000379); ♀, Morobe, Yawan, 26.X.2010, Nauclea sp. 1 (FSCA 00000369); 2♂, Morobe, Yawan, 30.X.2010, Nauclea sp. 1 (FSCA 00090254–00094591); ♀, Morobe, Yawan, 12.XI.2010, Nauclea sp. 1 (FSCA 00000354); ♂, Morobe, Yawan, 26.XI.2010, Nauclea sp. 1 (FSCA 00000411); ♀ ♂, Morobe, Yawan, 29.XI.2010, Nauclea sp. 1 (FSCA 00000343, 00090408); ♀, Morobe, Yawan, 26.I.2011, Nauclea sp. 1 (FSCA 00000375); ♀, Morobe, Yawan, 01.VIII.2012, Nauclea sp. 1 (FSCA 00000366); 15♀ 4♂, Morobe, Yawan, 13.XI.2012, Nauclea sp. 1 (FSCA 00000237, 00000355, 00034028, FSCA 00094485–00094492, 00094504–00094510); ♀, Morobe, Yawan, 21.XI.2012, Nauclea sp. 1 (FSCA 00000376).

Synopeas luli Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: slightly ovoid. Central keel: absent; partial; impressed. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: minute rugulae; reticulate microsculpture. Number of clypeal setae: 4. Length of median pair of clypeal setae: equal to lateral pair. Arrangement of clypeal setae: evenly spaced; medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): approximately 1 ocellar diameter. Hyperoccipital carina: complete. Hyperoccipital carina strength: uniformly weak. Distance between lateral ocellus and hyperoccipital carina: less than 1 ocellar diameter; approximately 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally, smooth band anterior to tegula. Lateral pronotal sculpture coverage: 1/4 to 1/2. Mesoscutellar spine: short and pointed. Mesoscutellar spine in lateral view: angled posterodorsally. Posterior margin of lateral propodeal carina in lateral view: pointed dorsally and ventrally. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint; marked by absence of sculpture. Setation of mesoscutum: sparse. Mesoscutal lamella: broad and rounded; roughly triangular. Setation of mesoscutellum: laterally dense, medially sparse.

Figure 34. 

Synopeas luli A lateral habitus (male paratype; FSCA 00000334) B dorsal habitus (male paratype; FSCA 00000334) C ventral habitus (female holotype; FSCA 00090412).

Metasoma. Microsculpture of S2: narrow band at posterior margin. Sculpture of T2: absent. Length of T2: shorter than mesosoma.

Wing. Length of setae on disc of forewing: longer than distance between setal bases. Density of setae on disc of forewing: moderately dense. Arrangement of setae on disc of forewing: setose distally, glabrous proximally; densely setose distally, sparsely setose proximally. Forewing marginal setae: approximately uniform in length on anterior and posterior margins.

Diagnosis

Synopeas luli has a short, pointed mesoscutellar spine that is angled upward, and a weak but complete hyperoccipital carina. It resembles S. kiki and may be distinguished by the presence of a partial central keel on the frons and by the mesoscutellar spine, which is posterodorsally angled in S. luli and posteriorly angled in S. kiki. The forewing of S. luli has setae of approximately equal length around the wing margin, while in S. kiki, the setae are distinctly longer on the posterior margin (Fig. 8).

Comments

The central keel is variable in this species. It can be absent, or present ventrally within a shallow longitudinal depression.

Etymology

The epithet “luli” is Numbani for “thorn” (Evans 2008) and refers to the shape of the mesoscutellar spine. The name is to be treated as a noun in apposition.

Plant associations

Reared from galls on Neuburgia corynocarpa (Apocynaceae) [GALL236] and Piper macropiper (Piperaceae) [GALL307].

Specimens examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 22.X.2012, Philip Butterill leg., Neuburgia corynocarpa (FSCA 00090412). Paratypes: Papua New Guinea: 5♀ 3♂, Morobe, Yawan, 22.IX.2010, Neuburgia corynocarpa (FSCA 00090427, 00090437–00090438, 00094466–00094470); 3♀ 2♂, Morobe, Yawan, 03.XI.2010, Piper macropiper (FSCA 00090426, 00090429, 00094482–00094484); ♀, Morobe, Yawan, 30.III.2012, Neuburgia corynocarpa (FSCA 00090440); ♂, Morobe, Yawan, 14.VI.2012, Neuburgia corynocarpa (FSCA 00090439); 7♀ 4♂, Morobe, Yawan, 22.X.2012, Neuburgia corynocarpa (FSCA 00094524–00094528, 00094548–00094549, 00000332–00000335).

