Research Article |
Corresponding author: Jeong Jae Yoo ( jjbiodiversity@gmail.com ) Academic editor: Ankita Gupta
© 2024 Jeong Jae Yoo, D. Christopher Darling.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Yoo JJ, Darling DC (2024) Integrative taxonomic revision of the Nearctic Perilampus hyalinus species complex (Hymenoptera, Chalcidoidea, Perilampidae) resolves 100 years of confusion about the host associations of P. hyalinus Say. Journal of Hymenoptera Research 97: 1301-1383. https://doi.org/10.3897/jhr.97.133255
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The enigmatic Nearctic parasitoid Perilampus hyalinus Say (Hymenoptera: Chalcidoidea: Perilampidae) has long been suspected as a species complex with a wide range of host associations and differing modes of parasitism. In this study we clarify the status of this species by combining morphological evidence, two genes (COI and ITS2) and host information and recognize ten species in the P. hyalinus species complex in the Nearctic region. Eight new species are described: Perilampus arcus Yoo & Darling, sp. nov., P. crassus Yoo & Darling, sp. nov., P. neodiprioni Yoo & Darling, sp. nov., P. monocteni Yoo & Darling, sp. nov., P. pilosus Yoo & Darling, sp. nov., P. seneca Yoo & Darling, sp. nov., P. sonora Yoo & Darling, sp. nov., and P. ute Yoo & Darling, sp. nov. A reared specimen with a COI sequence is designated as the Neotype of P. hyalinus Say establishing this species as either a hyperparasitoid that parasitizes dipteran parasitoids of Orthoptera or a parasitoid of dipteran kleptoparasites of Crabronidae and Sphecidae that provision their nests with Orthoptera. Perilampus sirsiris (Argaman) and four of the new species are hyperparastioids, parasitizing hymenopteran and dipteran parasitoids of Lepidoptera. Perilampus neodiprioni and P. monocteni can develop as both primary parasitoids of Diprionidae (Hymenoptera) and as hyperparasitoids, parasitizing dipteran and hymenopteran parasitoids of diprionid sawflies. Perilampus neodiprioni is strictly associated with Neodiprion sawflies, and P. monocteni is associated with Monoctenus sawflies.
COI, ITS2, key, morphology, new species, parasitoid, species delimitation, systematics
The Perilampus hyalinus species group contains the most conspicuous species of Perilampus in the Western Hemisphere. Specimens are large (3–5 mm) and brightly iridescent in color and are frequently collected on flowers. It is one of the 6 species groups recognized by
Perilampus hyalinus Say has been shrouded in confusion for at least 100 years, since biological studies of these parasitoids indicated divergent host associations and modes of parasitism (
Resolving the status of P. hyalinus could have important implications for the biological control of insect pests important to agriculture and forestry. Careful rearing studies have revealed P. hyalinus (s.l) as primary parasitoids attacking Neodiprion sawflies (Hymenoptera: Diprionidae), which are serious forest pests particularly in boreal forests (
To clarify the status of the P. hyalinus species group,
Maximum likelihood tree of Perilampus hyalinus species group retrieved from the combined analysis of COI and ITS2 (modified from
In this paper we revise the P. hyalinus species complex and recognize 10 Nearctic species using morphology, molecular data (COI and ITS2), and host information. We also designate a Neotype for Perilampus hyalinus Say. Most of Thomas Say’s type material is regarded as lost and the surviving Say specimens are in the Museum of Comparative Zoology (
Specimens were examined with a Leica MZ7.5 stereo zoom microscope and were illuminated with a Leica 20-watt halogen light source, filtered through a strip of translucent Mylar® film. Images of specimens were taken with a Keyence digital microscope VHX-7000 series, and edited with Adobe® Illustrator CC ver. 21.0.2 (Adobe Systems Inc., California, USA). The material examined and descriptions for each species were generated by a series of R coding commands and an Excel spreadsheet; the R-codes are freely available on GitHub (https://github.com/esdarling/r-taxonomy). The Excel spreadsheet for the species descriptions was output from Lucid Builder 3.3 (University of Queensland, Australia). Morphological terms mostly follow
The morphological characters of Perilampus hyalinus species group (A B D H F) and other Perilampus species groups (C E G I) A head, anterior-oblique view, P. seneca B vertex, dorsal view, P. neodiprioni C vertex, dorsal view, P. auratus D malar space, P. seneca E malar space, P. platigaster F axilla and axillula, P. hyalinus G axilla and axillula, Perilampus sp. H pronotum and mesoscutum, posterior oblique view, P. neodiprioni I pronotum and mesoscutum, posterior oblique view, P. platigaster. Abbreviations: ax, axilla; axl, axillula; axsh, axillular shelf; ca, carina; fc, frontal carina; ms, malar space; msl, malar sulcus; not, notaulus; psa, parascrobal area. [A ROME182769; B, H ROME162273; C ROME162320; D ROME199563; E ROME141508; F ROME162229; G ROME182831; I DHJPAR0038540]. Scale bars: 250 μm (A–C, H, I); 100 μm (D–G).
Sexual dimorphism in the Perilampus hyalinus species group A female head, anterior view, P. seneca B male head, anterior view, P. seneca C female scape anterior margin, P. seneca D female scape inner margin, P. seneca E female scape inner margin, P. neodiprioni F male scape anterior margin, P. neodiprioni G female head, dorsal view, P. seneca H male head, dorsal view, P. seneca I female flagellum, P. seneca J male flagellum, P. seneca K female habitus, dorsal view, P. neodiprioni L male habitus, dorsal view, P. neodiprioni. Abbreviation: scp, scape. [A ROME199563; B ROME183976; C, D ROME199531; E ROME198219; F ROME198147]. Scale bars: 250 μm (A, B, G–J); 100 μm (C–F); 500 μm (K, L).
