Biodiversity Data Journal : Single Taxon Treatment
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Single Taxon Treatment
Gryon ancinla Kozlov & Lê (Hymenoptera: Scelionidae): host association, expanded distribution, redescription and a new synonymy
expand article infoHuayan Chen, Elijah J Talamas§,|, Marie-Claude Bon, Matthew R. Moore#
‡ State Key Laboratory of Biocontrol, School of Life Sciences, Sun Yat-sen University, Guangzhou, China
§ Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, Gainesville, United States of America
| Systematic Entomology Laboratory, United States Department of Agriculture, Washington, DC, United States of America
¶ USDA-ARS-EBCL, Montpellier, France
# Florida Department of Agriculture and Consumer Services, Gainesville, FL, United States of America
Open Access

Abstract

Background

Gryon Haliday (Platygastroidea: Scelionidae) is a cosmopolitan genus of egg-parasitoid wasps primarily associated with Heteroptera.

New information

Gryon ancinla Kozlov & Lê is reported for the first time outside of Vietnam, in China and Cambodia, and as an egg parasitoid of the pestiferous leaf-footed bug, Acanthocoris scaber (L.). Gryon ancinla is redescribed based on recently collected specimens and compared to closely related species of Gryon in the region. Gryon clavaerus Kozlov & Lê is treated as a junior synonym and some characters found in the charon species group are discussed.

Introduction

Acanthocoris scaber (L.) (Hemiptera: Coreidae) is a pest that feeds on many economically important vegetables including chili, potato, tomato, and eggplant in China (Du et al. 2014), where it has become a serious problem for chili production in recent years. Adults and nymphs usually aggregate on leaves, stems and fruits (Fig. 1). They feed by sucking plant fluids that contain sugars and other nutrients, resulting in production losses. A recent survey conducted in China to assess parasitism of A. scaber eggs yielded numerous specimens of Gryon Haliday (Scelionidae), a cosmopolitan genus of parasitoid wasps whose species primarily parasitize the eggs of Heteroptera (Johnson et al. 1996, Masner 1983).

Figure 1.  

Nymphs and adults of Acanthocoris scaber feeding on a chili plant in Xiangtoushan, China.

The taxonomy of Gryon in southeast Asia is in need of thorough revision, as in much of the world. The conspicuous characters of the charon species group, to which G. ancinla belongs, enabled us to consider a limited number of species names when identifying the specimens. We considered the possibility that some species may have ranges that extend from Asia into Africa. Indeed, G. ancinla is similar to some African Gryon species but the molecular data at hand indicate that it is distinct from the African species in the current analysis. We assign the name G. ancinla to these specimens because we are confident that their morphology matches that of the holotype specimen and they are from the same geographic region. There remain a number of Asian species in the charon species group for which we have not yet examined primary types, and examination of more material is required to fully assess the morphological variation and geographic range of G. ancinla. This study should thus be considered as one of many steps forward in the advancement of the taxonomy of Gryon and the reader should be aware that future work will undoubtedly result in more nomeclatural adjustments.

The charon species group was proposed by Mineo 1983 for species with the frontal depression surrounded by a robust carina (Figs 2a, b, 4). Mineo also used other characters to define this group, many of which are located on the posterior head. While characters of the posterior head can be useful, their observation may require removal of the head from the mesosoma and it is not permissible to do with many primary type specimens. Mineo 1990 erected another species group, the letus group, listing only characters that this cluster shares with the charon group and did not provide characters to separate these groups. We consider this to be an unnecessary separation and, for simplicity, consider species with a frontal depression surrounded by a robust carina to be part of the charon group. A list of these species, while not exhaustive, is provided in Table 3 to facilitate meaningful comparisons between species. Phylogenetic analysis of Gryon is currently underway and we prefer not to further define any species groups without the context of a larger study. We also emphasize the use of characters that can be seen without dissection so that recently collected material can be matched to primary types. In Gryon there are many such characters to be exploited and during this study we found two that we have not previously encountered in taxonomic literature on the genus.

Table 1.

CO1 barcode K2P distances among species in the Gryon charon group.