Synopeas occultum Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: slightly ovoid. Central keel: absent. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: reticulate microsculpture. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Lateral ocellar depression: present laterally. Hyperoccipital carina: complete. Hyperoccipital carina strength: uniformly robust. Distance between lateral ocellus and hyperoccipital carina: less than 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally. Lateral pronotal sculpture coverage: 1/2 to 2/3. Mesoscutellar spine: short and pointed. Mesoscutellar spine in lateral view: pointing posteriorly. Posterior margin of lateral propodeal carina in lateral view: pointed ventrally. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: sparse. Mesoscutal lamella: short, projecting less than its width beyond margin. Setation of mesoscutellum: laterally dense, medially sparse.

Metasoma. Microsculpture of S2: present at posterior margin and in posterolateral corners. Sculpture of T2: narrow transverse band of microsculpture at posterior margin. Length of T2: approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: longer than distance between setal bases. Density of setae on disc of forewing: dense. Arrangement of setae on disc of forewing: uniformly setose distally, proximally glabrous with linea setosa. Forewing marginal setae: setae on posterior margin distinctly longer than setae on anterior margin.

Figure 35. 

Synopeas occultum (female holotype; FSCA 00090403) A lateral habitus B dorsal habitus C ventral habitus.

Diagnosis

Synopeas occultum has a short, pointed scutellar spine and a robust, complete hyperoccipital carina. It may be distinguished from similar-looking species by the patch of microsculpture on posterior and posterolateral S2 (Fig. 6B). It resembles S. sanga, but differs in the length of T2, which is much more elongate in S. occultum (compare Figs 35 and 41). Additionally, S. sanga has a long patch of microsculpture at the dorsal margin of T2, which in S. occultum is much shorter (Fig. 7).

Etymology

The epithet “occultum” is Latin for “hidden”. It is applied to this species because it was not initially recognized from morphology alone, and its hidden identity was revealed by DNA barcoding.

Plant associations

Reared from nodule-like galls on Cypholophus friesianus (Urticaceae) [GALL266] .

Material examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 24.V.2012, Philip Butterill leg., Cypholophus friesianus (FSCA 00090403). Paratypes: Papua New Guinea: ♀ 4♂, Morobe, Yawan, 04.X.2010, Cypholophus friesianus (FSCA 00090435, 00094460–00094463); 3♂, Morobe, Yawan, 07.X.2010, Cypholophus friesianus (FSCA 00090436, 00094464–00094465); ♂, Morobe, Yawan, 29.XI.2010, Cypholophus friesianus (FSCA 00090433); ♀, Morobe, Yawan, 24.V.2012, Cypholophus friesianus (FSCA 00094493).

Figure 36. 

Synopeas pattiae (female holotype; FSCA 00090423). A Lateral habitus B Dorsal habitus C Ventral habitus.

Synopeas pattiae Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: circular. Central keel: absent; partial. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: minute rugulae; reticulate microsculpture. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Hyperoccipital carina: complete. Hyperoccipital carina strength: uniformly robust. Distance between lateral ocellus and hyperoccipital carina: approximately 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally. Lateral pronotal sculpture coverage: 1/2 to 2/3. Mesoscutellar spine: short and pointed. Mesoscutellar spine in lateral view: pointing posteriorly; angled posterodorsally. Posterior margin of lateral propodeal carina in lateral view: pointed ventrally. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: absent medially, present laterally. Mesoscutal lamella: broad and rounded; roughly triangular. Setation of mesoscutellum: absent medially; laterally dense, medially sparse.

Metasoma. Microsculpture of S2: narrow band at posterior margin. Sculpture of T2: narrow transverse band of microsculpture at posterior margin. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: uncertain, longer than distance between setal bases. Density of setae on disc of forewing: moderately dense. Arrangement of setae on disc of forewing: setose distally, glabrous proximally. Forewing marginal setae: approximately uniform in length on anterior and posterior margins.

Diagnosis

Synopeas pattiae has a short, pointed mesoscutellar spine and a robust, complete hyperoccipital carina. Synopeas pattiae can be differentiated from S. pulupulu by the setation of the meoscutum and mesoscutellum, which are medially glabrous in S. pattiae and uniformly setose in S. pulupulu. Synopeas pattiae is similar to S. roncavei, but the forewing of S. pattiae lacks a linea setosa, which is present in S. roncavei (Fig. 23).

Comments

The central keel may be ventrally present or entirely absent. The forewing disc is proximally glabrous or with a few scattered setae.

Etymology

This species is named in honor of botanist Patti J. Anderson.

Plant associations

Reared from globoid galls on Syzygium decipiens (Myrtaceae) [GALL419].

Material examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 03.XII.2010, Philip Butterill leg., Syzygium decipiens (FSCA 00090423). Paratypes: Papua New Guinea: 3♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 03.XII.2010, Philip Butterill leg., Syzygium decipiens (FSCA 00094479–00094481).

Synopeas psychotriae Buhl, 2013

Description

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.