All measurements were taken with the Keyence Communication software, either directly from the specimens or from digital images, in which case care was taken to avoid problems associated with parallax. The measurements of each species were taken from four to five specimens per sex. Holotype and paratypes were measured for each new species. The primary type for the previously described Nearctic P. hyalinus complex species could not be borrowed for measurement (e.g. P. sirsiris). For this species, the measurements were taken from conspecific specimens that were sequenced and/or with collection localities close to that of the types. Most of the following abbreviations for measurements follow
Specimens were examined from the following collections: American Museum of Natural History, New York City, NY, United States (
Hind legs and/or middle legs were removed from the specimens of each morphospecies for DNA extraction. The molecular work was done either Canadian Centre for DNA Barcoding (CCDB) at the University of Guelph or in the Laboratory of Molecular Systematics (LMS) at Royal Ontario Museum. At LMS, DNA was extracted using a Qiagen DNeasy Blood and Tissue Kit following the manufacturer’s instructions and DNA was eluted with 50 μl of AE buffer. Each 25 μl PCR reaction consisted of 1 μl of template DNA, 18.89 μl ddH2O, 0.56 μl dNTPs [10 mM], 0.05 μl Platinum Taq (Invitrogen), 1 μl [0.01 mM] each of the universal COI primers LCO1490 and HCO2198 (
The Perilampus platigaster species group is regarded as the sister group to the P. hyalinus species group based on two putative morphological synapomorphies, frontal carina and finger-like axillula (
The pairwise uncorrected p-distances within and between preliminary species inferred from morphology were calculated in PAUP* 4.0a169 (
Three molecular species delimitation methods were employed for each locus to estimate the number of molecular operational taxonomic units (MOTUs): Automatic Barcoding Gap Discovery (ABGD) (
The species distribution maps were made with the online GIS tool SimpleMappr (
The host association of individual specimens was obtained from the associated labels, available host remains, and/or publications. Where necessary, the host names were updated to reflect the current nomenclature. In many instances, the dipteran or ichneumonid primary parasitoid hosts pupated inside the cocoons or pupae of their herbivore hosts. This necessitated a careful examination of the inner contents of the cocoons and pupae to determine whether the Perilampus had developed as primary or hyperparasitoids. The sawfly cocoons and lepidopteran pupae from which Perilampus exited were either dissected (Fig.
The molecular analysis provides strong support for both the P. hyalinus species group and the P. hyalinus species complex—both the ML and BI analyses yielded similar tree topologies (Fig.
The BINs assigned for each specimen by BOLD are shown in the material examined. The current BINs (December 28, 2023) and the number of BINed specimens are indicated in the remarks section for each species. The pairwise uncorrected distances for each gene for the Nearctic species are summarized in Table
Pairwise uncorrected p-distances for two genes among Nearctic members of the Perilampus hyalinus species complex. Intraspecific distances are along the diagonal, ITS2/COI. Interspecific distances for COI are below diagonal, for ITS2 are above diagonal.
P. arcus | P. crassus | P. neodiprioni | P. hyalinus | P. pilosus | P. seneca | P. sirsiris | P. sonora | P. ute | |
---|---|---|---|---|---|---|---|---|---|
P. arcus | 0/0–1.2 | 4.5 | 4.1 | 3.4–4.5 | 4.1–4.2 | 0.7 | 4.5–4.8 | 3.1 | 2.1–2.8 |
P. crassus | 4.0–5.5 | 0/0.2–0.5 | 2.8 | 1.0 | 2.4 | 3.8 | 2.4–2.8 | 3.4–3.5 | 4.5–5.2 |
P. neodiprioni | 4.0–5.3 | 2.7–3.7 | 0/0–0.9 | 1.7 | 2.4 | 3.4 | 1.7–2.1 | 2.8 | 4.5–4.9 |
P. hyalinus | 4.2–6.2 | 1.6–4.0 | 3.0–4.7 | 0/0–2.3 | 1.4 | 2.8 | 1.4–1.7 | 2.4 | 3.4–4.2 |
P. pilosus | 3.5–4.4 | 2.3–3.6 | 2.3–3.4 | 1.2–2.9 | 0/0–1.0 | 3.4 | 2.1–2.4 | 2.4 | 4.1–4.9 |
P. seneca | 2.5–4.0 | 4.0–5.3 | 3.4–4.7 | 4.3–6.7 | 3.8–5.2 | 0/0.2–0.7 | 3.8–4.1 | 2.4 | 1.4–2.1 |
P. sirsiris | 3.3–4.5 | 2.2–4.1 | 1.3–2.6 | 2.1–3.8 | 1.3–2.2 | 3.2–5.0 | 0–0.3/0–1.3 | 3.1–3.5 | 4.5–5.6 |
P. sonora | 3.9–5.8 | 4.2–6.3 | 4.0–5.2 | 4.6–6.5 | 3.7–4.8 | 3.4–5.4 | 3.5–4.7 | 0–0.3/1.0–1.3 | 3.8–4.5 |
P. ute | 3.1–4.4 | 3.7–5.5 | 3.2–4.2 | 4.1–6.5 | 4.1–6.5 | 2.2–3.0 | 2.9–4.7 | 4.0–5.2 | 0–0.7/0.3–2.0 |
Figs
Family Perilampidae Főrster, 1856
Taltonos Argaman, 1990 (subjective synonym, Darling, 1996).