Gryon Species Comparison

Range of Interspecific K2P Distances (%)

G. ancinla-G. drunoris

12.9–14.8

G. ancinla-G. sp. (BOLD:ADO2077)

14.8–16.7

G. ancinla-G. sp. 1 charon group

11.4–13.7

G. ancinla-G. sp. 2 charon group

14.0–14.6

G. drunoris-G. sp. (BOLD:ADO2077)

15.7–16.1

G. drunoris-G. sp. 1 charon group

14.6–14.8

G. drunoris-G. sp. 2 charon group

16.2–16.6

G. sp. 1 charon group-G. sp. (BOLD:ADO2077)

16.4–16.6

G. sp. 1 charon group-G. sp. 2 charon group

15.0

G. sp. 2 charon group-G. sp. (BOLD:ADO2077)

16.2

Table 2.

Specimens used in the molecular analysis

Species Origin Genbank Accession Collecting Unit Identifier Specimen Depository DNA Depository
Gryon ancinla China MT604053 FSCA 00094670 FSCA FSCA
Gryon ancinla China MT604054 FSCA 00094672 FSCA FSCA
Gryon ancinla China MT604055 FSCA 00094673 FSCA FSCA
Gryon ancinla China MT604056 USNMENT01335659 FSCA EBCL
Gryon ancinla China MT604057 USNMENT01335640 FSCA EBCL
Gryon ancinla China MT604058 SCAU 3017206 SYSU SYSU
Gryon ancinla China MT604059 FSCA 00094671 FSCA FSCA
Gryon drunoris Vietnam MT604060 FSCA 00094674 FSCA FSCA
Gryon drunoris Vietnam MT604061 FSCA 00094680 FSCA FSCA
Gryon drunoris Vietnam MT604062 FSCA 00094682 FSCA FSCA
Gryon drunoris Vietnam MT604063 FSCA 00094684 FSCA FSCA
Gryon sp. 1, charon group South Africa MT604064 SAM-HYM-P093268 SAMC FSCA
Gryon sp. 1, charon group South Africa MT604065 SAM-HYM-P093626 SAMC FSCA
Gryon sp. 2, charon group South Africa MT604066 SAM-HYM-P093671 SAMC FSCA
Table 3.

Species of Gryon in which the frontal depression is surrounded by carinae.

Species Basis of determination
G. ancinla Kozlov & Lê Examination of holotype
G. charon (Nixon) Examination of holotype
G. drunoris Kozlov & Lê Examination of holotype
G. dasyni (Nixon) Nixon 1934
G. hakonense (Ashmead) Examination of holotype
G. ingens Veenakumari & Rajmohana Kamalanathan et al. 2016
G. kenyotum Mineo Mineo 1982
G. krishnagiriense Sharma Examination of holotype
G. letus (Nixon) Examination of holotype
G. lucmon Mineo Examination of holotype
G. mudugeriense Sharma Examination of holotype
G. nigriclavatum (Dodd) Examination of holotype
G. odontogonusi (Risbec) Examination of lectotype
G. oophagum (Nixon) Nixon 1934
G. paracharontis Mineo Examination of holotype
G. parakenyotum Mineo Examination of holotype
G. philippinense (Ashmead) Examination of lectotype
G. sponus Kozlov & Lê Examination of holotype
G. urum Mineo Mineo 1982
Figure 2.

Gryon ancinla Kozlov & Lê, holotype female, images provided by Sergey Belokobylskij.

ahead, anterior view  
bhead and mesosoma, anterolateral view  
chabitus, lateral view  
dhabitus, dorsal view  
Figure 3.

Gryon clavaerus Kozlov & Lê, holotype female (IEBR 0170).

ahead, anterior view  
bhead and mesosoma, dorsal view  
chead and mesosoma, lateral view, image flipped horizontally.  
dhabitus, lateral view, image flipped horizontally  
Figure 4.  

Gryon ancinla, female (FSCA 00090950), head and mesosoma, ventrolateral view.

Figure 5.

Gryon sponus (IEBR 0138), holotype female.

ahabitus, lateral view  
bhead and mesosoma, lateral view  
Figure 6.

Gryon drunoris.

aholotype female (IEBR 0176), habitus, lateral view  
bfemale (FSCA 00094680), head and mesosoma, anterolateral view  
Figure 7.

Comparison of length of T1 and T3.

aGryon ancinla (FSCA 00090590), female, dorsolateral view  
bGryon drunoris (FSCA 00094680), female, dorsal view  
Figure 8.

Gryon ancinla, arrows indicate a pit at the junction of the mesoscutal suprahumeral sulcus and mesoscutal humeral sulcus.

afemale (FSCA 00090590), head and mesosoma, dorsolateral view  
bfemale (SCAU 3040175), mesosoma, dorsolateral view  
Figure 9.