Head. Shape of head in anterior view: circular; slightly ovoid. Central keel: complete and weakly developed. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: minute rugulae; reticulate microsculpture. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Hyperoccipital carina: complete. Hyperoccipital carina strength: uniformly weak. Distance between lateral ocellus and hyperoccipital carina: less than 1 ocellar diameter; approximately 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally. Lateral pronotal sculpture coverage: 1/4 to 1/2. Mesoscutellar spine: short and pointed. Mesoscutellar spine in lateral view: pointing posteriorly; angled posterodorsally. Posterior margin of lateral propodeal carina in lateral view: pointed dorsally and ventrally. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: well impressed. Setation of mesoscutum: sparse. Mesoscutal lamella: broad and square. Setation of mesoscutellum: laterally dense, medially sparse.

Figure 37. 

Synopeas psychotriae (male specimen; FSCA 00090261) A lateral habitus B dorsal habitus C ventral habitus.

Metasoma. Microsculpture of S2: narrow band at posterior margin. Sculpture of T2: absent; narrow transverse band of microsculpture at posterior margin. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: shorter than distance between setal bases. Density of setae on disc of forewing: sparse. Arrangement of setae on disc of forewing: setose distally, glabrous proximally. Forewing marginal setae: approximately uniform in length on anterior and posterior margins.

Diagnosis

The parapsidal lines of S. psychotriae are well impressed (Fig. 20A), distinguishing it from the similar S. luli, which has unmarked or very faint parapsidal lines. The metasomal microsculpture of S. psychotriae is minimal, being absent or restricted to very narrow bands at the posterior margins of each segment, whereas S. sanga, S. occultum, and S. kira have more extensive metasomal sculpturing (Fig. 6). There is no central keel, but rather an impressed line running from the toruli to the median ocellus. The mesoscutellar spine may be angled posteriorly or posterodorsally.

Comments

The female types of S. psychotriae have a posterodorsally pointed mesoscutellar spine. The male specimen has a posteriorly pointed mesoscutellar spine, while that of the female from the same gall is posterodorsally pointed. The angle of the mesoscutellar spine may be sexually dimorphic in this species, but more specimens are needed for confirmation.

Plant associations

Reared from galls on Psychotria ramuensis (Rubiaceae).

Material examined

Holotype: Papua New Guinea: ♀, Madang, 5°08'S, 145°47'E, 27.I–5.IV.2011, Philip Butterill leg., Psychotria ramuensis (USNMENT01335972). Paratype: Papua New Guinea: ♀, Madang, 5°08'S, 145°47'E, 27.I–5.IV.2011, Philip Butterill leg., Psychotria ramuensis (USNMENT01335982). Other material: Papua New Guinea: ♀ ♂, Morobe, Ohu, 5.233°S, 145.686°E, 22.II.2011, Philip Butterill leg., Psychotria ramuensis, (FSCA 00090261, 00094619).

Synopeas pterocarpi Buhl, 2013

Description

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.

Head. Shape of head in anterior view: circular. Central keel: absent. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: parallel, arched rugae. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: evenly spaced. Shape of mandible: unknown. Distance between lateral ocellus and compound eye (OOL): approximately 1 ocellar diameter. Hyperoccipital carina: complete. Hyperoccipital carina strength: uniformly robust. Distance between lateral ocellus and hyperoccipital carina: uncertain, approximately 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally. Lateral pronotal sculpture coverage: 1/2 to 2/3. Mesoscutellar spine: absent or inconspicuous. Posterior margin of lateral propodeal carina in lateral view: rounded. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: sparse. Mesoscutal lamella: broad and rounded. Setation of mesoscutellum: laterally dense, medially sparse.

Metasoma. Microsculpture of S2: unknown. Sculpture of T2: absent. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: shorter than distance between setal bases. Density of setae on disc of forewing: sparse. Arrangement of setae on disc of forewing: unknown. Forewing marginal setae: unknown.

Diagnosis

The frons of S. pterocarpi is characterized by parallel, arched rugae above the toruli, not found in other Synopeas from New Guinea.

Plant associations

Reared from galls on Pterocarpus indicus (Fabaceae).

Material examined

Holotype: Papua New Guinea: ♀, Madang, Ohu, 5°14'S, 145°41'E, 13.IX.2010–19.I.2011, Philip Butterill leg., Pterocarpus indicus (USNMENT01335974). Paratype: Papua New Guinea: ♀, Madang, Ohu, 5°14'S, 145°41'E, 13.IX.2010–19.I.2011, Philip Butterill leg., Pterocarpus indicus, ZMUC.

Figure 38. 

Holotype of Synopeas pterocarpi (USNMENT01335974) A lateral habitus B dorsal habitus C anterior head D dorsolateral habitus. Scale bar: 0.1 mm.

