6 urn:lsid:arphahub.com:pub:DE333E49-E4EE-5693-9BEF-D446E6301C98 urn:lsid:zoobank.org:pub:55528528-5C97-4D79-A718-8B3D153B37CC Journal of Hymenoptera Research JHR 1070-9428 1314-2607 Pensoft Publishers 10.3897/jhr.99.205393 205393 Research Article Chalcidoidea Pteromalidae Identification key Molecular systematics Phylogeny Taxonomy Cenozoic Asia Testing the generic status of Philoverdance and revising the Philotrypesis ravii species group (Otitesellini, Pteromalidae) associated with Ficus tinctoria (Moraceae) Li Yu-Ran https://orcid.org/0000-0003-2348-8224 1 2 Writing - original draft Writing - review and editing Data curation Formal analysis Visualization Cruaud Astrid https://orcid.org/0000-0001-8932-4199 3 Writing - review and editing Data curation Formal analysis Methodology Visualization Nidelet Sabine https://orcid.org/0000-0003-0437-319X 3 Writing - review and editing Methodology Peng Yan-Qiong pengyq@xtbg.ac.cn https://orcid.org/0000-0002-7453-9119 1 Conceptualization Funding acquisition Project administration Supervision Rasplus Jean-Yves jean-yves.rasplus@inrae.fr https://orcid.org/0000-0001-8614-6665 3 Conceptualization Writing - original draft Writing - review and editing Methodology Supervision Validation Visualization Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla 666303, China Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences Mengla China https://ror.org/02rz58g17 University of Chinese Academy of Sciences, Beijing, 10049, China CBGP, INRAE, CIRAD, IRD, Montpellier SupAgro, Université de Montpellier Montpellier France https://ror.org/051escj72 CBGP, INRAE, CIRAD, IRD, Montpellier SupAgro, Université de Montpellier, Montpellier, France University of Chinese Academy of Sciences Beijing China https://ror.org/05qbk4x57

Corresponding authors: Yan-Qiong Peng (pengyq@xtbg.ac.cn); Jean-Yves Rasplus (jean-yves.rasplus@inrae.fr)

Academic editor: Mircea-Dan Mitroiu

 2026 25 08 2026 99 915 954 0D18AD8F-2434-5B82-8D1B-09F687421F86 21F7F738-D972-47F1-9DDE-B0E969ABA863 25 06 2026 04 08 2026 Yu-Ran Li, Astrid Cruaud, Sabine Nidelet, Yan-Qiong Peng, Jean-Yves Rasplus This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. https://zoobank.org/21F7F738-D972-47F1-9DDE-B0E969ABA863

Philotrypesis Förster is one of the most species-rich genera of non-pollinating fig wasps associated with Old World fig trees (Moraceae, Ficus). Philoverdance Priyadarsanan was initially established to accommodate a morphologically distinctive species of Philotrypesis associated with Ficus tinctoria in India, but its phylogenetic position remained uncertain. We reassessed the status of Philoverdance using morphology and genomic data. Philoverdance was recovered nested within Philotrypesis, supporting its synonymization with Philotrypesissyn. nov. and its treatment as the P. ravii species group. In depth analysis of the species group revealed five species associated with F. tinctoria: P. ravii Priyadarsanan, P. minang Li & Rasplus, sp. nov. (Sunda), P. bannaensis Li & Rasplus, sp. nov. (Sino-Himalaya), P. paiwan Li & Rasplus, sp. nov. (Sino-Himalaya), and P. arfak Li & Rasplus, sp. nov. (Australasia). COI divergences among species ranged from 6.1 to 8.4%, supporting their distinctiveness. Both COI and BUSCO-based phylogenies recovered a western clade associated with F. tinctoria subsp. gibbosa and an eastern clade associated with F. tinctoria subsp. tinctoria. Molecular, morphological and biogeographic evidence were largely congruent, revealing a geographically structured diversification pattern across the range of the host plant. Diagnoses, descriptions and identification keys for females and males are provided for all species.

 Chalcidoidea DNA barcodes fig trees key new species the program Yunnan Key Research and Development Program (202503AP140027) and the program National Natural Science Foundation of China (32261123001). Introduction

Philotrypesis Förster, 1878 is a remarkable genus of Old World non-pollinating fig wasps belonging to the tribe Otitesellini of Pteromalidae (Förster 1878; Burks et al. 2022, 2025; Cruaud et al. 2024). The genus currently comprises 45 described species (UCD Community 2023), but its taxonomy remains poorly understood and is in urgent need of revision. Philotrypesis is a widespread genus, occurring from the Mediterranean Basin through Africa and tropical Asia to Australia. Like many lineages of non-pollinating fig wasps, Philotrypesis harbors an extraordinary amount of hidden species diversity. Over the last four decades, extensive collecting efforts covering more than 2,000 fig samples from approximately 160 host fig species have yielded more than 250 putative species of Philotrypesis, of which nearly 220 appear to be new to science. This remarkable and largely undocumented diversity highlights how poorly known non-pollinating fig wasp faunas remain despite their ecological importance and abundance in tropical ecosystems (Kerdelhué et al. 2000; Segar and Cook 2012; Wu et al. 2013).

Philotrypesis females oviposit from outside the syconium, usually after pollination and near the end of syconium development, inserting their long ovipositor through the fig wall into individual ovules (Abdurahiman 1980; Zhen et al. 2004). Ovipositor length relative to fig wall thickness appears to be a major ecological constraint shaping host use and diversification (Yu et al. 2018). The biology of the genus remains poorly understood and includes inquiline, possible parasitoid and even seed-predatory strategies (Joseph 1955; Wang et al. 2014). Males are generally wingless, often highly polymorphic, and engage in intense competition for mates involving fighting and alternative reproductive tactics, whereas winged males occur only rarely (Murray 1989, 1990; Jousselin et al. 2004; Cook and Bean 2006; Moore et al. 2009). Species have been recorded from most major Old World fig lineages, including Sycomorus, Oreosycea, Terega, Sycidium, Synoecia and multiple lineages of subgenus Urostigma (Wiebes 1966a; Wiebes 1967; Abdurahiman and Joseph 1975; Balakrishnan Nair et al. 1981; Bouček et al. 1981; Bouček 1988; McLeish et al. 2010). Although host specificity can be strong in some groups (Jousselin et al. 2008), molecular studies indicate frequent host switching, with several Philotrypesis species often co-occurring on a single host fig and some species exploiting multiple hosts (Jiang et al. 2006; Deng et al. 2021). Despite its abundance and ecological importance, Philotrypesis remains poorly studied, like most other non-pollinating fig wasp groups.

The genus is readily recognized by a set of distinctive morphological apomorphies (Grandi 1921; Grandi 1930; Wiebes 1966b). The terminal metasomal tergites (Mt7 and Mt8+9) are elongated and form a characteristic gastral extension or “tail”, while the ovipositor sheaths remain conspicuously visible beyond the apex of the gaster. Mt7 is broader than the terminal syntergum (Mt8+9), which may itself become strongly elongate. In addition, females typically possess bidentate mandibles, complete linear notauli, and antennae with three anelli. Males are highly characteristic and, in several species, display a subalate morphology remarkably similar to that observed in the related genus Watshamiella Wiebes, 1981 (Bouček et al. 1981), suggesting a close relationship between the two taxa.

The limits of Philotrypesis have nevertheless remained uncertain because of the existence of Philoverdance Priyadarsanan, 2000, a monotypic genus erected to accommodate Philoverdance ravii from India (Priyadarsanan 2000). Although Philoverdance exhibits all diagnostic apomorphies of Philotrypesis, it was distinguished on the basis of several morphological features, including a more compact body, less elongate terminal metasomal tergites, monodentate mandibles, a shorter pronotum, and a smoother body sculpture (Priyadarsanan 2000). However, overlap between these characters and the morphological variation observed within Philotrypesis was soon noted, raising doubts about the validity of the genus (Segar et al. 2012). The lack of comparative material and the absence of comprehensive phylogenetic analyses have further contributed to uncertainty regarding the status of Philoverdance. To date, its placement has been examined only once in a molecular framework, using a Sanger dataset (Segar et al. 2012). In that study, the species assigned to Philoverdance was recovered within a well-supported Philotrypesis clade, rendering Philotrypesis paraphyletic, although the position of the taxon remained weakly supported and somewhat ambiguous.

The primary objective of this study is to propose a preliminary phylogenomic hypothesis for Philotrypesis in order to test the validity and phylogenetic position of Philoverdance and determine whether it should be maintained as a distinct genus, reduced to a subgenus, or treated as a species group within Philotrypesis. We further analyze genomic (BUSCO), mitochondrial (COI), and morphological data to assess species diversity within this lineage, describe four previously unrecognized species, and investigate patterns of diversification across the geographic range of its host, Ficus tinctoria. More broadly, our analyses provide a first phylogenetic framework for Philotrypesis that will facilitate future taxonomic revision of this exceptionally species rich and still poorly known genus.

 Methods Sampling

Most specimens examined in this study were reared from figs (syconia) of different species of Ficus during the 2000s by J.-Y. Rasplus (JYR) and A. Cruaud (AC), and more recently by Y. Li (YL). Fully developed D-phase figs were collected and maintained in fine mesh bags until adult wasps emerged. Emerging individuals were carefully collected and fixed in 75% ethanol. Samples were stored at -20 °C until morphological examination, DNA extraction and sequencing. Additional material collected in the 1990s and 2000s was obtained from colleagues. Geographical coordinates and elevations were either recorded in the field or subsequently inferred, when possible, from specimen labels and associated field notes.

 Morphology

Specimens were dehydrated using hexamethyldisilazane (HMDS) following Heraty and Hawks (1998) and card-mounted following the procedure outlined by Noyes (1982). Mounted specimens were examined and curated using a Nikon SMZ25 stereomicroscope (Nikon Corp., Melville, NY, USA). High-resolution images and morphometric measurements were obtained from photographs acquired with a Keyence VHX-5000 digital multifocus imaging system (Keyence Corporation, Osaka, Japan). A total of 30 morphometric characters for females and 13 characters of males (Table 1, Fig. 1) were measured, whenever possible, from at least three individuals per sample using Fiji software (Schindelin et al. 2012) based on photographs. Image assembly, editing and plate preparation were performed using Affinity Photo.

10.3897/jhr.99.205393.figure1 51182A3F-9AE5-53DA-84DB-5BCFA35A75D4

Female P. ravii, main measurements. A habitus, lateral view, wings removed B head, dorsal view C head, frontal view D antenna E mesosoma, dorsal view F fore wing venation.

https://binary.pensoft.net/fig/1753542

Morphological measurements and abbreviations used in species descriptions and comparisons.

Morphological abbreviations Explanation
Head
Head L head length in dorsal view
Head W maximum head width in dorsal view
Head H head height in frontal view
TML (♂) temple length, measured from the posterior margin of the compound eye to the posterior margin of the head
Eye L maximum eye length
Eye H maximum eye height
POL postocellar line
OOL oculo-ocellar line
LOL lateral ocellar line
mls malar space
Antenna
Ped+Fla L combined length of pedicel and flagellum
Fl4 L length of fourth flagellar segment
Fl4 W width of fourth flagellar segment
Fl8 L length of eighth flagellar segment
Fl8 W width of eighth flagellar segment
Mesosoma
Mesosoma L (♂) maximum mesosoma length
Mesosoma W (♂) maximum mesosoma width
no1 L pronotum length
no1 W pronotum width
msc L mesoscutum length
mss L mesoscutellum length
Propodeum L (♂) median length of propodeum
Wings
mv L marginal vein length
stv L stigmal vein length
pmv L postmarginal vein length
FWr L (♂) length of fore wing remnant
Legs
tb3 L metatibia length
Body, metasoma and ovipositor
Body L body length from anterior margin of head to posterior margin of Mt8+9
OSL ovipositor sheath length
Mt2–Mt6 L combined length of metasomal tergites II–VI
Mt2–Mt6 W maximum width across metasomal tergites II–VI
Mt7 L length of metasomal tergite VII

Morphological terminology follows Bouček and Rasplus (1991), Gibson (1997), and Burks et al. (2025). Abbreviations used throughout the text are defined in Table 1, and measurements and ratios are illustrated in Figs 1, 2. For each new species, the description is based primarily on the female holotype and supplemented by variation observed among paratypes. Because multiple species of Philotrypesis may co-occur within the same syconium, male–female associations were established cautiously using shared morphological attributes and by comparing samples in which only a single species, or a reduced subset of species, was present. In this study, only one male individual (LYRA00012_0301) from sample LYRA00012 was sequenced and included in the COI-based phylogeny alongside the female individuals.