Gryon letus, holotype female (NHMUK 013378986).

ahead, anteroventral view, image flipped horizontally  
bhead, dorsolateral view, image flipped horizontally  

Materials and methods

Cybertaxonomy

The description of G. ancinla was generated using vSysLab (vsyslab.osu.edu), an online, matrix-based tool for generating species descriptions.

Imaging

Specimens SCAU 3040175–3040176 were photographed using CombineZP and AutoMontage extended-focus software, using a JVC KY-F75U digital camera, Leica Z16 APOA microscope and 1X objective lens. Specimens were photographed using a Macroscopic Solutions Macropod using a 20X Mitutoyo objective lens with images rendered using Helicon Focus. Scanning electron micrographs were produced using Phenom Pro Desktop SEM and aligned in Helicon focus.

Collections

Comprehensive identification tools are not presently available for Gryon in southeast Asia. Our determination of G. ancinla was made through direct comparison with holotypes and images of holotypes in the Institute of Ecology and Biological Resources (IEBR) (Hanoi, Vietnam) (Talamas and Pham 2017), The National Museum of Natural History (Washington, DC, USA) (Talamas et al. 2017), The Canadian National Collection of Insects (Ottawa, Canada), The Natural History Museum (NHMUK) (London, UK), Museo Civico di Storia Naturale Giacomo Doria (Genoa, Italy), National Museum of Natural History (Paris, France) and the Zoological Institute (St. Petersburg, Russia). The taxonomic and molecular portions of this study were based on material from the California Department of Food and Agriculture (CDFA), Florida State Collection of Arthropods, Gainesville, Florida (FSCA), Sun Yat-sen University, Guangzhou, China (SYSU), the Iziko Museum of Cape Town, South Africa (SAMC), the European Biological Control Laboratory, Montpellier, France (EBCL), and the College for Agriculture and Life Sciences, Seoul National University, Seoul, Korea (SNU).

CO1 barcoding

Genomic DNA was nondestructively isolated from the entire specimen using a Qiagen DNeasy Blood and Tissue kit (Hilden, Germany). The barcode region of the mitochondrial Cytochrome c Oxidase Subunit I (CO1) was amplified using the universal barcoding primer sets LCO1490/HCO2198 (Folmer et al. 1994) or LEPF1/FEPR1 (Hebert et al. 2004). Amplification and sequence editing were done at EBCL as described in Sabbatini Peverieri et al. 2018. At FSCA, PCRs used the following thermocycle: 1) initial denaturation at 950C for 2 minutes, 2) 980C for 30 seconds, 3) 500C for 30 seconds, 4) 720C for 40 seconds [32x steps 2-4], and a final extension at 720C for 7 minutes. Samples at FSCA were sequenced bidirectionally on the ABI SeqStudio platform with BigDye v3.1 chemistry. Sequences were trimmed and assembled into contigs using Sequencher 5.4.6.

All sequences generated from this study are deposited in GenBank (MT604053MT604066) and all residual DNAs are archived at EBCL, FSCA, and SYSU (Table 2). Voucher specimens which have been reexamined following the molecular analysis were deposited in public collections (Table 2). All barcode sequences were translated into amino acids to check for stop codons. The sequences obtained were compared with sequences present in GenBank using the Basic Local Alignment Search Tool (http://www.ncbi.nlm.nih.gov/BLASTn) and aligned with the barcode sequences of Gryon species previously available. BOLD (Ratnasingham and Hebert 2007) was similarly datamined for Gryon CO1 barcodes and evaluated for barcode identification success. Similar sequences not identified to at least genus-level were not included in neighbor-joining analyses or K2P distance calculations. Tree search was performed in MEGA7 using the K2P model with a 95% per site data coverage cutoff (Kumar et al. 2016, Kimura 1980). Branch support, calculated with 10,000 boostrap replicates, and distance calculations used the same parameters.

CO1 barcodes were translated into amino acids and aligned using the default settings of ClustalW (Thompson et al. 1994) as implemented in MEGA7 (Kumar et al. 2016). The amino acid alignment was then back-translated into DNA sequences for neighbor-joining and distance analyses. The resulting alignment contains 682 bp positions and 98 terminal taxa. A Psix species was chosen as an outgroup for neighbor-joining analysis based on the phylogenetic topologies recovered by Taekul et al. 2014.