Synopeas pulupulu Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: circular. Central keel: absent. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: minute rugulae; reticulate microsculpture. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Hyperoccipital carina: complete. Hyperoccipital carina strength: uniformly robust. Distance between lateral ocellus and hyperoccipital carina: approximately 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally. Lateral pronotal sculpture coverage: 2/3 to 3/4. Mesoscutellar spine: short and pointed. Mesoscutellar spine in lateral view: pointing posteriorly; angled posterodorsally. Posterior margin of lateral propodeal carina in lateral view: pointed dorsally and ventrally. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: moderately dense. Mesoscutal lamella: roughly triangular. Setation of mesoscutellum: present throughout.

Figure 39. 

Synopeas pulupulu (female holotype; FSCA 00090430) A lateral habitus B dorsal habitus C ventral habitus.

Metasoma. Microsculpture of S2: absent. Sculpture of T2: absent. Length of T2: approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: longer than distance between setal bases. Density of setae on disc of forewing: dense. Arrangement of setae on disc of forewing: uniformly setose distally, proximally glabrous with linea setosa. Forewing marginal setae: approximately uniform in length on anterior and posterior margins.

Diagnosis

Synopeas pulupulu has a short, pointed mesoscutellar spine and a robust, complete hyperoccipital carina. Synopeas pulupulu can be differentiated from S. roncavei and S. pattiae by the setation of the meoscutum and mesoscutellum, which are uniformly setose in S. pulupulu and medially glabrous in S. roncavei and S. pattiae. Additionally, the sculpture of the lateral pronotum in S. pulupulu is more extensive than that of S. roncavei and S. pattiae, covering more than two thirds of the sclerite.

Etymology

The epithet “pulupulu” is Manam for “hairy” (Osmond and Ross 2016) and refers to the relatively dense setae on the mesoscutum. The name is to be treated as a noun in apposition.

Plant associations

Reared from galls on Palmeria sp. (Monimiaceae) [GALL331].

Material examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 20.XI.2010, Philip Butterill leg., Palmeria sp. 1 (FSCA 00090430). Paratype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 20.XI.2010, Philip Butterill leg., Palmeria sp. 1 (FSCA 00094459).

Figure 40. 

Synopeas roncavei (female holotype; FSCA 00090249) A lateral habitus B dorsal habitus C ventral habitus.

Synopeas roncavei Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: circular. Central keel: absent. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: minute rugulae; reticulate microsculpture. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Hyperoccipital carina: complete. Hyperoccipital carina strength: uniformly robust. Distance between lateral ocellus and hyperoccipital carina: approximately 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally. Lateral pronotal sculpture coverage: 1/2 to 2/3. Mesoscutellar spine: short and pointed. Mesoscutellar spine in lateral view: pointing posteriorly. Posterior margin of lateral propodeal carina in lateral view: rounded. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: absent medially, present laterally. Mesoscutal lamella: broad and rounded; roughly triangular. Setation of mesoscutellum: laterally dense, medially sparse.

Metasoma. Microsculpture of S2: narrow band at posterior margin. Sculpture of T2: narrow transverse band of microsculpture at posterior margin. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: longer than distance between setal bases. Density of setae on disc of forewing: dense. Arrangement of setae on disc of forewing: uniformly setose distally, proximally glabrous with linea setosa. Forewing marginal setae: setae on posterior margin distinctly longer than setae on anterior margin.

Diagnosis

Synopeas roncavei has a short, pointed mesoscutellar spine and a robust, complete hyperoccipital carina. The metasomal sculpture on T2 is restricted to a narrow band at the posterior margin (Fig. 40). It is similar to S. kiki, and these can be differentiated by the hyperoccipital carina. In S. kiki, the hyperoccipital carina is medially weakened and laterally present as a thin carina, whereas in S. roncavei, the hyperoccipital carina is uniformly robust. Synopeas roncavei can be differentiated from S. pulupulu by the setation of the meoscutum and mesoscutellum, which are medially glabrous in S. roncavei and uniformly setose in S. pulupulu. Synopeas roncavei is similar to S. pattiae, but the forewing of S. pattiae lacks a linea setosa, which is present in S. roncavei (Fig. 23).

Etymology

This species is named in honor of the biological control researcher and beetle taxonomist Ronald D. Cave.

Plant associations

Primarily reared from blister galls on Piper amboinense and Piper celtidiforme (Piperaceae) [GALL273, GALL300]. One specimen reared from Cypholophus friesianus (Urticaceae) [GALL266].