10.3897/jhr.99.205393.figure2 607718EC-1F44-5E53-A51D-54DF9FCFEB05

Male P. ravii, main measurements. A, B habitus, dorsal view C habitus, lateral view D head, frontal view.

https://binary.pensoft.net/fig/1753543

Specimens examined during this study are deposited in the following institutions:

CBGP Centre de Biologie pour la Gestion des Populations, Montferrier-sur-Lez, France (Curator: Emma Artige);

MZB Museum Zoologicum Bogoriense, Research Center for Biology, Indonesian Institute of Sciences (BRIN), Cibinong, Indonesia;

MZDC Museum of Zoology, Department of Zoology, University of Calicut, Kerala, India;

XTBG Insect Collection, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Menglun, Yunnan, China.

 DNA extraction

DNA was extracted using the Qiagen DNeasy Blood and Tissue kit, following manufacturer’s instructions with minor modifications as described in Cruaud et al. (2019). Genomic DNA integrity was assessed by agarose gel electrophoresis, and DNA extracts were quantified using a Qubit 2.0 fluorometer (Invitrogen).

 Whole-genome sequencing (WGS)

For samples yielding less than 40 ng of genomic DNA, whole genome amplification was performed using the GenomiPhi V3 Ready-To-Go DNA Amplification Kit (Cytiva) following manufacturer’s instructions. Samples with sufficient DNA (> 500 ng) were sent to Berry Genomics Co., Ltd (http://www.berrygenomics.com/), Beijing, China to prepare genomic libraries and sequenced on the NGS (next generation sequencing) platform. Library preparation was conducted using kits developed by Berry Genomics Co., Ltd and included the following steps: DNA fragmentation by sonication, end-repair and A-tailing of DNA fragments, ligation of sequencing adapters, PCR amplification, and purification of PCR products using the AMPure XP bead system (Beckman Coulter). Library size distributions were assessed using an Agilent 2100 Bioanalyzer, and library concentrations were quantified by real-time PCR. The resulting libraries were sequenced on an Illumina NovaSeq 6000 platform. Multiple samples were loaded into one lane for sequencing, with approximately 1,000 Gb of PF (pass filter) data per lane. Samples were distinguished by unique index sequences, and raw data were demultiplexed accordingly after sequencing with a custom script. Raw paired-end reads were filtered using fastp (Chen et al. 2018) for quality control. Bases with a quality score below Q5 were defined as low-quality bases (-q 5), and reads were discarded if low-quality bases accounted for over 20% of the read length in either forward or reverse read (-u 20). Adapter sequences were automatically detected and removed based on the provided adapter sequences. After filtering, approximately 10 Gb of 150 bp paired-end clean reads were retained per sample.

 Barcoding

The 5' region of the cytochrome c oxidase subunit I (COI) gene, corresponding to the standard DNA barcode for insects, was either recovered from WGS data (sample codes beginning with LYRA in the COI tree; Fig. 4) or amplified by PCR and sequenced using Oxford Nanopore Technologies (ONT) (sample codes beginning with JRAS in the COI tree; Fig. 4). In addition, one outgroup taxon was sequenced for COI using the PCR-ONT approach (Philotrypesis sp. associated with F. parietalis). Generating COI barcodes for all species provides a reference framework for future species identification and the assignment of newly collected material. PCR-based amplification also enabled the inclusion of older or poorly preserved specimens that failed to yield usable DNA or sequences through the shotgun sequencing approach.

For the PCR-ONT approach: The COI barcode fragment was amplified with standard primer pairs that were asymmetrically indexed using Unique Molecular Identifiers (UMIs), according to the protocol of Hebert et al. (2025). Each PCR reaction (10 µl total volume) contained 5 µl of Qiagen Taq Multiplex Mastermix, 2 µl of ultrapure water, 1 µl of forward primer (2 µM), 1 µl of reverse primer (2 µM), and 1 µl of DNA template. Amplifications were carried out on an Eppendorf Mastercycler Nexus with the following thermal profile: an initial denaturation step at 95 °C for 15 min; 45 cycles of denaturation at 95 °C for 40 s, annealing at 45 °C for 40 s, and extension at 72 °C for 1 min; followed by a final extension at 72 °C for 2 min and a hold at 15 °C.

PCR products (1 µl) from all specimens were pooled together with amplicons generated in a larger experiment (N specimens total = 752). The combined pool was purified using AMPure XP magnetic beads (Agencourt). Library preparation, including end-repair, adapter ligation, and clean-up, followed the Oxford Nanopore Technologies Ligation Sequencing of Amplicons protocol V14 (SQK-LSK114), version V (ACDE_9163_v114_revW_30Jan2025), and the final library was loaded onto two Flongle flow cells at a concentration of 10 fmol for sequencing on a MinION Mk1B.

Basecalling and FASTQ file generation from POD5 files were performed using Dorado v0.9.0 with the super-accurate model (v4.3.0, 400 bps). After demultiplexing and removal of UMIs and primer sequences with a custom Python script, reads exceeding 600 bp were clustered using VSEARCH v2.29.3 (Rognes et al. 2016) at a similarity threshold of 0.90, and consensus sequences were generated for each cluster. To correct for over splitting caused by PCR or sequencing errors, clusters with consensus sequences sharing more than 98% identity were merged. Reads from merged clusters were subsequently realigned using MAFFT v7.505 (Katoh and Standley 2013) to produce 50% majority-rule meta-consensus sequences.

Only dominant meta-consensus sequences supported by more than 100 reads were retained, provided that they corresponded to protein coding sequences and showed at least 90% identity over their full length to a congeneric species in the NCBI database, as no exact reference sequences were available for the taxa analysed. For species represented by multiple individuals, consistency of the recovered barcode sequences was also verified.

 Phylogeny based on single-copy orthologs

The cleaned raw reads of WGS were first de novo assembled using MEGAHIT v1.2.9 (Li et al. 2015). Then, single-copy orthologs were identified and extracted from the assembled contigs using BUSCO v6.0.0 (Tegenfeldt et al. 2025) with the hymenoptera_odb12 lineage dataset, and retained as nucleotide and amino acid sequences for downstream analysis. A total of 2,471 genes present in at least 90% of samples were retained, while samples with a presence rate of less than 50% across all genes were removed. For each orthologous gene, the amino acid and nucleotide sequences were aligned using MAFFT with the L-INS-i option. Resulting amino acid alignments were trimmed using trimAL v1.5.revl (Capella-Gutiérrez et al. 2009) with the “automated1” option, and nucleotide alignments were trimmed accordingly by back-translation from the trimmed amino acid alignments. The nucleotide alignments were concatenated using AMAS (Borowiec 2016) to produce a supermatrix, which was subsequently subjected to phylogenetic analysis in IQ-TREE2 using the partitioned model option MFP+MERGE and 1,000 replicates of ultrafast bootstrap. The initial partitioning strategy comprised 7,413 partitions corresponding to the first, second and third codon positions of each of the 2,471 BUSCO loci.

 Phylogenetic based on <abbrev xlink:title="cytochrome c oxidase subunit I">COI</abbrev> sequences

For samples successfully sequenced using the WGS approach, the cleaned reads were also used to assemble mitochondrial genomes de novo with the GetOrganelle toolkit (Jin et al. 2020). Mitogenomes were annotated using MITOS (Bernt et al. 2013). Protein-coding genes were subsequently inspected and curated in Geneious Prime 2021.1.1 (Biomatters, New Zealand), allowing manual adjustment of start and stop codons where necessary. COI sequences were extracted from the assembled mitogenomes and combined with those obtained through the PCR–ONT approach. Sequences were aligned using MAFFT with the L-INS-i algorithm. Phylogenetic inference was performed in IQ-TREE 2, with the best-fitting substitution model selected by ModelFinder (Kalyaanamoorthy et al. 2017) using the standard model-selection procedure (-m TEST). Node support was assessed using 100 standard nonparametric bootstrap replicates (-b 100). Pairwise sequence similarities were calculated using Geneious Prime 2021.1.1.

 Results Phylogenomic relationships and assessment of the validity of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Philoverdance">Philoverdance</tp:taxon-name-part></tp:taxon-name></italic>

Phylogenomic analyses based on BUSCO loci supported the major lineages recognized within Philotrypesis. The sampled Oriental diversity was subdivided into approximately ten well-supported species groups, including the P. pilosa Mayr, 1906, P. longicaudata Mayr, 1906, P. dunia Joseph, 1954, P. thompsoni Grandi, 1926, P. taiwanensis Chen, 1999, P. emeryi Grandi, 1926, P. ravii (Priyadarsanan, 2000) and P. glaberrima species groups (Fig. 3). Philoverdance was nested within Philotrypesis with strong support. No clear phylogenetic structure associated with host-fig taxonomy, ecology, life-history traits or geographic distribution was apparent from the recovered topology (Fig. 3).

10.3897/jhr.99.205393.figure3 86281CC8-D8B8-5788-B010-3E17F498D18A

Phylogeny of selected Philotrypesis species based on BUSCOs. A maximum likelihood (ML) phylogeny including Philotrypesis bannaensis sp. nov. (LYRA00012_02, LYRA00150_02, Yunnan, China), Philotrypesis minang sp. nov. (JRAS06684_02, Sumatra, Indonesia), Philotrypesis paiwan sp. nov. (JRAS02406_02, Taiwan, China), along with additional congeners. Numbers above branches represent ultrafast bootstrap support values of ML. Only values < 100 are shown B overview of host-fig taxonomy, life history traits, breeding systems and geographic distribution.

https://binary.pensoft.net/fig/1753544 <abbrev xlink:title="cytochrome c oxidase subunit I">COI</abbrev> divergence and phylogeographic structure of the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Philotrypesis">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ravii">ravii</tp:taxon-name-part></tp:taxon-name></italic> species group

COI barcoding (Fig. 4; Table 2) recovered five well-defined lineages within the Philotrypesis ravii species group. Pairwise genetic distances among these lineages ranged from 6.1% between the most closely related taxa to 8.4% between the most divergent taxa (Table 2). These values are substantially higher than those commonly reported among closely related species of Hymenoptera and Pteromalidae and greatly exceed the levels usually associated with intraspecific variation. Together with the observed morphological differences, these results strongly support the recognition of all lineages as distinct species.

10.3897/jhr.99.205393.figure4 161EA6DE-4A47-5B65-9C15-A96C7235BED4

Phylogeny of the Philotrypesis ravii sp. gp. based on cytochrome oxidase subunit I (COI) sequences. Maximum likelihood (ML) phylogeny of 5 species within the Philotrypesis ravii sp. gp. associated with Ficus tinctoria, as well as one outgroup, Philotrypesis sp. (JRAS01864_04) associated with F. parietalis. Numbers above branches represent standard bootstrap support values.

https://binary.pensoft.net/fig/1753545

Average pairwise dissimilarity within and between species based on COI sequences.

P. minang P. ravii P. bannaensis P. paiwan P. arfak
P. minang 0.0% 6.1% 6.6% 8.4% 7.7%
P. ravii 6.1% NA 6.7% 7.0% 7.8%
P. bannaensis 6.6% 6.7% 1.5% 8.0% 7.7%
P. paiwan 8.4% 7.0% 8.0% 0.0% 7.8%
P. arfak 7.7% 7.8% 7.7% 7.8% NA

The COI topology further reveals a pronounced phylogeographic structure within the species group. A western lineage comprises P. bannaensis sp. nov. from southern China, P. ravii from India, and P. minang sp. nov. from Sumatra. All three species are associated with Ficus tinctoria subsp. gibbosa, the western representative of the F. tinctoria complex (Berg and Corner 2005). In contrast, an eastern lineage comprises the sister species P. paiwan sp. nov. from Taiwan and P. arfak sp. nov. from West Papua, both associated with F. tinctoria subsp. tinctoria. Although based on a more limited taxon sampling, the BUSCO phylogeny is fully consistent with this pattern and independently supports a major evolutionary split between western and eastern lineages.

Remarkably, this geographic partition is mirrored by morphology. The eastern species share a suite of characters, including a ventrally orange metasoma, a sculptured mesoscutellum and relatively elongate ovipositor sheaths, whereas the western species generally possess a dark ventral metasoma, a smooth mesoscutellum and shorter ovipositors. The concordance among mitochondrial data, genomic data, morphology, geography and host-plant subdivision provides compelling evidence that the P. ravii species group consists of several independently evolving species whose diversification may have been influenced by the historical biogeography of the Ficus tinctoria complex.