Taxon treatment

Gryon ancinla Kozlov & Lê, 1996

Nomenclature

Gryon ancinla Kozlov & Lê, 1996, Lê 1996: 11 (original description); Lê 2000: 98, 102 (description, keyed, type information).

Gryon clavaerus Kozlov & Lê syn. n., 1996, Lê 1996: 12 (original description); Lê 2000: 99, 108 (description, keyed, type information).

Materials    Download as CSV 
Holotypes:
  1. scientificName:
    Gryon ancinla
    ; kingdom:
    Animalia
    ; phylum:
    Arthropoda
    ; class:
    Inseca
    ; order:
    Hymenoptera
    ; family:
    Scelionidae
    ; genus:
    Gryon
    ; specificEpithet:
    ancinla
    ; scientificNameAuthorship:
    Kozlov & Lê
    ; country:
    Vietnam
    ; stateProvince:
    Bac Thai
    ; county:
    Phu Luong
    ; locality:
    Quan Chun Village
    ; locationRemarks:
    label transliteration "Bac Thai [Province], Phu Luong [District], Quan Chun Village, 16.IV.1986, A. Sharkov [leg.] " . "Holotypus Gryon ancinla sp. n. 86"
    ; eventDate:
    05/16/1986
    ; individualCount:
    1
    ; sex:
    female
    ; lifeStage:
    adult
    ; recordedBy:
    Elijah Talamas
    ; identifiedBy:
    Kozlov & Lę
    ; dateIdentified:
    1996
    ; institutionCode:
    Zoological Institute, St. Petersburg, Russia (ZIN)
    ; basisOfRecord:
    PreservedSpecimen
  2. scientificName:
    Gryon clavaerus
    ; scientificNameID:
    urn:lsid:biosci.ohio-state.edu:osuc_names:179785
    ; kingdom:
    Animalia
    ; phylum:
    Arthropoda
    ; class:
    Hexapoda
    ; order:
    Hymenoptera
    ; family:
    Scelionidae
    ; genus:
    Gryon
    ; specificEpithet:
    clavaerus
    ; country:
    Vietnam
    ; stateProvince:
    Hanoi
    ; locality:
    Hanoi, Hanoi Prov., Vietnam
    ; decimalLatitude:
    21.0333
    ; decimalLongitude:
    105.85
    ; georeferenceSources:
    GEOnet
    ; samplingProtocol:
    none specified
    ; eventDate:
    05/04/1979
    ; verbatimEventDate:
    May-04-1979
    ; individualCount:
    1
    ; sex:
    female
    ; lifeStage:
    adult
    ; catalogNumber:
    IEBR 0170
    ; identifiedBy:
    Kozlov, M. A. (Mikhail Alexeevich) & Lê, X. H. (Xuân Huê)
    ; dateIdentified:
    1996
    ; modified:
    2019-08-12T10:05:37Z
    ; language:
    en
    ; institutionCode:
    Institute of Ecology and Biological Resources, Hanoi, Vietnam (IEBR)
    ; collectionCode:
    Insects
    ; basisOfRecord:
    PreservedSpecimen
    ; source:
    http://hol.osu.edu/spmInfo.html?id=IEBR%200170
    ; occurrenceID:
    urn:lsid:biosci.ohio-state.edu:osuc_occurrences:IEBR__0170
Other materials:
  1. scientificName:
    Gryon ancinla
    ; kingdom:
    Animalia
    ; phylum:
    Arthropoda
    ; class:
    Inseca
    ; order:
    Hymenoptera
    ; family:
    Scelionidae
    ; genus:
    Gryon
    ; specificEpithet:
    ancinla
    ; scientificNameAuthorship:
    Kozlov & Lê
    ; country:
    China
    ; stateProvince:
    Guangdong
    ; municipality:
    Guangzhou
    ; locality:
    Huofoshan
    ; locationRemarks:
    [CHINA: Guangdong Prov., Guangzhou, Huofoshan, MT in jungle 7-15.VI.2017 23°10'50"N 113°23'26"E, Chen & Talamas][PL197][DNAVoucher]
    ; verbatimCoordinates:
    23°10'50"N 113°23'26"E
    ; decimalLatitude:
    23.180556
    ; decimalLongitude:
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    ; georeferenceProtocol:
    label
    ; samplingProtocol:
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    ; eventDate:
    2017-6-7/15
    ; individualCount:
    1
    ; sex:
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    ; lifeStage:
    adult
    ; catalogNumber:
    FSCA 00094673
    ; recordedBy:
    Elijah Talamas
    ; identifiedBy:
    Elijah Talamas
    ; institutionCode:
    Florida State Collection of Arthropods, Gainesville, Florida, USA (FSCA)
    ; basisOfRecord:
    PreservedSpecimen
  2. scientificName:
    Gryon ancinla
    ; kingdom:
    Animalia
    ; phylum:
    Arthropoda
    ; class:
    Inseca
    ; order:
    Hymenoptera
    ; family:
    Scelionidae
    ; genus:
    Gryon
    ; specificEpithet:
    ancinla
    ; scientificNameAuthorship:
    Kozlov & Lê
    ; country:
    China
    ; stateProvince:
    Guangdong
    ; municipality:
    Guangzhou
    ; locality:
    Huofoshan
    ; locationRemarks:
    [CHINA: Guangdong Prov., Guangzhou, Huofoshan, MT in jungle 7-15.VI.2017 23°10'50"N 113°23'26"E, Chen & Talamas][PL197][DNAVoucher]
    ; verbatimCoordinates:
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    ; decimalLatitude:
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    ; decimalLongitude:
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    ; eventDate:
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    ; sex:
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    ; catalogNumber:
    FSCA 00094672
    ; recordedBy:
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    ; identifiedBy:
    Elijah Talamas
    ; institutionCode:
    Florida State Collection of Arthropods, Gainesville, Florida, USA (FSCA)
    ; basisOfRecord:
    PreservedSpecimen
  3. scientificName:
    Gryon ancinla
    ; kingdom:
    Animalia
    ; phylum:
    Arthropoda
    ; class:
    Inseca
    ; order:
    Hymenoptera
    ; family:
    Scelionidae
    ; genus:
    Gryon
    ; specificEpithet:
    ancinla
    ; scientificNameAuthorship:
    Kozlov & Lê
    ; country:
    China
    ; stateProvince:
    Guangdong
    ; municipality:
    Guangzhou
    ; locality:
    Huofoshan
    ; locationRemarks:
    [CHINA: Guangdong Prov., Guangzhou, Huofoshan, MT in jungle 7-15.VI.2017 23°10'50"N 113°23'26"E, Chen & Talamas][PL197][DNAVoucher]
    ; verbatimCoordinates:
    23°10'50"N 113°23'26"E
    ; decimalLatitude:
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    ; eventDate:
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    ; individualCount:
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    ; sex:
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    ; catalogNumber:
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    ; recordedBy:
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    ; identifiedBy:
    Elijah Talamas
    ; institutionCode:
    Florida State Collection of Arthropods, Gainesville, Florida, USA (FSCA)
    ; basisOfRecord:
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  4. scientificName:
    Gryon ancinla
    ; kingdom:
    Animalia
    ; phylum:
    Arthropoda
    ; class:
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    Hymenoptera
    ; family:
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    ancinla
    ; scientificNameAuthorship:
    Kozlov & Lê
    ; country:
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    ; stateProvince:
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    ; municipality:
    Guangzhou
    ; locality:
    Huofoshan
    ; locationRemarks:
    [CHINA: Guangdong Prov., Guangzhou, Huofoshan, MT in jungle 7-15.VI.2017 23°10'50"N 113°23'26"E, Chen & Talamas][PL197][DNAVoucher]
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    Elijah Talamas
    ; institutionCode:
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  5. scientificName:
    Gryon ancinla