Female. Color: head and body entirely or at least partially brightly iridescent (Fig.
Head: in anterior view weakly transverse, slightly wider than high, HW/HH 1.2–1.3; slightly wider than pronotum, HW/PW 1.1–1.2. Frontal carina: distinct, extended from posterior margin of median ocellus (Fig.
Mesosoma: slightly longer than wide, ML/MW 1.2–1.3. Pronotum: carinulate (Fig.
Metasoma: petiole short and straplike, with weak transverse wrinkles; petiolar flange short with ventral margin of upper area with shallow emargination mesad; antecostal sulcus transverse, with weak vertical carinae laterad and smoothened mesad; Mt2 with trapezoidal demarcation and shallow median groove, imbricate and wrinkled anteriad, and smooth posterad without lateral protruberances along midline, posterior margin straight and sparsely setose; Gt3 smooth.
Male. As in female, except: Color: mesonotum sometimes nearly entirely black or with cupreous iridescence (Fig.
The P. hyalinus species group is characterized by: brightly iridescent coloration (Fig.
The P. hyalinus species group occurs exclusively in the Western Hemisphere, from the southern Canada to Argentina.
The previously published host records indicate that species are mostly hyperparasitoids, parasitizing dipteran (Tachinidae and Sarcophagidae) and hymenopteran parasitoids (Ichneumonoidea) of Lepidoptera, Orthoptera, and rarely, Phasmatoidea and Coleoptera (e.g.
This species group was hypothesized as a monophyletic group based on morphological characters (
Perilampus hyalinus Say, 1829:79. (original description, sex not indicated). Type locality: USA, Pennsylvania. Type material: Type lost. Neotype. “USA:OH: Montgomery Co. New Carlisle 39.989583, -84.029056, Ex. Ceracia dentate, Ex. Melanoplus femurrubrum prob. 26.x.2014 M. D. Sheaffer”. The neotype is point-mounted (Male ROME204130, USNM). BOLD:AEA0382. ROM Online Collection.
Perilampus entellus Walker, 1843:103 (original description). Type locality: Ohio, USA. Type Material. Lectotype, B.M. Type Hym. 5.2285, NHMUK014583126 (Images examined).
Perilampus aciculatus Provancher, 1889:199 (original description). Type locality: Ottawa, Canada. Type material. Lectotype, 1359, Université Laval, Québec City, Canada (Images examined). Note: Year of publication incorrect as 1887 in subsequent references to P. aciculatus (see Barron 1975:391).
Perilampus aciculatus, Lectotype,
Perilampus aciculatus Smuylan, 1936:380 (tentative synonym of P. hyalinus Say).
Perilampus aciculatus Peck, 1963:519 (subjective synonym of P. hyalinus Say).
Perilampus aciculatus Burks, 1963:1259 (subjective synonymy “probably correct”).
Perilampus entellus, Lectotype,
Taltonos hyalinus (Say). Argaman, 1990:205 (new combination).
Taltonos aciculatus (Provancher), Argaman, 1991:5 (new combination).
Taltonos entellus (Walker), Argaman, 1991:9 (new combination).
Perilampus hyalinus Say. Darling, 1996:113 (Taltonos, subjective synonym of Perilampus).
Perilampus aciculatus Darling, 1996:113 (Taltonos, subjective synonym of Perilampus).
Perilampus entellus Darling, 1996:113 (Taltonos, subjective synonym of Perilampus).
Perilampus aciculatus, New synonymy based on Neotype designation herein.
Perilampus entellus, New synonymy based on Neotype designation herein.
Canada: 77 females, 47 males. USA: 67 females, 31 males. (Suppl. materials).
Canada: 1 female. Ontario: 1 female. Durham R.M., Glen Major Forest: (1 female: ROME152664-ROME; BOLD:AEA0382; ITS2). Mexico: 2 females. Jalisco: 1 female. (1 female: ROME200751-HNHM). Sonora: 1 female. (1 female: ROME162260-USNM).
Female (Fig.
Perilampus hyalinus Female A habitus, lateral view B habitus dorsal view C lateral lobe of mesoscutum along notaulus D lateral panel of pronotum, posterior oblique view E axillula F axilla G head, dorsal view H head, anterior view I lower face J parascutal carina K mesoscutellum apex L, M mesofemoral depression. [A ROME185913; B, K ROME189058; C, M ROME185947; D, F ROME189056; E, G ROME162229; H ROME167636; I ROME189060; L ROME162246]. Scale bar: 1 mm (A).
Head (Fig.
Mesosoma (Fig.
Male (Fig.
Perilampus hyalinus Male A habitus, lateral view B lateral panel of pronotum and parascutal carina, posterior oblique view C head, anterior view D head, dorsal view E lower face F lateral lobe of mesoscutum along notaulus G, H scape. [A Neotype, ROME204130; B–D, F, G ROME182798; E ROME167638; H ROME189059]. Scale bars: 1 mm (A); 100 μm (G, H).
Perilampus hyalinus is morphologically similar to P. neodiprioni, but the axillula is always smooth dorsad without piliferous punctures (Fig.
(Fig.