 Taxonomic revision Animalia Hymenoptera Pteromalidae DA611479-6746-5CE3-A2B8-08B986A20A9D Philotrypesis Förster, 1878 Philotrypesis Förster, 1878: 59; type species by original designation or monotypy: Philotrypesis longicauda Förster, 1878. Philoverdance Priyadarsanan, 2000: 79–80; type species by original designation or monotypy: Philoverdance ravii Priyadarsanan, 2000, syn. nov. Animalia Hymenoptera Pteromalidae 7A428B21-5187-5B8B-AB4F-8E3326955E70 Philotrypesis ravii (Priyadarsanan, 2000) comb. nov. Figs 5A , 5C–F , 6A–D (♀), 5B, 6E, F (♂) Philoverdance ravii Priyadarsanan, 2000: 80–81. Holotype ♀ [MZDC], not examined. Type data: India, Calicut, 4.ii.1993; reared from figs of Ficus subsp. gibbosa (Blume) Corner [Terega (= Sycidium)] (Berg and Corner 2005; Pederneiras et al. 2015). Philoverdance ravii : Pramanik & Dey, 2016: SI. Type material.

India: • Calicut [Holotype ♀, MZDC].

 Distribution.

India [Oriental].

 Host <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ficus">Ficus</tp:taxon-name-part></tp:taxon-name></italic>.

F. tinctoria subsp. parasitica [Terega (=Sycidium)]. Series ♀♂, Calicut, coll. Priyadarsanan D.R., 4.ii.1993; 6 paratypes and 6 paratypes mounted on slides. (not examined).

 Additional material examined.

• 67♀, 62♂: India: Karnataka, Agumbe [13.5184 75.0898] 2.ii.2024 and ii.2005, Kjellberg F., ex F. tinctoria (JRAS01214_02 and JRAS01221_03 respectively) (CBGP).

 Diagnosis.

Female recognised by the weakly sculptured lower face and parascrobal area, antenna inserted just above the ocular line, straight clypeal margin, smooth mesoscutellum, dark ventral metasoma, and short ovipositor sheath, shorter than metasoma. The gastral tail is the shortest within the sp. gp., with Mt7 about 0.8× as long as Mt8+9. It belongs to the subgroup comprising P. ravii, P. minang sp. nov. and P. bannaensis sp. nov., characterized by a dark ventral metasoma, weak facial sculpture and a smooth mesoscutellum. It is closest to P. bannaensis sp. nov., but differs by its broader Mt7, and slightly longer ovipositor sheath. It differs from P. minang sp. nov. by its much shorter ovipositor sheath (less than metasomal length versus about 1.3–1.5× metasomal length) and by its less developed gastral tail (Mt7 about 0.8× versus 0.9–1.4× the length of Mt8+9). Male recognised by the broadly emarginate clypeal margin, 5-segmented funicle with undivided clava, and short pronotum, only about as long as the fused meso- and metanotum (1.0–1.2×). Closest to P. bannaensis sp. nov., but differing by the shorter pronotum, less punctate frontovertex, longer mesonotum relative to propodeum (1.6–1.9× versus 1.2–1.3×), and shorter fore wing remnants.

Female. Body length: 1.54–1.90 mm; ovipositor length: 0.95–1.09 mm.

Measurements (µm) [JRAS01214_0203]: Head L: 195, W: 450, H: 315; eye H: 204, L: 159; malar space L: 112; pedicel plus flagellum L: 492; fl4 L: 54, W: 46; fl8 L: 50, W: 49; POL: 148, OOL: 33. LOL: 65. Pronotum L: 155, W: 404; mesoscutum L: 196; mesoscutellum L: 218. mv: 246; stv: 67; pmv: 81. Tibia 3 L: 253. Metasoma: Mt2–Mt6 L: 804, W: 402; Mt7 L: 178, W: 67; Mt8+9 L: 239, W: 359. Body L: 1710, ovipositor sheath L: 1024.

Colour. Head and mesosoma black with bluish-greenish iridescence (Fig. 5A). Antenna yellowish, pedicel and flagellum brownish. Fore wing hyaline, venation yellowish (Fig. 5A). All legs entirely yellowish, including coxae (Fig. 5A). Gaster to Mt6 black with coppery iridescence; Mt7–Mt8+9 brownish. Ovipositor sheaths black, transversely striate. Hypopygium yellowish.

10.3897/jhr.99.205393.figure5 67A9BE1F-5A61-5667-9848-E8F0F72056A7

Philotrypesis ravii [JRAS01214_02]. A habitus B habitus, lateral view C antenna D head, frontal view E venation F head, dorsal view.

https://binary.pensoft.net/fig/1753546

Head. Head mostly smooth (Fig. 5D) with sparse punctures bearing a short seta, vertex transversely striate (Fig. 5F); lower face faintly alutaceous; scrobal depression and parascrobal area smooth. In dorsal view 2.27–2.31× as broad as long (Fig. 5F); in frontal view transverse 1.43–1.50× as broad as high (Fig. 5D). Eye height 1.17–1.28× eye length and 1.52–1.83× as long as malar space; malar sulcus distinct but shallow. Lateral ocellus closer to eye than its own transverse diameter (Fig. 5F). Clypeal margin straight (Fig. 5D). Antenna inserted just above ocular line (Fig. 5D); intertorular distance about as large as torulus diameter. POL 4.29–5.23× OOL and 1.91–2.02× LOL.

Antenna. Pedicel + flagellum L 1.06–1.13× as long as head breadth. Funicular segments, subquadrate, with sensilla arranged in one row (Fig. 5C); fl4 1.02–1.20× as long as wide; fl8 1.01–1.09× as long as wide (Fig. 5C).

Mesosoma. Pronotum dorsally smooth, transverse, 0.32–0.43× longer than wide and 0.64–0.79× as long as mesoscutum (msc) (Fig. 6B). Mesoscutum (both mesoscutal mid lobe and lateral lobe) conspicuously transversally alutaceous (Fig. 6B); notauli conspicuous, finely impressed (Fig. 6B), reaching transscutal articulation in front of scutoscutellar sutures (Fig. 6B). Axillae faintly alutaceous; mesoscutelllum (mss) smooth with 6–9 sparse punctures bearing setae (Fig. 6B), 1.10–1.18× as long as mesoscutum (msc). Prepectus smooth; upper mesepisternum (ues2) faintly engraved reticulate on its upper part, densely setose in lower third (Fig. 6A); mesopleural suture (pls) conspicuous, deep. Acron and mesepimeron smooth. Metapleuron (pl3) alutaceous (Fig. 6A). Dorsellum smooth. Propodeum smooth with a median longitudinal depression, bounded laterally by submedian longitudinal carinae (Fig. 6C); posterior propodeal carina sharp, bearing four or five anterior conspicuous carinae; two additional lateral carinae flanking propodeal spiracle: short external, long internal (Fig. 6C).

10.3897/jhr.99.205393.figure6 69CCE359-1429-59E2-849B-7542384D4DBC

Philotrypesis ravii [JRAS01214_02]. A mesosoma, lateral view B mesosoma, dorsal view C propodeum D metasoma, dorsal view E head, dorsal view F mesosoma, dorsal view.

https://binary.pensoft.net/fig/1753547

Wing. mv 3.25–3.68× longer than stv; pmv 1.21–1.22× longer than stv (Fig. 5E).

Metasoma. Mt2 smooth; Mt3-7 alutaceous at least laterally; Mt8+9 smooth (Fig. 6D). Length of Mt2–Mt6 1.89–1.95× length of reduced part of metasoma (Mt7+Mt8+9); Mt7 2.66–3.20× as long as broad and 0.75–0.79× as long as Mt8+9 L. Mt8+9 3.46–4.00× as long as broad. Ovipositor sheath 0.84–0.94× as long as metasoma (Mt2–Mt8+9 length) and 0.60–0.63× as long as body length (from head to posterior margin of Mt8+9) (Fig. 6A).

Male. Body length: 0.80–0.96 mm.

Measurements (µm) [paratype JRAS01214_0302]: Head L: 467, W: 471, eye H: 116; eye L: 87, malar space L: 60; TML (temple length): 387. Mesosoma L: 654, W: 583; pronotum L: 251; mesonotum L: 251; propodeum L: 130; fore wing remnant L: 234.

Colour. Body predominantly yellowish-brown, head and pronotum conspicuously darker, brown, mesonotum lighter. Mandible dark brown; antenna whitish, scape light brown (Fig. 6E). Tibia of all legs brown (Fig. 6F).

Head. In frontal view slightly transverse to subquadrate (Fig. 6E), 0.89–0.99× as long as wide with 7–10 medium-sized setae on vertex. Clypeal margin conspicuously emarginate (Fig. 6E). Malar space striate 0.37–0.51× the height of compound eye; temple 2.60–3.33× the height of compound eye; scrobal depression strongly and concentrically striate; frontovertex smooth; temple inconspicuously alutaceous, with sparse large punctures and a long, conspicuous sulcus posterior to eye, slightly exceeding eye length. Compound eye 0.40–0.75× as long as high. Funicle with 5 funicular segments becoming broader toward apex (Fig. 6E), and an undivided clava.

Mesosoma. 1.11–1.16 as long as broad. Pronotum 1.00–1.22× as long as fused mesonotum + metanotum (no2–no3) (Fig. 6F). Fused no2–no3 L 1.63–1.92× as long as propodeum (Fig. 6F). Propodeum, well delimitated by septa, propodeal spiracles located in the latero-posterior angle of the propodeum. Wings vestigial, fore wing remnant 0.94–1.28× as long as mesoscutum L. Protibia (tb1) (Fig. 5B) externally flattened and rugose over most of its surface; bearing 4–5 short apical or subapical spines in one row. Mesotibia (tb2) (Fig. 5B) externally with 9–10 short spines in 1–2 rows. Metatibia (tb3) with 14–16 short spines in 2–3 rows; first tarsomere digitally expanded, bearing a pair of long setae; second tarsomere slightly enlarged.

Metasoma. Metasoma weakly sclerotized, dorsoventrally flattened, each tergite with a row of setae.

 Host.

The species was reared from figs of Ficus tinctoria subsp. gibbosa (Blume) Corner (subgen. Terega), a senior synonym of Ficus tinctoria ssp. parasitica Willd. (Corner). This fig tree was pollinated by Kradibia rutherforti subsp. indica (Abdurahiman & Joseph, 1967).

 Distribution.

India (Karnataka, Kerala).

 Animalia Hymenoptera Pteromalidae A74E3DFC-097B-5B6E-8362-BA40F9CEAB2B Philotrypesis bannaensis https://zoobank.org/CB6ACF15-AC1C-48B3-B9B8-6F9E1CF9A575 Li & Rasplus sp. nov. Figs 7B–F , 8A–D (♀), 7A, 8E, F (♂) Type material.

Holotype • ♀: China, Xishuangbanna, Xishuangbanna Tropical Botanical Garden [21.928 101.256], xi.2005, Proffit M. leg [JRAS01451_0202], deposited in XTBG, Menglun. Condition: Glued on the right side to a rectangular card; in perfect condition, complete. Paratypes: • 7♀, 8♂, same data as holotype. • 21♀, same locality as holotype, 3.ix.2001 and 19.i.2002, Peng Y.Q. [JRAS00842_02]; • 5♀, 5♂, same locality but [21.929 101.257] (500 m), i.2023, Li Y. leg. [LYRA00012_02]; paratype specimens deposited in XTBG and CBGP.

 Diagnosis.

Female recognised by the weakly sculptured lower face and parascrobal area, antenna inserted just above the ocular line, straight clypeal margin, smooth mesoscutellum, dark ventral metasoma and short ovipositor sheath. The gastral tail is moderately developed, with Mt7 about same length as Mt8+9. Like P. ravii and P. minang sp. nov., it possesses a dark ventral metasoma, weak facial sculpture and a smooth mesoscutellum. It is morphologically intermediate between P. ravii and P. minang sp. nov., combining the short ovipositor sheath of the former with a more elongate gastral tail approaching the latter. It differs from P. ravii by its transverse fl8 and more elongate Mt7 (about 1.02–1.03× versus 0.8× the length of Mt8+9), and from P. minang sp. nov. by its much shorter ovipositor sheath (shorter than metasoma versus about 1.3–1.5× metasomal length). Male recognised by the broadly emarginate clypeal margin, 5-segmented funicle with undivided clava, frontovertex bearing numerous punctures, elongate pronotum (1.4–1.5× fused meso- and metanotum), and mesonotum only slightly longer than propodeum (1.2–1.3×). Closest to P. ravii, but differing by the longer pronotum, shorter mesonotum relative to propodeum, punctate frontovertex, and more developed fore wing remnants.

Female. Body length: 1.70–1.89 mm; ovipositor length: 0.90–0.94 mm.