Material examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 12.X.2010, Philip Butterill leg., Piper celtidiforme (FSCA 00090249). Paratypes: Papua New Guinea: ♀ ♂, Morobe, Yawan, 20.IX.2010, Piper amboinense (FSCA 00000407, 00000322); ♂, Morobe, Yawan, 05.X.2010, Piper sp. (FSCA 00090417); 2♀, Morobe, Yawan, 12.X.2010, Piper celtidiforme, FSCA 00094605–00094606); 4♀, Morobe, Yawan, 15.X.2010, Piper amboinense (FSCA 00000339–00000342); ♀, Morobe, Yawan, 15.X.2010, Cypholophus friesianus (FSCA 00090432); ♂, Morobe, Yawan, 18.X.2010, Piper amboinense (FSCA 00000408); 3♀ 3♂, Morobe, Yawan, 25.X.2010, Piper amboinense (FSCA 00090259, 00090411, FSCA 00094495–00094496, 00094607–00094608); ♂, Morobe, Yawan, 11.XI.2010, Piper amboinense (FSCA 00000253).

Synopeas sanga Awad, sp. nov.

Description

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.

Figure 41. 

Synopeas sanga (female holotype; FSCA 00000399) A lateral habitus B dorsal habitus C ventral habitus.

Head. Shape of head in anterior view: slightly ovoid. Central keel: absent. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: minute rugulae; reticulate microsculpture. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: evenly spaced. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Hyperoccipital carina: complete. Hyperoccipital carina strength: uniformly robust. Distance between lateral ocellus and hyperoccipital carina: less than 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally. Lateral pronotal sculpture coverage: 1/2 to 2/3. Mesoscutellar spine: short and pointed. Mesoscutellar spine in lateral view: pointing posteriorly. Posterior margin of lateral propodeal carina in lateral view: pointed ventrally. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: sparse. Mesoscutal lamella: long and narrow. Setation of mesoscutellum: laterally dense, medially sparse.

Metasoma. Microsculpture of S2: present at posterior margin and in posterolateral corners. Sculpture of T2: wide transverse band of microsculpture at posterior margin. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: longer than distance between setal bases. Density of setae on disc of forewing: dense. Arrangement of setae on disc of forewing: uniformly setose distally, proximally glabrous with linea setosa. Forewing marginal setae: setae on posterior margin distinctly longer than setae on anterior margin.

Diagnosis

Synopeas sanga has a short, pointed scutellar spine and a robust, complete hyperoccipital carina. It may be distinguished from similar species by the patch of microsculpture on posterior and posterolateral S2 (Fig. 6A) and the long band of reticulate microsculpture on dorsal T2 (Fig. 7A). It resembles S. occultum but differs in the length of T2, which is much more elongate in S. occultum (compare Figs 35 and 41). Synopeas sanga has sparser setation on the frons than does S. roncavei sp. 2 (compare Figs 40 and 41).

Etymology

The epithet “sanga” is Proto-Oceanic for “forked” (Evans, 2008) and refers to the forked microsculpture pattern on posterior S2. The name is to be treated as a noun in apposition.

Plant associations

Reared from blister galls on Saurauia conferta and Saurauia schumanniana (Actinidiaceae) [GALL397, GALL499].

Material examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 03.IX.2012, Philip Butterill leg., Saurauia conferta (FSCA 00000399). Paratypes: Papua New Guinea: 3♀ 2♂, Morobe, Yawan, 17.V.2012, Saurauia conferta (FSCA 00000345–00000349); 2♀, Morobe, Yawan, 03.IX.2012, Saurauia schumanniana (FSCA 00000397, 00094474); 7♀ 3♂, Morobe, Yawan, 03.IX.2012, Saurauia conferta (FSCA 00094511–00094514, 00094532–00094536, 00094615); 3♀ 3♂, Morobe, Yawan, 05.XI.2012, Saurauia conferta, (FSCA 00000356–00000361); ♂, Morobe, Yawan, 20.XI.2012, Saurauia conferta (FSCA 00000396).

Synopeas toto Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: circular. Central keel: complete and well developed. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: uncertain, minute rugulae; uncertain, reticulate microsculpture. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: medial setae closer to each other than to lateral setae. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Hyperoccipital carina: complete. Hyperoccipital carina strength: uniformly weak. Distance between lateral ocellus and hyperoccipital carina: approximately 1 ocellar diameter.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally. Lateral pronotal sculpture coverage: 1/2 to 2/3. Mesoscutellar spine: short and pointed. Mesoscutellar spine in lateral view: angled posterodorsally. Posterior margin of lateral propodeal carina in lateral view: pointed ventrally. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: sparse. Mesoscutal lamella: roughly triangular. Setation of mesoscutellum: absent medially.

Figure 42. 

Synopeas toto A lateral habitus (female paratype; FSCA 00000245) B dorsal habitus (female paratype; FSCA 00090421) C ventral habitus (female paratype; FSCA 00090421).

Metasoma. Microsculpture of S2: narrow band at posterior margin. Sculpture of T2: narrow transverse band of microsculpture at posterior margin. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: shorter than distance between setal bases. Density of setae on disc of forewing: sparse. Arrangement of setae on disc of forewing: densely setose distally, sparsely setose proximally. Forewing marginal setae: setae on posterior margin distinctly longer than setae on anterior margin.