Perilampus hyalinus is a hyperparasitoid, attacking dipteran parasitoids of Orthoptera and dipteran kleptoparasites of Crabronidae and Sphecidae provisioning with Orthoptera and rarely parasitoids of dipteran parasitoids attacking Phasmida (ROME204120). Hosts: Tachinidae (Diptera). Ceracia dentata (Coquillett) from Melanoplus femurrubrum (De Geer) (Acrididae). Tachinidae from Phasmatidae. Sarcophagidae (Diptera). Sarcophaga sp. from Melanoplus sanguinipes (Fabricius). Sarcophagids from Tettigoniidae and Oecanthinae collected in the nests of Isodontia mexicana (Saussure) (Sphecidae) (
A female from Ontario (ROME152664), Canada, has a smooth vertex. COI and ITS2 suggest that this specimen is a rare morphological variant of P. hyalinus.
The identity of P. hyalinus has long been obscured by the presumed lost type specimen (
Ichneumon cyaneus Brullé, 1846:21 (Plate V, #4). Type locality: USA, “Carolina”. Type material: Holotype. “Carolina”. (Female Paris EY35408, MHNH) (images examined).
Perilampus cyaneus Dalla Torre, 1898:355 (new combination ?).
Perilampus hyalinus
Viereck, 1910:647 (subjective synonym P. cyaneus ?, cited by
Taltonos sirsiris Argaman, 1990:15. Replacement name, Perilampus cyaneus Brullé (nec Fabricius 1798).
Perilampus sirsiris Darling, 1996:113 (Taltonos, subjective synonym of Perilampus).
Perilampus eucyaneus Özdikmen, 2011. Unnecessary replacement name.
Canada: 4 females, 8 males. USA: 17 females, 11 males. (Suppl. materials).
Female (Fig.
Perilampus sirsiris Female A habitus, lateral view B habitus, dorsal view C lateral lobe of mesoscutum along notaulus D, E lateral panel of pronotum, posterior oblique view F parascutal carina G head, dorsal view H head, anterior view I lower face J head, lateral view K mesoscutellum apex L mesofemoral depression. [A, B, D, F, H, J, K ROME182764; E, G, L ROME189054; I ROME182766]. Scale bar: 1 mm (A).
Head (Fig.
Mesosoma (Fig.
Male (Fig.
Perilampus sirsiris Male A habitus, lateral view B lateral panel of pronotum, parascutal carina, and axilla, posterior oblique view C head, dorsal view D head, anterior view E lateral lobe of mesoscutum along notaulus F, G scape. [A ROME162278; B ROME204099; C ROME199527; D ROME162281; F ROME152680; E ROME162281; G ROME185910]. Scale bars: 1 mm (A); 100 μm (F, G).
Perilampus sirsiris and P. arcus are the only Nearctic species with steeply curved or angulate parascutal carina often with a flange (Fig, 6F, 7B, 20E, 21B cf. Figs
(Fig.
Perilampus sirsiris is a hyperparasitoid, a parasitoid of dipteran and hymenopteran parasitoids of Lepidoptera, rarely of hymenopteran parasitoids of argid sawflies. Hosts: Tachinidae (Diptera) from Hyphantria cunea (Drury) (Erebidae) and Malacosoma disstria Hübner (Lasiocampidae). Sarcophagidae (Diptera) from Neophasia menapia (C. & R. Felder) (Pieridae). Braconidae (Hymenoptera). Cotesia hyphantriae (Riley) from Hyphantria cunea (Drury). Ichneumonidae (Hymenoptera) from Arge sp. (Hymenoptera).
There is a rare variant from Manitoulin Island, Ontario, a male (ROME152661) which has a wide bare area without setae on the clypeus similar to P. monocteni, but confirmed as P. sirsiris by the steeply curved parascutal carina and COI and ITS2.
The descriptions of P. sirsiris provided in
The steeply curved or angulate parascutal carina often with a flange (Figs
Perilampus sirsiris parasitizes dipteran and hymenopteran parasitoids of Lepidoptera, which feed on the leaves of deciduous trees. Interestingly, the hosts of P. sirsiris also include sarcophagid parasitoids of the pine butterfly, N. menapia (Pieridae)—this is the only species associated with pines other than P. neodiprioni, the hypothesized sister species of P. sirsiris (Fig.
Canada, Ontario, Haliburton County.
Holotype. “CANADA: ONT. Haliburton Hwy 16, 3.8 m. E. Minden Ex: Neodiprion lecontei in red pine plantation. VIII.28.93. DC Darling”, “Lab Reared 1994 S. Perlman M.Sc. thesis”, “BOLD COI-5P Sequence, 325bp”. The holotype is card-mounted (Female ROME183975,
Paratypes. Canada: 3 males. Ontario: 3 males. Nipissing Dist., Algonquin P.P., Cameron Road: (3 males: ROME152669-
Canada: 79 females, 97 males. USA: 36 females, 33 males. (Suppl. materials).
Belize: 9 females, 4 males. Stann Creek District: 9 females, 4 males. 4 1/2 mis., Stann Creek Valley: (4 females: ROME185928-USNM; ROME185929-USNM; ROME199572-USNM; ROME199573-USNM. 1 male: ROME185926-USNM); 5 1/2 mi Stann Creek Valley: (1 male: ROME201411-FSCA); Stann Creek Valley: (5 females: ROME185927-USNM; ROME199574-USNM; ROME199575-USNM; ROME199576-USNM; ROME199578-USNM. 2 males: ROME199571-USNM; ROME199577-USNM).
The specific epithet is a noun in the genitive case meaning “of Neodiprion”, in reference to the species’ predilection for pine sawflies and their primary parasitoids.
Female (Fig.