Measurements (µm) [Holotype, JRAS01451_0202]: Head L: 170, W: 375, H: 281; eye H: 221, L: 152; malar space: 123; pedicel plus flagellum L: 415; fl4 L: 47, W: 41; fl8 L: 40, W: 44; POL: 143, OOL: 28, POL: 70. Pronotum L: 110, W: 121; mesoscutum L: 176; mesoscutellum L: 204. mv: 245; stv: 69; pmv: 77. Tibia 3 L: 301. Metasoma: Mt2–Mt6 L: 733, W: 373; Mt7 L: 203, W: 58; Mt8+9 L: 199, W: 69; ovipositor sheath L: 937.

Colour. Head and mesosoma black with greenish iridescence (Fig. 7B). Antenna (flagellum and pedicel) brownish, scape lighter (Fig. 7C). Fore wing hyaline, venation yellowish (Fig. 7F). All legs entirely yellowish, including coxae (Fig. 7B). Gaster to Mt6 black with coppery iridescence (Fig. 8D); Mt7–Mt8+9 brownish. Ovipositor sheaths black, transversely striate. Hypopygium yellowish.

10.3897/jhr.99.205393.figure7 77772313-9B82-5465-8F4A-CFF32635C6A2

Philotrypesis bannaensis sp. nov. [JRAS01451_02]. A habitus B habitus, lateral view C antenna D head, frontal view E venation F head, dorsal view.

https://binary.pensoft.net/fig/1753548

Head. Head mostly smooth (Fig. 7D) with sparse small punctures bearing a short seta, vertex transversely striate (Fig. 7F); lower face faintly alutaceous; scrobal depression smooth. In dorsal view 2.17–2.20× as broad as long (Fig. 7F); in frontal view transverse 1.33–1.48× as broad as high (Fig. 7D). Eye height 1.09–1.19× eye length and 1.32–1.34× as long as malar space; malar sulcus distinct. Lateral ocellus closer to eye than its own transverse diameter (Fig. 7F). Clypeal margin straight (Fig. 7D). Antenna inserted just above ocular line, (Fig. 7D); intertorular distance about as large as torulus diameter. POL 4.70–5.17× OOL and 1.83–2.03× LOL.

Antenna. Pedicel + flagellum L 1.11–1.12× as long as head breadth. Funicular segments, longer than wide, with sensilla arranged in one row (Fig. 7C); fl4 1.14–1.24× as long as wide; fl8 0.91–0.96× as long as wide (Fig. 7C).

Mesosoma. Pronotum dorsally smooth to the exception of a few striae on posterior edge, laterally alutaceous, pronotum 0.30–0.36× longer than wide and 0.63–0.73× as long as mesoscutum (Fig. 8B). Mesoscutum (both mesoscutal mid lobe and lateral lobe) conspicuously transversally alutaceous; notauli weakly impressed, reaching transscutal articulation in front of scutoscutellar sutures (Fig. 8B). Axillae alutaceous; mss smooth with 8–10 sparse punctures (Fig. 8B), 1.16–1.19× as long as msc. Prepectus smooth; upper mesepisternum (ues2) faintly engraved reticulate on its upper part, setose in lower third (Fig. 8A); mesopleural suture (pls) conspicuous, deep (Fig. 8A). Acron and mesepimeron smooth. Metapleuron (pl3) alutaceous (Fig. 8A). Dorsellum smooth. Propodeum smooth with a median longitudinal depression, bounded laterally by submedian longitudinal carinae (Fig. 8C); posterior propodeal carina sharp, bearing four or five anterior conspicuous carinae; two additional lateral carinae flanking propodeal spiracle: short external, long internal (Fig. 8C).

10.3897/jhr.99.205393.figure8 8C41F1A8-17CF-5A9B-A4D9-6CDA0C6FF4F8

Philotrypesis bannaensis sp. nov. [JRAS01451_02]. A mesosoma, lateral view B mesosoma, dorsal view C propodeum D metasoma, dorsal view E head, dorsal view F mesosoma, dorsal view.

https://binary.pensoft.net/fig/1753549

Wing. mv 3.54× longer than stv; pmv 1.10–1.11× longer than stv.

Metasoma. Mt2 smooth; Mt3-7 alutaceous, sculpture becoming stronger on last Mt; Mt8+9 smooth (Fig. 8D). Length of Mt2–Mt6 1.70–1.83× length of reduced part of metasoma (Mt7+Mt8+9); Mt7 3.51–3.97× as long as broad and 1.02–1.03× as long as MT8+9. Mt8+9 2.89–2.93× as long as broad. Ovipositor sheath 0.83× as long as metasoma (Mt2–Mt8+9 length) and 0.58–0.60× as long as body length (from head to posterior margin of Mt8+9) (Fig. 7B).

Male. Body length: 0.80–0.96 mm.

Measurements (µm) [paratype JRAS01451_0303]: Head L: 420, W: 452, eye H: 80; eye L: 53, malar space L: 43; TML (temple length): 298. Mesosoma L: 585, W: 505; pronotum L: 261; mesonotum L: 186; propodeum L: 149; fore wing remnant L: 107.

Colour. Body yellowish-brown (Fig. 7A); metasoma yellowish, lighter than head and mesosoma. Mandible brown (Fig. 8E); flagellum whitish (Fig. 8E). All tibia brown (Fig. 7A).

Head. In frontal view slightly transverse to subquadrate (Fig. 8E), 0.93–0.97× as long as wide with 7–10 medium-sized setae on vertex. Clypeal margin conspicuously emarginate. Malar space striate 0.44–0.65× the height of compound eye; temple 3.01–3.73× the height of compound eye; scrobal depression strongly and concentrically striate; frontovertex smooth with multiple punctures; temple inconspicuously alutaceous, with sparse large punctures and a long, conspicuous sulcus posterior to eye, slightly exceeding eye length. Compound eye 0.56–0.67× as long as high. Funicle with 5 funicular segments becoming broader toward apex (Fig. 7A), and an undivided clava.

Mesosoma. 1.12–1.27 as long as broad. Pronotum 1.40–1.46× as long as fused meso- + metanotum (Fig. 8F). Fused no2–no3 (mesonotum + metanotum L) 1.20–1.25× as long as propodeum (Fig. 8F). Propodeum, well delimitated by septa even anteriorly, propodeal spiracles located in the latero-posterior angle of the propodeum. Wings vestigial, fore wing remnant 2.17–2.83× as long as mesoscutum L. Protibia (tb1) (Fig. 7A) externally flattened and rugose over most of its surface; bearing 4–5 short apical or subapical spines in one row. Mesotibia (tb2) (Fig. 7A) externally with 12–13 short spines in 2–3 rows. Metatibia (tb3) with 15–16 short spines in 3–4 rows (Fig. 7A); first tarsomere digitally expanded, bearing a pair of long setae; second tarsomere slightly enlarged (Fig. 7A).

Metasoma. Metasoma weakly sclerotized, dorsoventrally flattened, each tergite with one-two rows of setae.

 Host.

The type series was reared from syconia of Ficus tinctoria subsp. gibbosa (Blume) Corner. This subspecies is distributed from Sri Lanka and India through southern China and mainland Southeast Asia to western Malesia (including Sumatra, Java and Borneo) and is pollinated, in the type locality, by Kradibia gibbosae (Hill, 1967).

 Distribution.

Oriental region, sino-himalayan subregion. China.

 Etymology.

The specific epithet bannaensis sp. nov. refers to Xishuangbanna Autonomous Prefecture, Yunnan Province (China) often abbreviated as “Banna”, where the type series was collected.

 Animalia Hymenoptera Pteromalidae 09AE608A-55AB-58CB-A67F-E59C6D411CBD Philotrypesis minang https://zoobank.org/FAF45711-746D-4186-B198-A34B0EE631DC Li & Rasplus sp. nov. Figs 9A , 9C–F , 10A–D (♀), 9B, 10E, F (♂) Type material.

Holotype • ♀: Indonesia, Sumatra, Bukittinggi [-0.304929 100.358216] 6.iv.1990 Tanjung P. [JRAS06684_0202]. Condition: glued by right side on rectangular card; perfect state, entire. Paratypes • 13 ♀, 1 ♂, same data as holotype. All type specimens deposited at CBGP, Montpellier.

 Diagnosis.

Female recognised by the weakly sculptured lower face and parascrobal area, antenna inserted above the ocular line, slightly convex clypeal margin, smooth mesoscutellum, dark ventral metasoma, and strongly developed gastral tail. The ovipositor sheath is remarkably long, about 1.3–1.5× metasomal length, and Mt7 is about 0.9–1.4× as long as Mt8+9. The propodeum bears complete submedian carinae and four conspicuous anterior carinae. It belongs to the dark-bodied subgroup (with P. ravii, P. minang sp. nov., P. bannaensis sp. nov.) sharing a smooth mesoscutellum and weak facial sculpture, but differs from P. ravii and P. bannaensis sp. nov. by its much longer ovipositor sheath (about 1.3–1.5× versus less than metasomal length) and more strongly developed gastral tail (Mt7 about 0.9–1.4× versus 0.8× and 1.0× the length of Mt8+9, respectively). Although approaching P. arfak sp. nov. in ovipositor length and tail development, it differs by the dark ventral metasoma, smooth mesoscutellum, weak facial sculpture and complete propodeal carination. Male recognised by the elongate head (longer than broad), very long vertex setae, 5-segmented funicle with subdivided clava, and extremely elongate pronotum, nearly twice as long as the fused meso- and metanotum. Closest to P. paiwan sp. nov., but differing by the elongate head, 5-segmented rather than 4-segmented funicle, broadly emarginate rather than triangularly emarginate clypeus, and shorter mesonotum relative to propodeum.

Female. Body length: 1.56–1.75 mm; ovipositor length: 1.47–1.73 mm.

Measurements (µm) [Holotype JRAS06684_0201]: Head L: 167, W: 390, H: 255; eye H: 167, L: 145; malar space: 127; pedicel plus flagellum L: 419; fl4 L: 45, W: 41; fl8 L: 38, W: 34; POL: 136, OOL: 37. LOL: 73. Pronotum L: 105, W: 294; mesoscutum L: 193; mesoscutellum L: 209. mv: 245; stv: 66; pmv: 87. Tibia 3 L: 239. Metasoma: Mt2–Mt6 L: 727, W: 378; Mt7 L: 209, W: 55; Mt8+9 L: 234, W: 48. Body L: 1713; ovipositor sheath L: 1732.

Colour. Head and mesosoma with slight greenish iridescence (Fig. 9A). Antenna entirely brownish, scape lighter. Fore wing hyaline, venation yellowish (Fig. 9E). All legs entirely yellowish, including coxae. Gaster ventrally and dorsally black, with coppery iridescence; Mt7–Mt8+9 brownish. Ovipositor sheaths black, transversely striate. Hypopygium yellowish.

10.3897/jhr.99.205393.figure9 FCF7133F-F59E-51CF-80B8-829A783BBCA2

Philotrypesis minang sp. nov. [JRAS06684_02]. A habitus B habitus, lateral view C antenna D head, frontal view E venation F head, dorsal view.

https://binary.pensoft.net/fig/1753550

Head. Head mostly smooth (Fig. 9D) with sparse punctures bearing a short seta; lower face inconspicuously alutaceous; scrobal depression smooth. In dorsal view 2.34–2.41× as broad as long, vertex transversely striate (Fig. 5F); in frontal view transverse 1.38–1.53× as broad as high (Fig. 9D). Eye height 1.15–1.38× eye length and 1.00–1.17× as long as malar space; malar sulcus distinct but shallow. Lateral ocellus separated from the eye by a distance equal to its own transverse diameter (Fig. 9F). Clypeal margin slightly convex (Fig. 9D). Antenna inserted above ocular line (Fig. 9D); intertorular distance about as large as torulus diameter. POL 3.71–4.17× OOL and 1.85–2.00× OOL.

Antenna. Pedicel + flagellum L 1.00–1.16× as long as head breadth. Funicular segments, slightly elongated, with sensilla arranged in one row (Fig. 9C); fl4 1.14–1.46× as long as wide; fl8 1.03–1.21× as long as wide (Fig. 9C).