Diagnosis

Synopeas toto has a strong, distinct central keel on the frons. Synopeas psychotriae also has a line on the frons that extends from between the toruli to the median ocellus, but it is impressed rather than raised. Additionally, the parapsidal lines of S. psychotriae are well impressed (Fig. 20A) and are very weak in S. toto (Fig. 42). Synopeas butterilli can have a complete central keel, but it is weakly developed. The sculpture of the lateral pronotum in S. butterilli is restricted to a narrow medial band (Fig. 27), rather than covering the dorsal half of the lateral pronotum as in S. toto.

Etymology

The epithet “toto” is Proto-Oceanic for “cut” (Ross et al. 1998) and refers to the complete central keel dividing the frons. The name is to be treated as a noun in apposition.

Plant associations

Reared from galls on Nauclea (Rubiaceae) [GALL217, GALL218].

Specimens examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 01.XII.2010, Philip Butterill leg., Nauclea sp. 1 (FSCA 00000252). Paratypes: Papua New Guinea: ♀♂, Morobe, Yawan, 11.XI.2010, Nauclea sp. 1 (FSCA 00090421, 00094597); 2♂, Morobe, Yawan, 25.XI.2010, Nauclea sp. 1 (FSCA 00090420, 00094494); ♂, Morobe, Yawan, 26.XI.2010, Nauclea sp. 1, FSCA 00000412); ♀, Morobe, Yawan, 01.XII.2010, Nauclea sp. 1 (FSCA 00000330); ♀ 2♂, Morobe, Yawan, 26.I.2011, Nauclea sp. 1 (FSCA 00000406, 00094592–00094593); 5♀ 4♂, Morobe, Yawan, 21.XI.2012, Nauclea sp. 1 (FSCA 00000245, 00000258, 00000336–00000338, 00000393, 00094563–00094565).

Synopeas valavala Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: triangular. Central keel: absent. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: reticulate microsculpture. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: evenly spaced. Shape of mandible: bidentate. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Hyperoccipital carina: absent. Hyperoccipital carina strength: unknown. Distance between lateral ocellus and hyperoccipital carina: unknown. Postgenal region: medially impressed.

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally, smooth band anterior to tegula. Lateral pronotal sculpture coverage: 1/2 to 2/3. Mesoscutellar spine: absent or inconspicuous. Posterior margin of lateral propodeal carina in lateral view: rounded. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: moderately dense. Mesoscutal lamella: short, projecting less than its width beyond margin. Setation of mesoscutellum: laterally dense, medially sparse.

Metasoma. Microsculpture of S2: narrow band at posterior margin. Sculpture of T2: uncertain, absent. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: longer than distance between setal bases. Density of setae on disc of forewing: dense. Arrangement of setae on disc of forewing: uniformly setose distally, proximally glabrous with linea setosa. Forewing marginal setae: setae on posterior margin distinctly longer than setae on anterior margin.

Diagnosis

Synopeas valavala has an exceptionally large, triangular head, unlike any other known species of Synopeas. The malar space is long, about equal to the height of the compound eye, and the mesoscutellar spine is inconspicuous to absent.

Comments

A similar head shape is found in Inostemma macarangae Buhl, 2013 (Fig. 44), which has also been reared from Macaranga galls, suggesting convergent evolution. There is no central keel, but the angle of the frons may give the impression of a keel when viewed at certain angles.

Figure 43. 

Synopeas valavala A lateral habitus (female holotype; FSCA 00033890) B dorsal habitus (female paratype; FSCA 00000350) C ventral habitus (female holotype; FSCA 00033890).

Etymology

The epithet “valavala” is Maenge for “Macaranga” (Ross, 2008) and refers to the identity of the gall host plant. The name is to be treated as a noun in apposition.

Plant associations

Reared from round, fuzzy galls on Macaranga strigosa (Euphorbiaceae) [GALL357].

Material examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 21.XI.2012, Philip Butterill leg., Macaranga strigosa (FSCA 00033890). Paratypes: Papua New Guinea: 2♀, Morobe, Yawan, 21.XI.2012, Philip Butterill leg., Macaranga strigosa (FSCA 00000259–00000350).

Synopeas zhangi Awad, sp. nov.

Description

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.

Head. Shape of head in anterior view: distinctly ovoid. Central keel: absent; partial. Sculpture on frons: reticulate microsculpture. Epitorular sculpture: minute rugulae; reticulate microsculpture. Number of clypeal setae: 4. Length of median pair of clypeal setae: longer than lateral pair. Arrangement of clypeal setae: uncertain, medial setae closer to lateral setae than to each other. Shape of mandible: unidentate. Distance between lateral ocellus and compound eye (OOL): less than 1 ocellar diameter. Hyperoccipital carina: absent.

Figure 44. 

Inostemma macarangae (male specimen; FSCA 00090262) A lateral habitus B dorsal habitus C anteroventral habitus.