Perilampus neodiprioni Female A habitus, lateral view B habitus, dorsal view C lateral lobe of mesoscutum along notaulus D lateral panel of pronotum, posterior oblique view E axillula F axilla G head, dorsal view H head, anterior view I clypeus J parascutal carina K mesoscutellum apex L, M mesofemoral depression. [A–E, G, H, J, L Paratype, ROME162273; F, M Paratype, ROME198146; I ROME181757; K ROME189061]. Scale bar: 1 mm (A).
Head (Fig.
Mesosoma (Fig.
Male (Fig.
Perilampus neodiprioni Male A habitus, lateral view B lateral panel of pronotum, parascutal carina, and axilla C head, anterior view D head, dorsal view E lower face F lateral lobe of mesoscutum along notaulus G, H scape. [A–D, F Paratype, ROME162274; E ROME185922; G ROME152634; H Paratype, ROME97556]. Scale bars: 1 mm (A); 100 μm (G, H).
Perilampus neodiprioni can usually be distinguished by an axillula with one or more piliferous punctures dorsad (Fig.
(Fig.
Perilampus neodiprioni can develop as a primary parasitoid attacking Neodiprion sawflies (Fig.
Both COI and ITS2 support P. neodiprioni as a distinct species (Fig.
Genetic analysis suggests there are at least two distinct COI clades of P. neodiprioni in the Nearctic region: Ontario and Massachusetts; and Virginia, West Virginia, and Texas (Suppl. material
Perilampus neodiprioni is the only species in the P. hyalinus species complex that exhibits an exclusive association with pine sawflies, more commonly as a primary parasitoid but also as a hyperparasitoid. An exception is a single P. neodiprioni specimen reared from Diprion similis (Hartig) in Ontario (ROME207314), but it lacks associated host remains and the collector had noted the uncertainty in their identification in the collection form. While it isn’t surprising that P. neodiprioni can develop on D. similis, this sawfly species is non-native in the Nearctic region and not relevant to the evolutionary history of P. neodiprioni.
A large number of P. neodiprioni specimens were reared from Neodiprion lecontei cocoons in the 1940s at the Dominion Parasite Laboratory (DPL) in Belleville, Ontario and subsequently transferred to the
Neodiprion species are often serious pests in boreal forests (
The shift in ecology from hyperparasitoid associated with Lepidoptera to primary or hyperparasitoid associated with pine sawflies is suggested by the sister species relationship between P. neodiprioni and P. sirsiris (Suppl. material
Canada, Ontario, Peterborough County, Aspley.
Holotype. “CANADA, Ontario, Aspley No. S64-3469-01, 22.III. 1965 Ex. Monoctenus juniperinus On e.w. cedar, Lot 65.418”. The Monoctenus associated with the holotype was later re-identified as M. fulvus (Kevin Barber, personnel communication). The holotype is point-mounted (Female ROME201079,
Paratypes. Canada: 7 females, 2 males. Ontario: 7 females, 2 males. City of Ottawa., Bells Corners: (5 females: ROME207334-
Canada: 1 female, 1 male. Ontario: 1 female, 1 male. City of Ottawa., Bells Corners: (1 male: ROME207330-
The specific epithet is a noun in the genitive case meaning “of Monoctenus”, in reference to the species’ host preference for Monoctenus sawflies and their parasitoids.
Female (Fig.
Perilampus monocteni Female A habitus, lateral view B habitus, dorsal view C lateral lobe of mesosucutm along notaulus D lateral panel of pronotum, posterior oblique view E parascutal carina F axilla G head, dorsal view H head, anterior view I, J clypeus K mesoscutellum apex L mesofemoral depression. [A, B, D, K Paratype, ROME201078; C Paratype, ROME207336; E, J, L Holotype, ROME201079; F Paratype, ROME207331; G–I Paratype, ROME207334]. Scale bar: 1 mm (A).
Head (Fig.
Mesosoma (Fig.
Male (Fig.
Perilampus monocteni can be distinguished by a clypeus with a wide and bare area without setae near the dorsal margin, which is extended ventrad medially (Figs
(Fig.
Perilampus monocteni can develop as a primary parasitoid of cedar sawflies (Fig.
A female and male reared from Monoctenus spp. have slight variations in the clypeal setae. One has two setae on the mostly bare area of the clypeus dorsad (ROME201101), and the other lacks a distinctive bare area on clypeus dorsad (ROME207330). The host records and absence of diagnostic characters specific to the other species suggest that these two specimens are P. monocteni.
The available specimens of P. monocteni are unsuitable for the sequencing method used in this study. Despite the lack of genetic data, the species hypothesis of P. monocteni is supported by the distinctive setal distribution pattern on the clypeus and a unique host association. Perilampus monocteni is a paraisitoid strictly associated with Diprionidae similar to P. neodiprioni. However, P. monocteni parasitizes cedar sawflies and their parasitoids, unlike P. neodiprioni which parasitizes pine sawflies and their parasitoids. Of the total six specimens associated with Monoctenus cocoons, three were primary parasitoids of cedar sawflies (ROME207330, 207332, and 207336) and three were hyperparasitoids that parasitized dipteran parasitoids of cedar sawflies. (ROME207328, 207333, and 207335). The known distribution is restricted to eastern Ontario but this may be due to the solitary larval feeding behaviour of Monoctenus spp. (Rose et al. 2010)—in comparison, many Neodiprion spp. are characterized by gregarious larval feeding behaviour (
USA, Florida, Gainesville.