Mesosoma. Pronotum dorsally smooth, transverse, 0.36–0.40× longer than wide and 0.54–0.84× as long as mesoscutum (msc) (Fig. 10B). Mesoscutal mid lobe conspicuously alutaceous (Fig. 10B), lateral lobe nearly smooth, faintly alutaceous near notauli; notauli conspicuous, finely impressed (Fig. 10B), reaching transscutal articulation slightly internal of scutoscutellar sutures (Fig. 10B). Axillae weakly alutaceous; mesoscutelllum (mss) smooth with 6–9 sparse punctures bearing setae (Fig. 10B, C), 1.06–1.28× as long as mesoscutum (msc). Prepectus smooth; upper mesepisternum (ues2) smooth, densely setose on lower third (Fig. 10A); mesopleural suture (pls) conspicuous, deep. Acron and mesepimeron smooth. Metapleuron (pl3) alutaceous (Fig. 10A). Dorsellum smooth. Propodeum smooth with a median area, bounded laterally by complete submedian longitudinal carinae (Fig. 10C); posterior propodeal carina sharp and bearing, in addition to submedian carinae, four conspicuous anterior carinae; two additional lateral carinae flanking propodeal spiracle: short external, long internal (Fig. 10C).

10.3897/jhr.99.205393.figure10 28D09664-68E0-5213-B624-07A21461C3C0

Philotrypesis minang sp. nov. [JRAS06684_02]. A mesosoma, lateral view B mesosoma, dorsal view C propodeum D metasoma, dorsal view E head, dorsal view F mesosoma, dorsal view.

https://binary.pensoft.net/fig/1753551

Wing. mv 3.31–3.70× longer than stv; pmv 1.00–1.31× longer than mv (Fig. 5E).

Metasoma. Mt2 smooth; Mt3-7 alutaceous (Fig. 10D); Mt8+9 smooth. Length of Mt2–Mt6 1.43–1.64× length of reduced part of metasoma (Mt7+Mt8+9); Mt7 3.80–5.25× as long as broad and 0.89–1.38× as long as Mt8+9 L. Mt8+9 3.57–4.84× as long as broad. Ovipositor sheath 1.33–1.48× as long as metasoma (Mt2–Mt8+9 length) and 0.90–1.02× as long as body length (from head to posterior margin of Mt8+9) (Fig. 9A).

Male. Body length: 0.83 mm.

Measurements (µm) [paratype JRAS06684_0501]: Head L: 453, W: 427, eye H: 73, L: 60, malar space L: 52; TML (temple length): 297. Mesosoma L: 584, W: 495; pronotum L: 333; mesonotum L: 177; propodeum L: 125; fore wing remnant L: 234.

Colour. Body yellowish-brown (Fig. 9B); Lower face and mandible brown (Fig. 10E); flagellum whitish (Fig. 9B). All tibia brown darker than body (Fig. 9B).

Head. In frontal view slightly elongated (Fig. 10E), 1.18× as long as broad with 8 long-sized setae on vertex (about as long as head length). Clypeal margin conspicuously emarginate (Fig. 10E). Malar space striate 0.71× the height of compound eye; temple 4.07× the height of compound eye (Fig. 9B); scrobal depression strongly and concentrically striate; frontovertex smooth with sparse puncture (Fig. 10E).; temple inconspicuously alutaceous, with sparse large punctures and a long, conspicuous sulcus posterior to eye, about 1.5× longer than eye height (Fig. 9B). Compound eye 0.84× as long as high. Funicle with 5 funicular segments becoming broader toward apex (Fig. 9B), and a subdivided clava.

Mesosoma. 1.18 as long as broad. Pronotum 1.88× as long as fused meso- + metanotum (no2–no3) (Fig. 10F). Fused no2–no3 1.42× as long as propodeum (Fig. 10F). Propodeum, well delimitated by septa, except in its anterior margin where septum vanished, propodeal spiracles located in the latero-posterior angle of the propodeum. Wings vestigial, fore wing remnant 1.32× as long as mesoscutum L. Protibia (tb1) (Fig. 9B) externally flattened and rugose over most of its surface; bearing 4–5 short apical or subapical spines in one row. Mesotibia (tb2) (Fig. 9B) externally with 8 short spines in 2 rows. Metatibia (tb3) with 12 short spines in 3 rows (Fig. 9B); first tarsomere digitally expanded, bearing a pair of long setae; second tarsomere slightly enlarged.

Metasoma. Metasoma weakly sclerotized, each tergite with a row of setae.

 Host.

The species was also reared from figs of Ficus tinctoria subsp. gibbosa (Blume) Corner, pollinated by an undescribed species of Kradibia (sp046 in our database) closely related to Kradibia rutherforti subsp. rutherforti (Waterston, 1920).

 Distribution.

Oriental region, Indonesia (Sumatra).

 Etymology.

The specific epithet minang is a noun in apposition referring to the Minang (Minangkabau) people of West Sumatra.

 Animalia Hymenoptera Pteromalidae 88773753-90A3-5470-A0F8-B9788EED390B Philotrypesis paiwan https://zoobank.org/7A34E037-48D5-4E5F-83FA-2BDE4F299C3C Li & Rasplus sp. nov. Figs 11A , 11C–F , 12A–D (♀), 11B, 12E, F (♂) Type material.

Holotype • 1♀: China, Taiwan, Kenting, Hengchun TBG [21.964 120.813], 8.ix.2008, Cruaud A. & Rasplus J.Y. leg. [JRAS02406_0201], deposited in CBGP, Montpellier. Condition: glued by right side on rectangular card; perfect state, entire. Paratypes: • 3♀, 1♂, same data as holotype. Paratype specimens deposited in CBGP, Montpellier.

 Diagnosis.

Female recognised by the entirely yellow antennae, strongly alutaceous lower face and parascrobal area, antenna inserted distinctly above the ocular line, regularly convex clypeal margin, orange ventral metasoma, and densely sculptured mesoscutellum with elongate meshes and a median longitudinal declivity. The propodeum is distinctive in possessing winding submedian carinae and a median depression crossed by transverse septa. The ovipositor sheath slightly exceeds metasomal length and Mt7 is moderately elongate, being about 1.0–1.1× as long as Mt8+9. Together with P. arfak sp. nov., it forms a distinctive subgroup characterized by an orange ventral metasoma, strong facial sculpture and a sculptured mesoscutellum. It differs from P. arfak sp. nov. by its entirely yellow antennae, denser mesoscutellar sculpture, more elaborate propodeal sculpture, shorter ovipositor sheath (about 1.2× versus 1.4–1.5× metasomal length) and less developed gastral tail (Mt7 about 1.0–1.1× versus 1.2–1.4× the length of Mt8+9). It differs from P. ravii, P. minang sp. nov. and P. bannaensis sp. nov. by the orange ventral metasoma, strongly sculptured lower face and sculptured mesoscutellum.

Male recognised by the triangularly emarginate clypeal margin, 4-segmented funicle with subdivided clava, large eyes (eye length about 0.8× eye height), and extremely elongate pronotum (about 1.8× as long as mesonotum + metanotum). Closest to P. arfak sp. nov., but differing by the much longer pronotum and mesonotum, and by the larger eyes.

Female. Body length: 2.32–2.35 mm; ovipositor length: 1.61–1.66 mm.

Measurements (µm) [holotype JRAS02406_0201]: Head L: 217, W: 477, H: 503; eye H: 221, L: 187; malar space: 112; pedicel plus flagellum L: 537; fl4 L: 57, W: 45; fl8 L: 59, W: 42; POL: 169, OOL: 39, LOL: 88. Pronotum L: 165, W: 447; mesoscutum L: 214; mesoscutellum L: 230. mv: 304; stv: 79; pmv: 86. Tibia 3 L: 330. Metasoma: Mt2–Mt6 L: 888, W: 461; Mt7 L: 252, W: 63; Mt8+9 L: 253, W: 69; ovipositor sheath L: 1657.

Colour. Head and mesosoma black with greenish iridescence (Figs 11A, 11D, 12B). Antenna (flagellum, pedicel and scape) yellowish (Fig. 11C). Fore wing hyaline, venation yellowish (Fig. 11E). All legs entirely yellowish, including coxae (Fig. 11A). Gaster to Mt6 anteriorly yellowish-orange, dorsally black with bluish-greenish iridescence (Fig. 11A); ventrally entirely yellowish-orange; Mt7–Mt8+9 blackish dorsally, yellowish ventrally. Ovipositor sheaths black, transversely striate. Hypopygium yellowish.

10.3897/jhr.99.205393.figure11 2B9E5023-F627-5744-BDFE-73841B2E6BB2

Philotrypesis paiwan sp. nov. [JRAS02406_02]. A habitus B habitus, lateral view C antenna D head, frontal view E venation F head, dorsal view.

https://binary.pensoft.net/fig/1753552

Head. Lower face and parascrobal area conspicuously alutaceous (Fig. 11D) with punctures bearing a short seta, vertex transversely striate (Fig. 11F); scrobal depression and frontovertex smooth. Head in dorsal view 2.23–2.32× as broad as long (Fig. 11F); in frontal view transverse, 1.46–1.55× as broad as high (Fig. 11D). Eye height 1.18–1.25× eye length and 1.42–1.68× as long as malar space; malar sulcus distinct. Lateral ocellus closer to eye than its own transverse diameter (Fig. 11F). Clypeal margin regularly convex (Fig. 11D). Antenna inserted distinctly above the ocular line (Fig. 11D); intertorular distance about as large as torulus diameter. POL 4.32–4.54× OOL and 1.93–2.03× LOL.

Antenna. Pedicel + flagellum L 1.05–1.07× as long as head breadth. Funicular segments, longer than wide, with sensilla arranged in one row (Fig. 11C); fl4 1.06–1.25× as long as wide and fl8 1.13–1.41× as long as wide.

Mesosoma. Pronotum dorsally smooth to the exception of a few striae on posterior edge, laterally alutaceous (Fig. 12B); pronotum transverse 0.36–0.37× longer than wide and 0.72–0.77× as long as mesoscutum (Fig. 12B). Mesoscutum (both mesoscutal mid lobe and lateral lobe) conspicuously transversally alutaceous (Fig. 12B). Axillae densely alutaceous; mesoscutellum longitudinally alutaceous, medially with denser elongated meshes and a longitudinal declivity (Fig. 12B), mss 1.07–1.08× as long as msc. Prepectus smooth; upper mesepisternum (ues2) faintly engraved reticulate on its upper part, densely setose in lower third (Fig. 12A); mesopleural suture (pls) conspicuous, deep (Fig. 12A). Acron and mesepimeron smooth. Metapleuron (pl3) alutaceous (Fig. 12A). Dorsellum smooth. Propodeum smooth with a median longitudinal depression, with transverse septa, and laterally bounded by winding submedian longitudinal carinae (Fig. 12C); posterior propodeal carina sharp, bearing four or five anterior conspicuous carinae; two additional lateral carinae flanking propodeal spiracle: short external, long internal (Fig. 12C).

10.3897/jhr.99.205393.figure12 3B6875B6-9D6F-57F1-A651-9E0B848C6562

Philotrypesis paiwan sp. nov. [JRAS02406_02]. A mesosoma, lateral view B mesosoma, dorsal view C propodeum D metasoma, dorsal view E head, dorsal view F mesosoma, dorsal view.

https://binary.pensoft.net/fig/1753553

Wing. mv 3.86–4.06× longer than stv; pmv 1.10–1.13× longer than mv (Fig. 11E).

Metasoma. Mt2 smooth; Mt3-7 alutaceous at least laterally; Mt8+9 smooth. Length of Mt2–Mt6 1.57–1.76× length of reduced part of metasoma (Mt7+Mt8+9); Mt7 3.56–4.00× as long as broad and 1.00–1.14× as long as Mt8+9 L. Mt8+9 3.21–3.67× as long as broad. Ovipositor sheath 1.17–1.19× as long as metasoma (Mt2–Mt8+9 length) and 0.80–0.81× as long as body length (from head to posterior margin of Mt8+9) (Fig. 11A).

Male. Body length: 0.97 mm.

Measurements (µm) [paratype JRAS02406_0601]: Head L: 484, W: 521, eye H: 83; eye L: 70, malar space L: 62; TML (temple length): 281. Mesosoma L: 719, W: 630; pronotum L: 406; mesonotum L: 224; propodeum L: 115; fore wing remnant L: 234.

Colour. Body yellowish-brown. Mandible dark brown; antenna whitish (Fig. 12E). All tibia brown (Fig. 11B).

Head. In frontal view slightly transverse to subquadrate (Fig. 12E), 0.93× as long as wide with 6 medium-sized setae on vertex. Clypeal margin conspicuously, triangularly emarginate (Fig. 12E). Malar space striate 0.75× the height of compound eye; temple 3.38× the height of compound eye; scrobal depression strongly and concentrically striate; frontovertex smooth with sparse puncture; temple inconspicuously alutaceous, with sparse large punctures and a long, conspicuous sulcus posterior to eye, about 1.5× longer than eye height. Compound eye 0.84× as long as high. Funicle with 4 funicular segments becoming broader toward apex (Fig. 12E), and a subdivided clava.