Figure 45. 

Synopeas zhangi A lateral habitus (female paratype; FSCA 00000291) B dorsal habitus (female paratype; FSCA 00000383) C ventral habitus (female paratype; FSCA 00000383).

Mesosoma. Epomial carina: present, complete or nearly so. Microsculpture of lateral pronotum: present dorsally, absent ventrally. Lateral pronotal sculpture coverage: 1/4 to 1/2. Mesoscutellar spine: absent or inconspicuous. Posterior margin of lateral propodeal carina in lateral view: pointed dorsally and ventrally. Mesosomal dorsum in lateral view: convex. Notauli: unmarked or faintly suggested. Parapsidal lines: very faint. Setation of mesoscutum: sparse. Mesoscutal lamella: roughly triangular. Setation of mesoscutellum: laterally dense, medially sparse.

Metasoma. Microsculpture of S2: narrow band at posterior margin. Sculpture of T2: absent; narrow transverse band of microsculpture at posterior margin. Length of T2: shorter than mesosoma; approximately as long as mesosoma.

Wing. Length of setae on disc of forewing: shorter than distance between setal bases. Density of setae on disc of forewing: sparse. Arrangement of setae on disc of forewing: setose distally, glabrous proximally. Forewing marginal setae: setae on posterior margin distinctly longer than setae on anterior margin.

Diagnosis

Synopeas zhangi may be recognized by its head shape, which is distinctly wider than high, the unidentate mandible, and the absence of a hyperoccipital carina. The dorsal mesosoma is moderately convex, allowing it to be easily distinguished from an undescribed species known from a single male specimen, which has an extremely convex mesosoma.

Etymology

This species is named in honor of the hymenopterist Yuanmeng Miles Zhang.

Plant associations

Reared from galls on Nauclea spp. (Rubiaceae) [GALL218, GALL219, GALL222], Macaranga strigosa (Euphorbiaceae) [GALL357], Ficus adenosperma (Moraceae) [GALL318], and Elaeocarpus dolichostylus (Elaeocarpaceae) [GALL241].

Material examined

Holotype: Papua New Guinea: ♀, Morobe, Yawan, 6.14141°S, 146.87415°E, 26.I.2011, Philip Butterill leg., Macaranga strigosa (FSCA 00090250). Paratypes: Papua New Guinea: ♂, Morobe, Yawan, 15.XI.2010, Nauclea sp. 1 (FSCA 00000367); ♂, Morobe, Yawan, 16.XI.2010, Nauclea sp. 1 (FSCA 00090407); ♂, Morobe, Yawan, 01.I.2011, Elaeocarpus dolichostylus (FSCA 00090422); ♀ 4♂, Morobe, Yawan, 24.I.2011, Nauclea sp. 1 (FSCA 00033887, 00094611–00094614); 3♀ 9♂, Morobe, Yawan, 25.I.2011, Nauclea tenuiflora (FSCA 00000291, 00000370–00000373, 00000383–00000384, 00033886, 00094475–00094478); ♀, Morobe, Yawan, 25.I.2011, Ficus adenosperma (FSCA 00090424); 1♀ 4♂, Morobe, Yawan, 26.I.2011, Macaranga strigosa (FSCA 00090252, 00094472–00094473, 00094497, 00094582); ♀ ♂, Morobe, Yawan, 26.I.2011, Nauclea sp. 1 (FSCA 00000324, 00000353); ♂, Morobe, Yawan, 08.X.2012, Nauclea tenuiflora (FSCA 00000387).

CO1 Results

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 46, 47). Some clusters were strongly supported. Synopeas sanga, S. occultum, S. kira, and the specimen from the UK formed a cluster with over 99% support. Synopeas amandae and S. codex formed a cluster with over 97% support. A large and well-supported cluster (over 97%) included S. zhangi, S. kiki, S. toto, S. luli, S. psychotriae, S. pattiae, and S. butterilli. Synopeas klingunclum, S. anunu, and the specimen from Myanmar formed a cluster with over 75% support in the neighbor-joining analysis.

The New Guinea material did not all cluster together. Synopeas specimens from the US, the UK, and Myanmar were placed relatively far apart from one another. The US and Myanmar specimens clustered with S. klingunculum and S. anunu. As stated above, the UK specimen clustered with S. sanga, S. occultum, and S. kira, although it was distinct from the New Guinea species. The specimen from the UK was tentatively identified as S. larides (Walker) by comparison to a paralectotype specimen photographed by Elijah Talamas.

The CO1 sequences of three species, S. luli, S. sanga, and S. zhangi, had 99–100% matches with undetermined species of Synopeas collected in South Korea. The plant genus from which S. sanga was reared, Saurauia, is widespread in tropical and subtropical Asia. The plant species from which S. luli was reared, Neuburgia corynocarpa and Piper macropiper, are distributed from Taiwan to the western Pacific (POWO 2020). Synopeas zhangi was reared from diverse plant taxa and may be a generalist.