    ; kingdom:
    Animalia
    ; phylum:
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    Hymenoptera
    ; family:
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    ; specificEpithet:
    ancinla
    ; scientificNameAuthorship:
    Kozlov & Lê
    ; country:
    China
    ; stateProvince:
    Guangdong
    ; municipality:
    Huizhou
    ; locality:
    Nan Kun Shan Forest Park
    ; locationRemarks:
    [CHINA: Guangdong Prov., Huizhou, Nan Kun Shan Forest Park, MT in tea field beside stream & forest 8-14.VI.2017 23°38'8"N, 113°51'4"E Chen & Talamas][GRYON115][DNA Voucher]
    ; verbatimCoordinates:
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    ; identifiedBy:
    Elijah Talamas
    ; institutionCode:
    Florida State Collection of Arthropods, Gainesville, Florida, USA (FSCA)
    ; basisOfRecord:
    PreservedSpecimen
  6. scientificName:
    Gryon ancinla
    ; kingdom:
    Animalia
    ; phylum:
    Arthropoda
    ; class:
    Inseca
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    Hymenoptera
    ; family:
    Scelionidae
    ; genus:
    Gryon
    ; specificEpithet:
    ancinla
    ; scientificNameAuthorship:
    Kozlov & Lê
    ; country:
    China
    ; stateProvince:
    Guangdong
    ; municipality:
    Huizhou
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    Nan Kun Shan Forest Park
    ; locationRemarks:
    [CHINA: Guangdong Prov., Huizhou, Nan Kun Shan Forest Park, YPT 9-10.VI.2017 trail along stream, 23°37'6"N 113°51'33"E, Chen & Talamas][GRYON118][DNA Voucher]
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    Elijah Talamas
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    Elijah Talamas
    ; institutionCode:
    Florida State Collection of Arthropods, Gainesville, Florida, USA (FSCA)
    ; basisOfRecord:
    PreservedSpecimen
  7. scientificName:
    Gryon ancinla
    ; kingdom:
    Animalia
    ; phylum:
    Arthropoda
    ; class:
    Inseca
    ; order:
    Hymenoptera
    ; family:
    Scelionidae
    ; genus:
    Gryon
    ; specificEpithet:
    ancinla
    ; scientificNameAuthorship:
    Kozlov & Lê
    ; country:
    Cambodia
    ; stateProvince:
    Khum Ta Tey Leu
    ; locality:
    Cardamom Mtns.
    ; locationRemarks:
    [CAMBODIA: Khum Ta Tey Leu, Cardamom Mtns. MT 11°59'48.4"N 103°25'12.4"E 7-JUL-2010, Seunghwan Lee][PL077][DNA Voucher]
    ; verbatimCoordinates:
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    ; decimalLatitude:
    11.996778
    ; decimalLongitude:
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    ; georeferenceProtocol:
    label
    ; samplingProtocol:
    Malaise trap
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    07/07/2010
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    1
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    adult
    ; catalogNumber:
    FSCA 00090590
    ; recordedBy:
    Elijah Talamas
    ; identifiedBy:
    Elijah Talamas
    ; institutionCode:
    College for Agriculture and Life Sciences, Seoul National University, Seoul, Korea (SNU)
    ; basisOfRecord:
    PreservedSpecimen
  8. scientificName:
    Gryon ancinla
    ; kingdom:
    Animalia
    ; phylum:
    Arthropoda
    ; class:
    Inseca
    ; order:
    Hymenoptera
    ; family:
    Scelionidae
    ; genus:
    Gryon
    ; specificEpithet:
    ancinla
    ; scientificNameAuthorship:
    Kozlov & Lê
    ; country:
    China
    ; stateProvince:
    Zhejiang
    ; locality:
    Tianmu Mountain (West)
    ; verbatimElevation:
    930 m
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Description