Mesosoma. 1.15× as long as broad. Pronotum 1.81× as long as fused meso- + metanotum (Fig. 12F). Fused no2–no3 (mesonotum + metanotum L) 1.95× as long as propodeum (Fig. 12F). Propodeum, well delimitated by septa, except in its anterior margin where septum vanished in the middle; propodeal spiracles located in the latero-posterior angle of the propodeum. Wings vestigial, fore wing remnant 1.05× as long as mesoscutum L. Protibia (tb1) (Fig. 11B) externally flattened and rugose over most of its surface; bearing 4–5 short apical or subapical spines in one row. Mesotibia (tb2) (Fig. 11B) externally with 11 short spines in 2 rows. Metatibia (tb3) with 15 short spines in 3 rows (Fig. 11B); first tarsomere digitally expanded, bearing a pair of long setae; second tarsomere slightly enlarged (Fig. 11B).

Metasoma. Metasoma weakly sclerotized, each tergite with a row of setae.

 Host.

The type series was reared from syconia of Ficus tinctoria subsp. tinctoria G. Forst., a widespread Indo-Pacific fig extending from Taiwan and eastern Malesia to New Guinea, the Pacific Islands and northern Australia. The figs are pollinated by an undescribed species of Kradibia (sp026 in our database), occurring from Taiwan to Solomon Islands.

 Distribution.

Oriental region, sino-himalayan subregion. Taiwan, China.

 Etymology.

The specific epithet paiwan is a noun in apposition honoring the Paiwan people, one of the major Indigenous groups of southern Taiwan, China. The type series was collected in the Hengchun Tropical Botanical Garden, situated within the traditional Paiwan region of the Hengchun Peninsula.

 Animalia Hymenoptera Pteromalidae 8A4AC569-3FCE-5BF2-B1DF-02F453A9916B Philotrypesis arfak https://zoobank.org/3666077F-9893-4C7C-A42C-1AA8313D5113 Li & Rasplus sp. nov. Figs 13A , 13C–F; 14A–D (♀); 13B, 14E, F (♂) Type material.

Holotype • ♀: Indonesia: West Papua, road from Manokwari to Maruni [-0.925 134.044], 11.v. 2008, Cruaud A.; Ubaidillah R. & Rasplus J.Y. leg. [JRAS02350_0310], deposited in MZB, Bogor. Condition: glued by right side on rectangular card; perfect state, entire. Paratypes: • 22 ♀, 28 ♂, same data as holotype. Paratype specimens deposited in MZB, Bogor and CBGP, Montpellier.

 Diagnosis.

Female recognized by the alutaceous lower face and parascrobal area, antenna inserted on the ocular line, orange ventral metasoma extending dorsally above the midline of the gaster, weakly sculptured mesoscutellum with a broad median longitudinal declivity, and long ovipositor sheath. The propodeum is distinctive in possessing discontinuous submedian carinae and only two conspicuous anterior carinae. The gastral tail is the most strongly developed within the P. ravii sp. gp., with Mt7 about 1.2–1.4× as long as Mt8+9. Together with P. paiwan sp. nov., it belongs to the subgroup characterized by strong facial sculpture, orange metasomal coloration and a sculptured mesoscutellum. It differs from P. paiwan sp. nov. by its weaker mesoscutellar sculpture, simplified propodeal ornementation, markedly longer ovipositor sheath (about 1.4–1.5× versus 1.2× metasomal length) and more elongate gastral tail (Mt7 about 1.2–1.4× versus 1.0–1.1× the length of Mt8+9). Although approaching P. minang sp. nov. in ovipositor length and tail development, it differs by the orange metasoma, sculptured face and mesoscutellum, and reduced propodeal carination. Male recognised by the triangularly emarginate clypeal margin, long malar space (0.5–0.8× eye height), long temples (3.3–4.0× eye height), 4-segmented funicle with a 3-segmented clava, and pronotum only moderately longer than the mesonotum + metanotum (1.1–1.6×). Closest to P. paiwan sp. nov., but differing by the shorter pronotum, smaller eyes (eye length only 0.5–0.6× eye height), and less elongate mesonotum relative to propodeum.

Female. Body length: 1.94–2.23 mm; ovipositor length: 1.80–1.97 mm.

Measurements (µm) [Holotype, JRAS02350_0301]: Head L: 205, W: 425, H: 287; eye H: 193, L: 186; malar space: 116; pedicel plus flagellum L: 489; fl4 L: 50, W: 40; fl8 L: 42, W: 40; POL: 140, OOL: 36, POL: 64. Pronotum L: 137, W: 367; mesoscutum L: 194; mesoscutellum L: 213. mv: 263; stv: 72; pmv: 77. Tibia 3 L: 306. Metasoma: Mt2–Mt6 L: 835, W: 418; Mt7 L: 315, W: 54; Mt8+9 L: 226, W: 63; ovipositor sheath L: 1973.

Colour. Head and mesosoma black with greenish iridescence (Figs 13A, 14B). Antenna (flagellum, pedicel and scape) brownish (Fig. 13C). Fore wing hyaline, venation yellowish (Fig. 13E). All legs entirely yellowish light, including coxae (Fig. 13A). Gaster to Mt6 anteriorly yellowish-orange, dorsally black with bluish-greenish iridescence (Fig. 13A); ventrally entirely yellowish-orange, the yellow coloration extending above the median line; Mt7–Mt8+9 blackish dorsally, yellowish ventrally. Ovipositor sheaths black, transversely striate. Hypopygium yellowish.

10.3897/jhr.99.205393.figure13 604E995E-D8DE-52AA-93AE-A4179F7F992F

Philotrypesis arfak sp. nov. [JRAS02350_03]. A habitus B habitus, lateral view C antenna D head, frontal view E venation F head, dorsal view.

https://binary.pensoft.net/fig/1753554

Head. Lower face and parascrobal area conspicuously alutaceous (Fig. 13D) with punctures bearing a short seta, vertex with few transverse striae (Fig. 13F); scrobal depression and frontovertex smooth. In dorsal view 2.08–2.39× as broad as long (Fig. 13F); in frontal view transverse, 1.39–1.48× as broad as high (Fig. 13D). Eye height 1.04–1.26× eye length and 1.60–1.67× as long as malar space; malar sulcus distinct. Lateral ocellus closer to eye than its own transverse diameter (Fig. 13F). Clypeal margin slightly convex (Fig. 13D). Antenna inserted on ocular line, (Fig. 13D). Intertorular distance equal to diameter of torulus. POL 3.33–4.40× OOL and 1.87–2.19× LOL.

Antenna. Pedicel + flagellum L 0.91–1.15× as long as head breadth. Funicular segments, longer than wide, with sensilla arranged in one row (Fig. 13C); fl4 1.04–1.26× as long as wide and fl8 1.05–1.11× as long as wide (Fig. 13C).

Mesosoma. Pronotum dorsally smooth to the exception of a few striae on posterior edge, laterally alutaceous (Fig. 14B); pronotum transverse 0.37–0.43× longer than wide and 0.71–0.95× as long as mesoscutum (Fig. 14B). Mesoscutum (both mesoscutal mid lobe and lateral lobe) conspicuously transversally alutaceous (Fig. 14B); notauli conspicuous, finely impressed (Fig. 14B); axillae and mesoscutellum weakly alutaceous medially nearly smooth with large longitudinal declivity (Fig. 14B), mss 1.10–1.32× as long as msc. Prepectus smooth; upper mesepisternum (ues2) faintly engraved reticulate on its upper part, densely setose in lower third (Fig. 14A); mesopleural suture (pls) conspicuous, deep (Fig. 14A). Acron and mesepimeron smooth. Metapleuron (pl3) alutaceous (Fig. 14A). Dorsellum smooth. Propodeum smooth with a median longitudinal depression, and laterally bounded by submedian and discontinuous longitudinal carinae (Fig. 14C); posterior propodeal carina sharp, bearing two anterior conspicuous carinae; two additional lateral carinae flanking propodeal spiracle: short external, long internal (Fig. 14C).

10.3897/jhr.99.205393.figure14 75AC8FE1-FC85-505A-A320-C8BED87E386A

Philotrypesis arfak sp. nov. [JRAS02350_03]. A mesosoma, lateral view B mesosoma, dorsal view C propodeum D metasoma, dorsal view E head, dorsal view F mesosoma, dorsal view.

https://binary.pensoft.net/fig/1753555

Wing. mv 3.67–4.02× longer than stv; pmv 1.07–1.32× longer than stv (Fig. 13E).

Metasoma. Mt2 smooth; Mt3-7 alutaceous at least laterally, the sculpture becoming stronger on posterior tergite; Mt8+9 smooth (Fig. 14D). Length of Mt2–Mt6, 1.45–1.67× length of reduced part of metasoma (Mt7+Mt8+9). Mt7 4.71–5.78× as long as broad and 1.21–1.39× as long as Mt8+9. Mt8+9 3.51–3.60× as long as broad. Ovipositor sheath 1.41–1.53× as long as metasoma (Mt2–Mt8+9 length) and 1.00–1.07× as long as body length (from head to posterior margin of Mt8+9).

Male. Body length: 0.79–0.94 mm.

Measurements (µm) [paratype JRAS02350_0501]: Head L: 407, W: 461; eye H: 72, L: 33; malar space L: 56; TML (temple length): 289. Mesosoma L: 651, W: 590; pronotum L: 299; mesonotum L: 209; propodeum L: 125; fore wing remnant L: 230.

Colour. Body entirely yellowish-brown excepted mandible and all tibiae dark brown (Fig. 13B); antenna whitish (Fig. 14E).

Head. In frontal view slightly transverse to subquadrate (Fig. 14E), 0.83–0.91× as long as wide with 6–8 long and setae (as long as head) on vertex. Clypeal margin conspicuously, triangularly emarginate (Fig. 14E). Malar space striate 0.54–0.84× the height of compound eye; temple 3.32–4.00× the height of compound eye; scrobal depression strongly and concentrically striate; frontovertex smooth with sparse large punctures; temple inconspicuously alutaceous, with sparse large punctures and a long, conspicuous sulcus posterior to eye, slightly exceeding eye length. Compound eye 0.47–0.59× as long as high. Funicle with 4 funicular segments becoming broader toward apex (Fig. 14E), and a three-segmented clava.

Mesosoma. 1.10–1.25 as long as broad. Pronotum 1.14–1.56× as long as fused meso- + metanotum (no2+no3) (Fig. 14F). Fused no2–no3 L 1.67–1.83× as long as propodeum (Fig. 14F). Propodeum, well delimitated by septa, propodeal spiracles located in the latero-posterior angle of the propodeum. Wings vestigial, fore wing remnant 0.98–1.10× as long as mesoscutum L. Protibia (tb1) (Fig. 13B) externally flattened and rugose over most of its surface; bearing 4–5 short apical or subapical spines in one row. Mesotibia (tb2) (Fig. 13B) externally with 9–10 short spines in 1–2 rows. Metatibia (tb3) with 14–16 short spines in 2–3 rows (Fig. 13B); first tarsomere digitally expanded, bearing a pair of long setae; second tarsomere slightly enlarged (Fig. 13B).

Metasoma. Metasoma weakly sclerotized, each tergite with a row of setae.

 Host.

The type series was also reared from syconia of Ficus tinctoria subsp. tinctoria G. Forst., pollinated by an undescribed species of Kradibia (sp026 in our database).

 Distribution.

Australasian region, Papua.

 Etymology.

The specific epithet arfak is a noun in apposition referring to the Arfak people of West Papua, Indonesia.