We recognize the possibility that these species are widespread and may have already been described. However, Synopeas species from the Palearctic cannot be reliably identified without a thorough taxonomic treatment, a task far beyond the scope of this project. In the event that these species have been described before, we believe we have made it straightforward to treat them as junior synonyms.

Discussion

Some morphological characters formed discernible patterns. The cluster including S. klingunculum, S. anunu, and the specimen from Myanmar was morphologically identifiable. The notauli of species in this cluster are distinctly grooved, and there is striate sculpture on the mesoscutal lamella and medial mesoscutellum. Therefore, future analysis may find that the grooved notauli identify a distinct species group within Synopeas. Similar notauli are also found in Inostemma, an early-diverging group, which could mean that grooved notauli represent the plesiomorphic state for Platygastrinae. Masner and Huggert (1989) applied the same polarity to this character at the genus and subfamily levels.

Other small clusters have more subtle morphological similarities. Synopeas zhangi and an undescribed species with similar CO1 are characterized by a distinctly ovoid head in frontal view and a unidentate mandible. All other Synopeas species in the analysis have a bidentate mandible. Synopeas amandae and S. codex both have the lateral pronotum with microsculpture throughout, whereas the other species in the analysis have the lateral pronotum only partially sculptured. Synopeas sanga, S. occultum, and S. kira have more extensive sculpturing on the metasoma than do most other species treated here, although this character is also present in S. codex, which is not closely related.

Figure 46. 

Neighbor-joining tree of Synopeas 5’-CO1 barcodes. Terminal taxa are labeled to their narrowest identification level. Numbers in parentheses after terminal taxa indicate how many sequences are represented in each cluster. Bootstrap support values greater than 75% are indicated on the tree.

Figure 47. 

Maximum likelihood tree of Synopeas 5’-CO1 barcodes. Colored boxes indicate gall host plant genera from which multiple specimens were barcoded.

Of the 19 Synopeas species with known plant associations, twelve were reared from a single gall host and seven from at least two gall hosts. From the latter, five species came from galls on two or more plant genera. Thus we can report evidence that some Synopeas species use multiple host galls and plants, and that individual gall types (morphospecies) can support multiple platygastrid species.

For example, the cluster of five gall types formed on Nauclea spp. (Rubiaceae) supported 8 species of Synopeas, and at least one, but likely several, species of other platygastrine genera. The most structurally complex gall (a globoid leaf gall, GALL218) supported five Synopeas species. Only S. anunu, S. kalubia, and S. kiki were reared from a single gall type. The others were found in at least two Nauclea gall types, and both S. zhangi and S. codex were reared from galls formed on different plant genera (7 and 6 gall types, respectively, including the Nauclea galls).

The number of reared parasitoids is directly related to the number of host galls reared, and the Nauclea galls were by far the most abundant and frequently reared galls throughout the project, which may in part explain the apparently rich platygastrid fauna associated with them. Despite this, the emerging picture does suggest a richer, more complex, and more generalised community associated with galls than perhaps was expected. Additionally, it is possible that some Synopeas species are associated with cecidomyiid inquilines or predators rather than with the gall formers themselves.

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 Synopeas diversity.

Acknowledgements

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 (ZMUC), Frederique Bakker (RMNH), Zoltán Vas (HMNH), and Derek Binns. We thank Jackie Miller and Debbie Matthews (McGuire Center for Lepidoptera and Biodiversity) for providing access to imaging resources. We thank all field assistants from Yawan and Madang, and their surrounding villages, especially Gibson Maiah, the field lab assistant to Phil Butterill in Yawan. Thanks also to the staff and para-ecologists of the New Guinea Binatang Research Centre, Madang, and Inga Freiberga of the Czech Academy of Sciences for lab work in Czech Republic. We thank Vojtech Novotny and George Weiblen for project supervision and plant identifications, respectively. Elijah Talamas, Jonathan Bremer, and Matthew Moore were supported by FDACS-DPI. Philip Butterill was supported by European Research Council grant 669609.

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Supplementary materials

Supplementary material 1 

List of specimens and their depositories

Jessica Awad, Jonathan S. Bremer, Philip T. Butterill, Elijah J. Talamas

Data type: material examined

Explanation note: List of specimens examined, depositories, type status, and image links. Link to holotype image repository: https://zenodo.org/communities/platygastroidea_primary_types/

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (87.00 kb)
Supplementary material 2 

CO1 data in Genbank

Jessica Awad, Jonathan S. Bremer, Philip T. Butterill, Elijah J. Talamas

Data type: COI data

Explanation note: List of barcode vouchers, internal identifiers, and Genbank accession numbers. Link to PNG image repository: https://zenodo.org/communities/synopeas_from_png/

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (43.00 kb)