Size: Female body length: 1.26–1.86 mm (n=9).

Color: Color of body: dark brown to black, rarely with reddish-brown areas. Color of legs: coxae dark brown to black, otherwise yellow.

Head: Number of papillary sensilla on A7: 2. Number of papillary sensilla on A8: 2. Color of antenna in female: A1–A6 yellow, A7–A12 brown. Number of mandibular teeth: 3. Shape of mandibular teeth: dorsal tooth distinctly the largest. Shape of clypeus: roughly rectangular with rounded corners. Number of clypeal setae: 6. Epiclypeal carina: present. Facial striae: absent. Central keel: present. Line of setae above interantennal process: absent. Setation of compound eye: short and sparse, often appearing absent with light microscopy. Setation of orbital furrow: present along dorsal half of compound eye. Macrosculpture of frontal depression: transversely rugose. Sculpture of frons outside of frontal depression: areolate rugose. Lateral margin of frontal depression: delimited by carina. Dorsal margin of frontal depression: delimited by carina. Smooth area at base of mandible: present. Malar striae: absent. Genal carina: present. Hyperoccipital carina: present between lateral ocelli. Anterior margin of occipital carina on gena: crenulate. Occipital carina: terminating near dorsal margin of compound eye.

Mesosoma: Epomial carina: present. Sculpture of lateral pronotum: transversely rugose. Netrion sulcus: absent. Mesoscutal suprahumeral sulcus: absent. Mesoscutal humeral sulcus: present as a smooth furrow, anteriorly terminating in a pit. Sculpture of mesoscutum: coarsely rugose, with rugae oriented longitudinally at posterior margin. Posterior mesoscutellar sulcus: foveate. Posterior margin of mesoscutellum: extending over metanotum, metascutellum not visible in dorsal view. Posterior margin of metascutellum: straight. Sculpture on posteroventral surface metascutellum: antero-posteriorly strigose. Sculpture of metanotal trough: foveate. Length of postmarginal vein in fore wing: about twice as long as stigmal vein. Length of marginal vein in fore wing: about half as long as stigmal vein. Lateral propodeal carina: continuous across posterior propodeum, forming flange around metasomal depression. Sculpture of metasomal depression: radially strigose. Preacetabular sulcus: present as a line of punctures. Orientation of acetabular carina: parallel to mesopleural carina. Posterior limit of acetabulum: nearly reaching ventral mesopleural carina. Postacetabular sulcus: crenulate. Episternal foveae: absent. Mesopleural carina: present. Cells or foveae along ventral margin of mesopleural carina: present. Sculpture of femoral depression: irregularly rugose. Prespecular sulcus: indicated by crenulae. Sculpture of speculum: transversely rugose. Mesepimeral sulcus: comprised of circular foveae. Sculpture of posterior mesepimeral area: weakly rugulose in ventral half. Paracoxal sulcus: indicated by large, irregular cells along anterior margin of metapleuron. Anteroventral extension of the metapleuron: long, reaching base of mesocoxa. Metapleural structure: dorsoventrally divided by carina, posterior portion densely setose.

Metasoma: Form of sulcus on anterior T1: simple line of foveae. Lateral pit on anterior T1: absent. Macrosculpture of T1: longitudinally striate. Setation of T1: present in a posterolateral triangular area. Smooth area on anterior T2: present. Setation of T2: sparse medially, dense laterally. Macrosculpture of T2: irregularly rugose. Posterior margin of T6: concave. Lateral pit on anterior S1: absent. Transverse sulcus on anterior S2: absent. Macrosculpture of S2: sparsely striate, striae attenuating posteriorly.

Variation: Specimens FSCA 00094670 and FSCA 00094672 (Fig. 10) are both female, are from the same Malaise trap sample and have identical CO1 barcode sequences. They are also notably different in size (1.67 and 1.27 mm, respectively) and exhibit differences in the sculpture of the frons between the frontal depression and the inner orbit of the compound eye. Specimen FSCA 00094670, which is the larger specimen, has a ridge extending from the orbital carina to the margin of the frontal depression (Fig. 10a). Interestingly, the location of this ridge along the inner orbit corresponds to the transition point between the setose and glabrous portions of the orbital furrow. This ridge can clearly be seen in the holotype specimen of G. ancinla (Fig. 2b). The smaller of the two specimens, FSCA 00094672, has the frons evenly rugose between the orbital furrow and the frontal depression, without a transverse ridge (Fig. 10b).

Figure 10.

Variation in the sculpture on the frons in Gryon ancinla.

aGryon ancinla, female (FSCA 00094670), head, anterolateral view, image flipped horizontally  
bGryon ancinla, female (FSCA 00094672), head, anterior view  

Diagnosis

Females of G. ancinla have a 6-merous clava (Fig. 4), which is found in other species of the charon group, including the African species G. charon and G. paracharontis. The holotype of G. sponus, from Vietnam, is missing its antennae (Fig. 5), but the illustration of the female antenna in Lê 1996 suggests that it has 6 clavomeres. Each of these species can also be separated from G. ancinla by the shape of the mesoscutellum. In G. ancinla, the posterior margin of the mesoscutellar disc is directly above the posterior margin of the scutellar rim. In G. charon, G. paracharontis, and G. sponus, the mesoscutellar disc extends posteriorly well beyond the scutellar rim (Fig. 5). Gryon drunoris, which is sympatric with G. ancinla, has a mesoscutellum that is evenly convex and the clava is 5-merous (Fig. 6). Gryon ancinla and G. drunoris may also be separated by the relative lengths of the metasomal tergites: T1 is distinctly longer than T3 in G. ancinla (Fig. 7a) and they are roughly equal in G. drunoris (Fig. 7b). In the females of the charon species group that we have examined so far, the clava tends to be distinctly darker than the preceding antennomeres. This makes the clava easily distinguishable from the funicle in most cases, but we caution that using color to differentiate the clava from the funicle may not be reliable in all species or specimens and unambiguous determination of the number of clavomeres requires examination of the papillary sensilla.