Key to females of the <italic>ravii</italic> species group
1 Ovipositor sheath distinctly longer than metasoma, more than 1.15× metasomal length (Figs 9A, 11A, 13A); ovipositor sheath more than 5× metatibial length; Mt7 at least 1.8× as long as Mt8+9 2
– Ovipositor sheath shorter than metasoma (Figs 5A, 7A); ovipositor sheath less than 4× metatibial length; Mt7 less than 1.8× as long as Mt8+9 4
2(1) Ovipositor sheath more than 5× metatibial length (Fig. 11A); Mt7 less than 4× as long as wide; length of Mt7 about 1.8× the length of Mt8+9; ventral metasoma orange; mesoscutellum finely engraved (Fig. 11A) P. paiwan sp. nov.
– Ovipositor sheath more than 6× metatibial length; Mt7 more than 4.4× as long as wide; ventral metasoma orange or dark; mesoscutellum finely engraved or smooth to weakly sculptured 3
3(2) Mt7 more than 5× as long as wide (Fig. 13A); length of Mt7 about 2.5× length of Mt8+9; ventral metasoma orange; mesoscutellum finely engraved P. arfak sp. nov.
– Mt7 less than 4.7× as long as wide (Fig. 9A); length of Mt7 about 2.2× the length of Mt8+9; ventral metasoma dark; mesoscutellum smooth to weakly sculptured P. minang sp. nov.
4(1) Mt7 less than 3.3× as long as wide; length of Mt7 about 1.45× the length of Mt8+9 (Fig. 5A); ovipositor sheath about 3.8× metatibial length and about 0.9× metasomal length P. ravii (Priyadarsanan, 2000)
– Mt7 more than 3.5× as long as wide; length of Mt7 about 1.65× the length of Mt8+9 (Fig. 7A); ovipositor sheath about 3.4× metatibial length and about 0.8× metasomal length P. bannaensis sp. nov .
Tentative key to males of the <italic>ravii</italic> species group

Males are particularly challenging to separate, and accurate identification often relies on morphometric measurements. As some species are currently represented by few specimens, the key should be considered provisional and may require modification when additional material becomes available.

1 Pronotum long, at least 1.8× as long as mesonotum 2
– Pronotum shorter, less than 1.6× as long as mesonotum 3
2(1) Head slightly longer than wide, head L/W about 1.06; temple long, about 4.1× eye height; mesonotum about 1.4× as long as propodeum P. minang sp. nov.
– Head wider than long, head L/W about 0.93; temple shorter, about 3.4× eye height; mesonotum about 2.0× as long as propodeum P. paiwan sp. nov.
3(1) Mesonotum short relative to propodeum, only 1.2–1.3× as long as propodeum; pronotum 1.4–1.5× as long as mesonotum P. bannaensis sp. nov.
– Mesonotum longer relative to propodeum, 1.6–1.9× as long as propodeum; pronotum 1.0–1.5× as long as mesonotum 4
4(3) Malar space short, 0.37–0.51× eye height; temple 2.6–3.3× eye height P. ravii
– Malar space longer, 0.54–0.84× eye height; temple 3.3–4.0× eye height P. arfak sp. nov.
Discussion

Although originally described as a distinct genus, Philoverdance is not recovered as the sister lineage of all other Philotrypesis in the BUSCO-based phylogeny. Instead, it is nested within Philotrypesis. The erection of Philoverdance was justified by a combination of unusual morphological traits, including the low insertion of the antennae, falcate mandibles, shortened terminal metasomal tergites, and the highly modified morphology of brachypterous males (Priyadarsanan 2000). However, none of these characters appears to represent an isolated evolutionary lineage when considered in a broader phylogenetic context (Segar et al. 2012). Furthermore, the host association of these species provides no evidence for an ancient or independent origin. Species of the former Philoverdance are associated with Ficus tinctoria, a widespread member of subgenus Terega, one of the most diverse and broadly distributed Old World fig lineages (Pederneiras et al. 2015). The host is neither phylogenetically isolated nor associated with a peculiar ecological syndrome, and its distribution extends continuously from South Asia to the Pacific Islands. Likewise, the geographical distribution of the species group spans a large portion of the Oriental and Australasian regions and does not correspond to a relictual or geographically isolated lineage. Rather than supporting generic status, the available evidence suggests that the former Philoverdance represents a morphologically distinctive but deeply nested lineage within Philotrypesis. Its members share a unique combination of characters that readily distinguishes them from other species groups of the genus, but these differences are best interpreted as derived specializations at the species-group level. We therefore treat Philoverdance as the P. ravii species group and provide a diagnosis to facilitate its recognition within Philotrypesis.

Diagnosis of the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Philotrypesis">Philotrypesis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ravii">ravii</tp:taxon-name-part></tp:taxon-name></italic> species group.

Females are recognized by the antennae inserted low on the face, at or slightly above the ocular line and close to the epistomal margin, the strongly convergent cheeks, and the falcate, apparently unidentate mandibles. Head and mesosoma black with metallic green to bluish-green reflections; legs entirely yellowish; fore wing hyaline with yellowish venation. Lower face and parascrobal areas smooth to distinctly alutaceous, but never strongly sculptured; mesosoma predominantly smooth or weakly engraved, with the mesoscutum transversely alutaceous and the mesoscutellum smooth to weakly sculptured. Pronotum transverse and distinctly shorter than the mesoscutum. Propodeum with a median depression bounded by submedian carinae and bearing a posterior carina associated with two to five anterior carinae. Metasoma characterized by a relatively short gastral tail, with Mt7 and Mt8+9 together distinctly shorter than the remainder of the metasoma, Mt8+9 not strongly elongate, and ovipositor sheaths relatively short to moderately long (approximately 0.8–1.5× metasomal length), markedly shorter than in most other Philotrypesis species groups. The group comprises two morphological sublineages: a western lineage (P. ravii, P. minang sp. nov., P. bannaensis sp. nov.) characterized by a dark ventral metasoma, weak facial sculpture and a smooth mesoscutellum, and an eastern lineage (P. paiwan sp. nov., P. arfak sp. nov.) characterized by a ventrally orange metasoma, more conspicuous facial sculpture and a weakly sculptured mesoscutellum with a median longitudinal declivity. Males brachypterous, with vestigial fore wings, enlarged temples, strongly striate scrobal depression, and a pronotum as long as to markedly longer than the fused meso- and metanotum. Antennal scrobes roofed by an overgrowth of the cuticle; antennae inserted beneath this projection by a long radicle. Antenna with four or five funicular segments and a simple to segmented clava. Clypeal margin strongly emarginate, rounded to triangularly so. Mesosoma compact and convex, with free thoracic terga and a distinct hump at the pronoto-metanotal angle. Metasoma weakly sclerotized and dorsoventrally flattened.

 Barcoding of the species within the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Philotrypesis">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ravii">ravii</tp:taxon-name-part></tp:taxon-name></italic> sp.gp.

DNA barcoding has proven particularly effective for species discovery and delimitation in parasitoid Hymenoptera, where closely related species often differ by only a few percent in COI sequence divergence, whereas deeper divergences generally correspond to well-established species boundaries (Stahlhut et al. 2013). Nevertheless, no universal threshold exists, and species hypotheses are best evaluated using an integrative framework combining molecular, morphological and ecological evidence (Ratnasingham and Hebert 2007). In Pteromalidae, DNA barcoding has repeatedly uncovered substantial cryptic diversity, including previously unrecognized species complexes and host-associated lineages (Cruaud et al. 2013; Pollmann et al. 2023). In many cases, closely related species exhibit only subtle morphological differences despite clear genetic differentiation (Kin et al. 2026). This situation contrasts markedly with that observed in the Philotrypesis ravii species group. Although the species recognized here are genetically close, they can be readily distinguished by a consistent suite of morphological characters involving ovipositor length, metasomal proportions, sculpture, coloration and male morphology.

The BUSCO-based phylogeny, although based on a more limited sampling, provides an independent line of evidence supporting a deep split within the species group. The genomic dataset includes only P. minang sp. nov., P. bannaensis sp. nov. and P. paiwan sp. nov., but nonetheless identifies P. paiwan sp. nov. as distinct from the two western taxa. While the absence of both P. ravii and P. arfak sp. nov. prevents a direct comparison with the complete COI topology, the genomic results are fully consistent with the existence of a major phylogenetic break separating western and eastern lineages. Importantly, this signal is recovered independently from morphology, mitochondrial data and host biogeography. Therefore, despite the relatively shallow COI divergences observed among some taxa, the concordance between mitochondrial and genomic data, morphology, geography and host associations provides strong evidence that all entities recognized within the P. ravii species group represent independently evolving lineages deserving species status, and must be described as such.

 Morphological lability in host associated lineages

The phylogenetic, morphological and biogeographical evidence collectively supports the existence of a distinct P. ravii sp. gp., which comprises two sublineages associated correspondingly with two geographically structured subspecies of F. tinctoria, suggesting that diversification within the ravii sp. gp. may have been influenced by the historical fragmentation and expansion of host populations across the Indo-Pacific region.

An interesting pattern emerging from both morphology and geography is the repeated occurrence of short- and long-ovipositor species within the major host-associated lineages. In the western lineage associated with Ficus tinctoria subsp. gibbosa, the southern Chinese species P. bannaensis sp. nov. and the Indian P. ravii possess relatively short ovipositors, whereas the Sumatran P. minang sp. nov. exhibits a much longer ovipositor. A similar trend is observed in the eastern lineage associated with subsp. tinctoria, where the Taiwanese P. paiwan sp. nov. has a shorter ovipositor than its Papuan sister species P. arfak sp. nov. Although direct measurements of syconium size and wall thickness across geographical population of F. tinctoria are currently lacking, this pattern is suggestive of repeated ecological adaptation to local host traits, as has been documented in other Ficus lineages (Yu et al. 2018).

Our results points to a scenario in which diversification within the P. ravii sp. gp. has been shaped by both geographic isolation and the evolutionary history of the F. tinctoria complex, producing a series of closely related species distributed across the Indo-Pacific and exhibiting parallel trends in ovipositor length, gastral morphology and external sculpture.

 Acknowledgments

We thank Ma Li, Deng Yao, and Li Cheng-Fang at Yunnan Agriculture University for assistance with using Keyence imaging system.

We are grateful to our friend Rosichon Ubaidillah (LIPI, Bogor, Indonesia) for sharing our field trip in northern Papua, to Magali Proffit and Finn Kjellberg (CEFE, CNRS, France) for sampling figs and to Anthony Bain, Lien Siang Chou and Hsy-Yu Tzeng for facilitating fieldwork and access to collections in Taiwan.