Holotype. “USA: FL: Alachua Co.: nr. Gainesville airport, 45 m 29°42'0"N, 82°15'40"W 2.Oct.2016 A. Baker, A. Knyshov, J. Zhang swp AB16.028". The holotype is point-mounted (Female ROME182771,
Paratypes. USA: 1 female, 2 males. Florida: 1 female, 2 males. Putnam Co., Ordway-Swisher Biol. Station, Rd. C6: (1 female: ROME189115-
USA: 9 females, 4 males. (Suppl. materials).
Cuba: 1 female. (1 female: ROME189093–USNM).
The specific epithet is the Latin adjective crassus (coarse), in reference to the punctate sculpture on the lateral lobes of the mesoscutum along notaulus.
Female (Fig.
Perilampus crassus Female A habitus, lateral view B habitus, dorsal view C lateral lobe of mesoscutum along notaulus D lateral panel of pronotum, posterior oblique view E parascutal carina F axilla G head, dorsal view H head, anterior view I lower face J head, lateral view K mesoscutellum apex L mesofemoral depression. [A–H, J, L Holotype, ROME182771 I Paratype, ROME189085 K Paratype, ROME189115]. Scale bar: 1 mm (A).
Head (Fig.
Mesosoma (Fig.
Male (Fig, 13). Length: usually smaller, 3.0–3.8 mm. As in female, except: Frontal carina (Fig.
Perilampus crassus Male A habitus, lateral view B lateral panel of pronotum, parascutal carina, and axilla, posterior oblique view C head, anterior view D head, dorsal view E lower face F lateral lobe of mesoscutum along notaulus G, H scape. [A, B, G Paratype, ROME189063; C, E, F Paratype, ROME189062; H Paratype, ROME189117]. Scale bars: 1 mm (A); 100 μm (G, H).
Perilampus crassus can be distinguished by a weakly iridescent and nearly straight ventral margin of clypeus (Figs
(Fig.
Hosts unknown.
This species is supported by both genes (Fig.
USA, Texas, 3.5 mi SE La Sauceda.
Holotype. “USA, Texas, Presidio Co. Big Bend Ranch SNA McGuirks Tanks on desert willows 12.V.1990, R Wharton”. The holotype is point-mounted (Female ROME182765, TAMU). BOLD:AEF0151/ITS2. ROM Online Collection.
Paratypes. USA: 3 females, 3 males. Arizona: 2 females. Graham Co., Pinaleno Mountains, Ash Creek near Cluff Ranch Wildlife Area, 14 km SW Pima, 32°47.69'N, 109°51.42'W: (1 female: ROME152679-
Mexico: 5 females. USA: 19 females, 29 males. (Suppl. materials).
Mexico: 1 female. Sonora: 1 female. (1 female: ROME189096-
The specific epithet is the Latin adjective pilosus (hairy), in reference to the densely setose face.
Female (Figs
Perilampus pilosus Female A habitus, lateral view B habitus, dorsal view C lateral lobe of mesoscutum along notaulus D lateral panel of pronotum, posterior oblique view E parascutal carina F axilla G head, dorsal view H head, anterior view I lower face J head, lateral view K mesoscutellum apex L mesofemoral depression. [A, H Holotype, ROME182765; B, C, E–G, L Paratype, ROME182819; I, J Paratype, ROME189067; K ROME189129; L Paratype, ROME152679]. Scale bar: 1 mm (A).
Head (Fig.
Mesosoma (Fig.
Male (Fig.
Perilampus pilosus can be distinguished by the combination of an advanced median ocellus (Figs
Perilampus pilosus Female A habitus, dorsal view. Male B habitus, lateral view C head, anterior view D head, dorsal view E lower face F, G scape. [A Paratype, ROME189067; B Paratype, ROME152727; C, D, F Paratype, ROME182757; E Paratype, ROME152714; G Paratype, ROME152725]. Scale bars: 1 mm (B); 100 μm (F, G).
(Fig.
Perilampus pilosus is a hyperparasitoid, parasitizing dipteran parasitoids of Lepidoptera. Hosts: Tachinidae (Diptera). Chaetogena sp. from Hemileuca juno Packard (Saturniidae).
A female from Sonora, Mexico (ROME189096) has the frontal carina close to the lateral ocellus (FCLO/OD about 0.5) and iridescent olivaceous head.
This species is supported by both genes (Fig.
Canada, Ontario, Chaffey’s Locks.
Holotype. “CANADA, ONT: Frontenac Co., Chaffey’s Locks Oct. 6, 1987 DC Darling. Ex: birch”, “LAB REARED Ex: tachinid parasite of Hyphantria cunea”. The holotype is point-mounted (Female ROME183977, ROME). BOLD:ACF3436. ROM Online Collection.
Paratypes. Canada: 1 male. Ontario: 1 male. Essex Co., Windsor: (1 male: ROME162263-ROME; BOLD:ACF3436; ITS2). USA: 5 females, 1 male. Indiana: 1 female. Posey Co., Harmonie State Park: (1 female: ROME182769-TAMU; BOLD:ACF3436; ITS2). Kentucky: 1 female, 1 male. Jessamine Co., S. of Nicholasville, 37°47'4"N, 84°34'11"W: (1 female: ROME158541-ROME; BOLD:ACF3436; ITS2). Missouri: 1 female. St. Louis Co., St. Louis: (1 female: ROME185906-ROME; BOLD:ACF3436; ITS2). Texas: 1 female. Walker Co., Huntsville: (1 female: ROME185911-TAMU; BOLD:ACF3436; ITS2). West Virginia: 1 female. Hardy Co., 3 mi NE Mathias, 38.9098, -78.8881: (1 female: ROME198142-USNM; BOLD:ACF3436; ITS2).