Notes

Gryon clavaerus (Fig. 3) was described in the same publication as G. ancinla. Our examination of the type specimens finds no differences that justify keeping them as separate species and we thus treat G. clavaerus as a junior synonym.

Analysis

Characters in the charon species group

Humeral pit

The mesoscutal humeral pit (Fig. 8) is located at the junction of the mesoscutal humeral sulcus and the mesoscutal suprahumeral sulcus. This pit is found in all species of the charon group that we have examined. It is also present in species that are not part of the charon group as it is currently defined, and these species have varying forms of carinae surrounding the frontal depression. The mesoscutal humeral pit thus may be useful for determining affinities between the charon group and other lineages within Gryon.

Setation of the orbital furrow

Setation of the orbital furrow can separate some species in the charon group and perhaps other species groups. Gryon ancinla has setation only in the dorsal part of the orbital furrow (Figs 2b, 4), whereas some species, including the African G. letus, have setation throughout the orbital furrow (Fig. 9).

Molecular analysis

The neighbor-joining analysis revealed relatively large sequence divergences between clusters of Gryon CO1 barcodes (Fig. 11). Interpretation of sequence divergence in Gryon is currently hampered by the lack of species-level identifications that are necessary to define intra- and interspecific variation. We included 14 new CO1 barcodes from members of the Gryon charon species group and our neighbor-joining analysis recovered a cluster of these species with an additional unidentified Gryon from South Africa (BIN: BOLD:ADO2077; SAFRA3055-18, SAFRA4239-18) (Ratnasingham and Hebert 2013). We found two haplogroups of G. ancinla, indicated in Fig. 11 by the red and magenta branches. Specimens from each lineage were collected in a single Malaise trap sample in Guangzhou (FSCA 00094670–00094673), demonstrating that the haplogroups are sympatric. These Gryon ancinla haplogroups differ by K2P distances ranging from 9.6–10.4% (Table 1) and they are each other’s nearest neighbor. BLASTn searches yielded poor matches to the G. ancinla barcodes (86–87% identity to other hymenopteran barcodes). In BOLD, G. ancinla from haplogroup 1 were a 97% match to an unidentified specimen from Bangladesh (GMBCB2151-15) and the available image of this specimen is consistent with our concept of G. ancinla. This suggests that G. ancinla has a wide geographical distribution in southeast Asia.

Figure 11.  

K2P neighbor-joining tree demonstrating the clustering of Gryon CO1 barcodes. The larger cluster in blue highlights the Gryon charon species group. Clusters in red and magenta highlight the Gryon ancinla haplogroups. Bootstraps values of 80 and above are indicated.

Discussion

Gryon contains widespread species and geographically broad analysis is needed to identify synonyms. This study characterizes a species found in southeast Asia to facilitate comparison with similar species of Gryon in the region and to associate ecological data with a taxonomic name.

Acknowledgements

We extend our thanks to the CanaColl Foundation, which funded a visit by Elijah Talamas to the Canadian National Collection and to Lubomír Masner (CNCI) who hosted this visit. We are extremely grateful to Sergey Bolokobilskij (ZIN) for providing the images of the holotype of Gryon ancinla and to Yingqi Liu (China Agricultural University, Beijing), who provided images of the holotyope of Gryon charon. Specimens from South Africa were made available on loan thanks to the efforts of Simon van Noort and Aisha Mayekiso (SAMC) and specimens from Cambodia were made available by Nam Sangheok (SNU). We thank Cheryl Roberts and Lynn Combee (FDCAS-DPI) for their assistance with generating CO1 barcodes for this project. Elijah Talamas and Matthew Moore were supported by the Florida Department of Agriculture and Consumer Services, Division of Plant Industry. Huayan Chen was supported by the National Natural Science Foundation of China (31900346). Taxonomic research on Gryon in Asia and Africa is supported by a USDA-APHIS Farm Bill, Biological Control of Bagrada Bug. The USDA is an equal opportunity employer and does not endorse any commercial product mentioned in this research.

References