 References Abdurahiman UC (1980) Observations on the oviposition behaviour in Philotrypesis pilosa Mayr (Torymidae: Hymenoptera). Proceedings of the All-India Symposium on Environmental Biology: 146–150. Abdurahiman UC, Joseph KJ (1975) New fig insects [Hym.: Chalcidoidea] from India: Three torymids parasitic on agaonids. Entomophaga: 73–80. https://doi.org/10.1007/bf02373452 Balakrishnan Nair P, Abdurahiman UC, Joseph M (1981) Two new torymidae (Hymenoptera: Chalcidoidea) from Ficus hirta. Oriental Insects 15: 433–442. https://doi.org/10.1080/00305316.1981.10434341 Berg CC, Corner EJH (2005) Flora Malesiana: Moraceae (Ficus). National Herbarium Nederland, Universiteit Leiden branch, Leiden, The Netherlands, 293–296. Bernt M, Donath A, Jühling F, Externbrink F, Florentz C, Fritzsch G, Pütz J, Middendorf M, Stadler PF (2013) MITOS: Improved de novo metazoan mitochondrial genome annotation. Molecular Phylogenetics and Evolution 69: 313–319. https://doi.org/10.1016/j.ympev.2012.08.023 Borowiec ML (2016) AMAS: A fast tool for alignment manipulation and computing of summary statistics. PeerJ 4: e1660. https://doi.org/10.7717/peerj.1660 Bouček Z (1988) Australian Chalcidoidea (Hymenoptera): A biosystematic revision of genera and fourteen families, with a reclassification of species. CAB International Institute of Entomology, Wallingford, Oxon. Bouček Z, Rasplus J-Y (1991) Illustrated key to West-Palearctic Genera of Pteromalidae (Hymenoptera – Chalcidoidea). INRA Editions, série Techniques et Pratiques, 140 pp. Bouček Z, Watsham A, Wiebes JT (1981) The fig wasp fauna of the receptacles of Ficus thonningii (Hymenoptera, Chalcidoidea). Tijdschrift voor Entomologie 124: 149–233. Burks R, Mitroiu M-D, Fusu L, Heraty JM, Janšta P, Steve H (2022) From hell’s heart I stab at thee! A determined approach towards a monophyletic Pteromalidae and reclassification of Chalcidoidea (Hymenoptera). Journal of Hymenoptera Research 94: 13–88. https://doi.org/10.3897/jhr.94.94263 Burks RA, Gibson GAP, Heraty JM (2025) External morphology of adult Chalcidoidea. In: Heraty JM, Woolley JB (Eds) Chalcidoidea of the World. CABI Press, Oxford, UK, 69–110. https://doi.org/10.1079/9781800623545.0005 Burks RA, Mitroiu MD, Cruaud A, Rasplus J-Y (2025) Pteromalidae. In: Heraty JM, Woolley JB (Eds) Chalcidoidea of the World. CABI Press, Oxford, UK, 568–595. Capella-Gutiérrez S, Silla-Martínez JM, Gabaldón T (2009) trimAl: A tool for automated alignment trimming in large-scale phylogenetic analyses. Bioinformatics 25: 1972–1973. https://doi.org/10.1093/bioinformatics/btp348 Chen SF, Zhou YQ, Chen YR, Gu J (2018) fastp: An ultra-fast all-in-one FASTQ preprocessor. Bioinformatics 34: i884–i890. https://doi.org/10.1093/bioinformatics/bty560 Cook JM, Bean D (2006) Cryptic male dimorphism and fighting in a fig wasp. Animal Behaviour 71: 1095–1101. https://doi.org/10.1016/j.anbehav.2005.07.027 Cruaud A, Underhill JG, Huguin M, Genson G, Jabbour-Zahab R, Tolley KA (2013) A multilocus phylogeny of the world Sycoecinae fig wasps (Chalcidoidea: Pteromalidae). PLOS ONE 8(11): e79291. https://doi.org/10.1371/journal.pone.0079291 Cruaud A, Nidelet S, Arnal P, Weber A, Fusu L, Gumovsky A, Huber J, Polaszek A, Rasplus J-Y (2019) Optimized DNA extraction and library preparation for minute arthropods: Application to target enrichment in chalcid wasps used for biocontrol. Molecular Ecology Resources 19: 702–710. https://doi.org/10.1111/1755-0998.13006 Cruaud A, Rasplus J-Y, Zhang JX, Burks R, Delvare G, Fusu L (2024) The Chalcidoidea bush of life: Evolutionary history of a massive radiation of minute wasps. Cladistics 40: 34–63. https://doi.org/10.1111/cla.12561 Deng XX, Chen LF, Tian EW, Zhang DY, Wattana T, Yu H, Kjellberg F, Segar ST (2021) Low host specificity and broad geographical ranges in a community of parasitic non- pollinating fig wasps (Sycoryctinae; Chalcidoidea). Journal of Animal Ecology 90: 1678–1690. https://doi.org/10.1111/1365-2656.13483 Förster A (1878) Kleine Monographien parasitischer Hymenopteren. Verhandlungen des Naturhistorischen Vereines de Preussischen Rheinlande und Westfalens 35: 42–82. Gibson GAP (1997) Morphology and Terminology. In: Gibson GAP, Huber JT, Woolley JB (Eds) Annotated Keys to the Genera of Nearctic Chalcidoidea (Hymenoptera). National Research Council Research Press, Ottawa, Ontario, Canada, 16–44. Grandi G (1921) Richerche sul gen. Philotrypesis (Hymenoptera-Chalcididae). Bollettino del Laboratorio di Zoologia Portici 15: 33–190. Grandi G (1930) Monographia del gen. Philotrypesis Först. Bollettino del Laboratorio di Entomologia Bologna 1: 107–235. Hebert PDN, Floyd R, Jafarpour S, Prosser SWJ (2025) Barcode 100K specimens: in a single nanopore run. Molecular Ecology Resources 25. https://doi.org/10.1111/1755-0998.14028 Heraty JM, Hawks DC (1998) Hexamethyldisilazane – A Chemical Alternative For Drying Insects. Entomological News 109: 369–374. https://doi.org/10.5281/zenodo.15983478 Jiang ZF, Huang DW, Chen LL, Zhen WQ, Fu YG, Peng YQ (2006) Rampant host switching and multiple female body colour transitions in Philotrypesis (Hymenoptera: Chalcidoidea: Agaonidae). European Society for Evolutionary Biology 19: 1157–1166. https://doi.org/10.1111/j.1420-9101.2006.01087.x Jin JJ, Yu WB, Yang JB, Song Y, dePamphilis CW, Yi TS, Li DZ (2020) GetOrganelle: A fast and versatile toolkit for accurate de novo assembly of organelle genomes. Genome Biology 21: 1–31. https://doi.org/10.1186/s13059-020-02154-5 Joseph KJ (1955) Observations sur la biologie de Philotrypesis caricae (Hym. Chalcidiens Callimomidae). Comptes Rendus Hebdomadaires des Séances de l’Académie des Sciences 241: 1624–1625. Jousselin E, Noort Sv, Greeff JM (2004) Labile male morphology and intraspecific male polymorphism in the Philotrypesis fig wasps. Molecular Phylogenetics and Evolution 33: 706–718. https://doi.org/10.1016/j.ympev.2004.08.008 Jousselin E, van Noort S, Berry V, Rasplus J-Y, Rønsted N, Erasmus JC, Greeff JM (2008) One fig to bind them all: host conservatism in a fig wasp community unraveled by cospeciation analyses among pollinating and nonpollinating fig wasps. Evolution 62: 1777–1797. https://doi.org/10.1111/j.1558-5646.2008.00406.x Kalyaanamoorthy S, Minh BQ, Wong TKF, Haeseler Av, Jermiin LS (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14: 587–589. https://doi.org/10.1038/nmeth.4285 Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution 30: 772–780. https://doi.org/10.1093/molbev/mst010 Kerdelhué C, Rossi J-P, Rasplus J-Y (2000) Comparative community ecology studies on old world figs and fig wasps. Ecology 81: 2832–2849. https://doi.org/10.2307/177345 Kin OC, Dario NN, Shahrul A, Rafe MB, Carl RH, Indraneil D, Stefan TH (2026) A genomic perspective on cryptic species reveals complex evolutionary dynamics in the gray zone of the speciation continuum. Systematic Biology: syag001. https://doi.org/10.1093/sysbio/syag001 Li D, Liu CM, Luo R, Sadakane K, Lam TW (2015) MEGAHIT: An ultra-fast single-node solution for large and complex metagenomics assembly via succinct de Bruijn graph. Bioinformatics 31: 1674–1676. https://doi.org/10.1093/bioinformatics/btv033 McLeish MJ, van Noort S, Tolley KA (2010) African parasitoid fig wasp diversification is a function of Ficus species ranges. Molecular Phylogenetics and Evolution 57: 122–134. https://doi.org/10.1016/j.ympev.2010.05.025 Moore JC, Obbard DJ, Reuter C, West SA, Cook JM (2009) Male morphology and dishonest signalling in a fig wasp. Animal Behaviour 78: 147–153. https://doi.org/10.1016/j.anbehav.2009.04.006 Murray MG (1989) Environmental constraints on fighting in flightless male fig wasps. Animal Behaviour 38: 186–193. https://doi.org/10.1016/S0003-3472(89)80081-8 Murray MG (1990) Comparative morphology and mate competition of flightless male fig wasps. Animal Behaviour 39: 434–443. https://doi.org/10.1016/S0003-3472(05)80406-3 Noyes JS (1982) Collecting and preserving chalcid wasps (Hymenoptera: Chalcidoidea). Journal of Natural History 16: 315–334. https://doi.org/10.1080/00222938200770261 Pederneiras LC, Carauta JPP, Neto SR, Mansano VdF (2015) An overview of the infrageneric nomenclature of Ficus (Moraceae). Taxon 64: 589–594. https://doi.org/10.12705/643.12 Pollmann M, Kuhn D, König C, Homolka I, Paschke S, Reinisch R, Schmidt A, Schwabe N, Weber J, Gottlieb Y, Steidle JLM (2023) New species based on the biological species concept within the complex of Lariophagus distinguendus (Hymenoptera, Chalcidoidea, Pteromalidae), a parasitoid of household pests. Ecology and Evolution 13: 1–22. https://doi.org/10.1002/ece3.10524 Priyadarsanan DR (2000) Fig. insects of Kerala. Records of the Zoological Survey of India, Occasional Paper 182: 1–175. Ratnasingham S, Hebert PDN (2007) BOLD: The Barcode of Life Data System (www.barcodinglife.org). Molecular Ecology Notes 7: 355–364. https://doi.org/10.1111/j.1471-8286.2007.01678.x Rognes T, Flouri T, Nichols B, Quince C, Mahé F (2016) VSEARCH: A versatile open source tool for metagenomics. PeerJ 4. https://doi.org/10.7717/peerj.2584 Schindelin J, Arganda-Carreras I, Frise E (2012) Fiji: An open-source platform for biological-image analysis. Nature Methods 9: 676–682. https://doi.org/10.1038/nmeth.2019 Segar ST, Cook JM (2012) The dominant exploiters of the fig/pollinator mutualism vary across continents, but their costs fall consistently on the male reproductive function of figs. Ecological Entomology 37: 342–349. https://doi.org/10.1111/j.1365-2311.2012.01370.x Segar ST, Lopez-Vaamonde C, Rasplus J-Y, Cook JM (2012) The global phylogeny of the subfamily Sycoryctinae (Pteromalidae): parasites of an obligate mutualism. Molecular Phylogenetics and Evolution 65: 116–125. https://doi.org/10.1016/j.ympev.2012.05.030 Stahlhut JK, Fernandez-Triana J, Adamowicz SJ, Buck M, Goulet H, Hebert PD (2013) DNA barcoding reveals diversity of Hymenoptera and the dominance of parasitoids in a sub-arctic environment. BMC Ecology 13. https://doi.org/10.1186/1472-6785-13-2 Tegenfeldt F, Kuznetsov D, Manni M, Berkeley M, Zdobnov EM, Kriventseva EV (2025) OrthoDB and BUSCO update: Annotation of orthologs with wider sampling of genomes. Nucleic Acids Research 53: D516–D522. https://doi.org/10.1093/nar/gkae987 UCD Community (2023) UCD Community: Universal Chalcidoidea Database (UCD) curated in TaxonWorks. https://sfg.taxonworks.org/api/v1/ Wang R, Matthews AVG, Ratcliffe JV, Barwell LJ, Peng YQ, Chou LS, Yu H, Yang HW, Compton SG (2014) First record of an apparently rare fig wasp feeding strategy: obligate seed predation. Ecological Entomology 39: 492–500. https://doi.org/10.1111/een.12122 Wiebes JT (1966a) Bornean fig wasps from Ficus stupenda Miquel (Hymenoptera, Chalcidoidea). Tijdschrift voor Entomologie 109: 163–192. https://doi.org/10.5281/zenodo.16485072 Wiebes JT (1966b) The structure of the ovipositing organs as a tribal character in the Indo-Australian SycophaginaeTorymidae (Hymenoptera, Chalcidoidea). Zoologische Mededelingen 41: 151–159. Wiebes JT (1967) Provisional host catalogue of fig wasps (Hymenoptera, Chalcidoidea). Zoologische Verhandelingen 83: 1–44. https://doi.org/10.1163/9789004742772 Wu T, Dunn DW, Hu H-Y, Niu LM, Xiao JH, Pan XL, Feng G, Fu YG, Huang DW (2013) The occurrence of fig wasps in the fruits of female gynodioecious fig trees. Acta Oecologica 46: 33–38. https://doi.org/10.1016/j.actao.2012.10.009 Yu H, Liang D, Tian EW, Zheng LN, Kjellberg F (2018) Plant geographic phenotypic variation drives diversification in its associated community of a phytophagous insect and its parasitoids. BMC Evolutionary Biology 18. https://doi.org/10.1186/s12862-018-1239-5 Zhen WQ, Huang DW, Yang DR, Xiao H, Zhu CD (2004) Oviposition behavior of Philotrypesis pilosa Mayr (Hymenoptera: Agaonidae). Acta Entomologica Sinica 47: 365–371. Additional information Conflict of interest

The authors have declared that no competing interests exist.

 Ethical statement

No ethical statement was reported.

 Artificial Intelligence (AI) use

The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.

No AI tools were used in the preparation of this manuscript.

 Funding

This study was funded by the program Yunnan Key Research and Development Program (202503AP140027) and the program National Natural Science Foundation of China (32261123001).

 Author contributions

Jean-Yves Rasplus conceived the study, and both he and Peng Yan-Qiong supervised the work. Peng Yan-Qiong acquired the funding. Sabine Nidelet and Yu-Ran Li extracted DNA, and Sabine Nidelet conducted the PCR and COI sequencing. Astrid Cruaud and Yu-Ran Li performed the data analyses. Yu-Ran Li and Jean-Yves Rasplus prepared the specimens and took morphological measurements. Jean-Yves Rasplus and Yu-Ran Li wrote the manuscript, and all authors revised and approved the final version.

 Author ORCIDs

Y.-R. Li https://orcid.org/0000-0003-2348-8224

A. Cruaud https://orcid.org/0000-0001-8932-4199

S. Nidelet https://orcid.org/0000-0003-0437-319X

Y.-Q. Peng https://orcid.org/0000-0002-7453-9119

J.-Y. Rasplus https://orcid.org/0000-0001-8614-6665

 Data availability

The COI sequences used for phylogenetic analysis have been deposited in the NCBI GenBank database under accession numbers PZ789829 and PZ794766–PZ794774.