Canada: 13 females, 17 males. USA: 38 females, 29 males. (Suppl. materials).
USA: 2 females, 3 males. Florida: 2 females, 3 males. Alachua Co., Gainesville: (1 male: ROME204136-SEMC). (2 males: ROME207380-FSCA; ROME207381-FSCA). Orange Co., Orlando, UCF Campus: (1 female: ROME152729-
The specific epithet is a noun in apposition—a reference to both the Seneca people who are the original inhabitants of the core range of the species and to the county in New York State where most of the specimens were reared.
Female (Fig.
Perilampus seneca Female A habitus, lateral view B habitus, dorsal view C lateral lobe of mesoscutum along notaulus D lateral panel of pronotum, posterior oblique view E parascutal carina F axilla G head, dorsal view H head, anterior view I lower face J head, lateral view K mesoscutellum apex L mesofemoral depression. [A, H, I, J, L Paratype, ROME199563; B, F, G Paratype, ROME182769; C Paratype, ROME158541; D Paratype, ROME199544; E Holotype, ROME183977; K Paratype, ROME198142]. Scale bar: 1 mm (A).
Head (Fig.
Mesosoma (Fig.
Male (Fig.
Perilampus seneca Male A habitus, lateral view B lateral panel of pronotum, parascutal carina, and axilla, posterior oblique view C head, anterior view D head, dorsal view E habitus, dorsal view F lateral lobe of mesoscutum along notaulus G, H scape. [A–C, F, G Paratype, ROME183976; D Paratype, ROME199551; E Paratype, ROME162263; H Paratype, ROME199552]. Scale bars: 1 mm (A); 100 μm (G, H).
Perilampus seneca is most similar to P. ute, but females can be distinguished by the lateral panel of pronotum which has a small triangular flange (Fig.
(Fig.
Perilampus seneca is a hyperparasitoid, primarily parasitizing dipteran and hymenopteran parasitoids of Lepidoptera (Fig.
An unsequenced male from Florida (ROME204136), has a violet mesonotum and two sequenced females from Florida (ROME152728, ROME152729) lack black coloration between the frontal carina and median ocellus. And two unsequenced males from Florida (ROME207380, ROME207381) reared from Anisomorpha buprestoides (Pseudophasmatidae, Phasmida) have a violet mesonotum and the lateral lobe of mesoscutum is weakly punctate along notaulus.
There are uncertainties about the species limit of P. seneca. Although P. seneca is differentiated from the other Nearctic species in COI (10 BINed specimens on BOLD, ACF3436), there are Neotropical clades (from Argentina, Costa Rica, and Venezuela) with BIN ACF3436 that cannot be delimited from P. seneca and each other. Specimens either morphologically indistinguishable from P. seneca (“P. hyalinus 1” from Costa Rica and Venezuela) or differ only in the body coloration or subtle male scape morphology (“P. hyalinus 2, 3, and 16, from Argentina, Costa Rica, and Venezuela, respectively). Perilampus seneca and these Neotropical groups are placed together as a single clade in COI (Suppl. material
Despite these uncertainties we hypothesize that P. seneca is a recently diverged Nearctic species distinct from the Neotropical groups with similar COI sequences—the populations north of Florida are monophyletic for COI albeit with poor support in BI (Suppl. materials
The relationships between P. seneca and the Neotropical specimens shown in COI unsupported by ITS2 may be due to retention of ancestral polymorphism via incomplete lineage sorting (ILS), introgression events between the Neotropical lineages, and/or insufficient phylogenetic information in ITS2 for this group. The paraphyly of P. seneca may be explained by ILS instead of hybridization due to the geographical distance between the two groups. However, two females from Florida (ROME152729, ROME152728) without black coloration between the frontal carina and lateral ocellus, and unsequenced males from Florida (ROME204136, ROME207380, ROME207381) with entirely violet body color in contrast to cupreous body color of northern P. seneca suggest there could be two independently evolving lineages in the Nearctic region.
A single Perilampus was reared as a parasitoid of Tachinidae in Kentucky, USA (ROME174210) from a series of 1,139 tachinid primary parasitoids of Acalymma vittatum and Diabrotica undecimpunctata howardi (Chrysomelidae) (
USA, Colorado, Idledale.
Holotype. “USA, Colorado, Jefferson Co., Idledale, Sawmill Gulch, 1981 m, 39°40'N, 105°14'W, 20–27.viii.2001, Malaise, Irwin, Lambkin, Metz & Hauser”. The holotype is point-mounted (Female ROME182768, TAMU). BOLD:AEE9091/ITS2. ROM Online Collection.
Paratypes. USA: 3 females, 1 male. Arizona: 1 female. Cochise Co., Coronado National Forest, Huachuca Mts., Copper Canyon, 31°21'44"N, 110°18'02"W: (1 female: ROME182763-TAMU; BOLD:AEE9091; ITS2). California: 1 male. San Bernardino Co., Kellers Peak, 34°12'22"N, 117°02'36"W: (1 male: ROME182781-
USA: 9 females, 5 males. (Suppl. materials).
Mexico: 1 male. Jalisco: 1 male. (1 male: ROME200745-HNHM). USA: 1 male. California: 1 male. Mono Co., Golden Gate Mine, 4.6 mi NW Walker: (1 male: ROME201998-
The specific epithet is a noun in apposition—a reference to the Ute, indigenous people of the Great Basin regions of present-day Utah and Colorado where the holotype was collected.
Female (Fig.
Perilampus ute Female A habitus, lateral view B habitus, dorsal view C lateral lobe of mesoscutum along notaulus D, E lateral panel of pronotum, posterior oblique view F axilla G head, dorsal view H head, anterior view I lower face J head, lateral view K mesoscutellum apex L mesofemoral depression. [A, C, D, F, G, H Holotype, ROME182768; B, E, I–L Paratype, ROME182763]. Scale bar: 1 mm (A).
Head (Fig.
Mesosoma (Fig.
Male (Fig.
Perilampus ute Male A habitus, lateral view B, C lateral panel of pronotum, posterior oblique view D head, anterior view E head, dorsal view F habitus, dorsal view G lateral lobe of mesoscutum along notaulus H, I Scape. [A, B, D–F, H Paratype, ROME182781; C, G, I Paratype, ROME198139]. Scale bars: 1 mm (A); 100 μm (H, I).
Perilampus ute can usually be distinguished by the lateral panel of pronotum with an expanded triangular flange in posterior oblique view (Figs
(Fig.
Perilampus ute is a hyperparasitoid, parasitizing dipteran parasitoids of Lepidoptera. Hosts: Tachinidae (Diptera), Lespesia aletiae (Riley) from Apatelodes pudefacta Dyar (Apatelodidae).
An unsequenced male from California (ROME201998) has a greenish iridescence along the midline of a mesonotum with a weak cupreous iridescence laterad.
Perilampus ute is recovered as monophyletic (Fig.
USA, West Virginia, Hardy County, 3 mi NE Mathias.
Holotype. “WEST VIRGINIA: Hardy Co. 3 mi NE Mathias 38.9098, -78.8881, 14–31.VII.2007, Malaise David R. Smith”. The holotype is point-mounted (Female ROME189051, USNM). BOLD:AEE7608/ITS2. ROM Online Collection.
Paratypes. Canada: 1 male. Ontario: 1 male. Norfolk Co., Normandale Fish Hatchery, 42°43'08"N, 80°20'23"W: (1 male: ROME198214-
Canada: 2 females, 1 male. USA: 6 females, 1 male. (Suppl. materials).
The specific epithet is the Latin noun arcus (arch), in reference to the steeply curved parascutal carina.
Female (Fig.
Perilampus arcus Female A habitus, lateral view B habitus, dorsal view C lateral lobe of mesoscutum along notaulus D lateral panel of pronotum, posterior oblique view E parascutal carina F axilla G head, dorsal view H head, anterior view I lower face J head, lateral view K mesoscutellum apex L mesofemoral depression. [A, D, E Holotype, ROME189051; B, C, H, J, L Paratype, ROME185944; F Paratype, ROME189050; G Paratype, ROME158551; I, K Paratype, ROME189131]. Scale bar: 1 mm (A).
Head (Fig.
Mesosoma (Fig.
Male (Fig.
Perilampus arcus Male A habitus, lateral view B lateral panel of pronotum, parascutal carina, and axilla C head, anterior view D head, dorsal view E lower face F lateral lobe of mesoscutum along notaulus G, H scape. [A–E, G Paratype, ROME185954; F, H Paratype, ROME198214]. Scale bars: 1 mm (A); 100 μm (G, H).
The steeply curved parascutal carina often with a flange distinguishes P. arcus from the majority of the species of the P. hyalinus species complex (Figs
(Fig.
Unknown.
This species is supported by both genes (Fig.
USA, Arizona, Santa Cruz County, Sonoita.
Holotype. “ARIZONA: Santa Cruz Co. Sonoita, 2 mi S of town center 31°38'N, 110°39'W, 16–22. VI.2008 Malaise trap in juniper/oak grasslands, EE Grissell”. The holotype is point-mounted (Female ROME152670, USNM). BOLD:AEO0861/ITS2. ROM Online Collection.
Paratypes. Mexico: 1 male. Oaxaca: 1 male. 19 mi S San Miguel Suchixtepec at Puente Jalatengo: (1 male: ROME182754-TAMU; BOLD:AEO0861; ITS2). USA: 1 female, 1 male. Arizona: 1 female, 1 male. Cochise Co., Coronado National Forest, Chiricahua Mts., 1 mi N Rustler Park, 31°54'53"N, 109°16'07"W: (1 female: ROME186060-TAMU; BOLD:AEO0861; ITS2). Santa Cruz Co., (1 male: ROME198215-USNM; ITS2).
Mexico: 15 females, 3 males. USA: 3 females. (Suppl. materials).
USA: 1 female. Arizona: 1 female. Cochise Co., Bisbee, 1429 Franklin St., 31°24'23.8"N, 109°55'57.6"W: (1 female: ROME152671-USNM; BOLD:AEO0861; ITS2).
The specific epithet is a noun in apposition—a reference to both the Sonoran Desert and to the state of Mexico where most of the specimens were collected.
Female (Fig.
Perilampus sonora Female A habitus, lateral view B habitus, dorsal view C lateral lobe of mesoscutum along notaulus D lateral panel of pronotum, posterior oblique view E parascutal carina F axilla G head, dorsal view H head, anterior view I lower face J head, lateral view K mesoscutellum apex L mesofemoral depression. [A, B, D, G, H, J, K Holotype, ROME152670; C, E, I Paratype, ROME186060; L Paratype, ROME189110]. Scale bar: 1 mm (A).
Head (Fig.
Mesosoma (Fig.
Male (Fig.
Perilampus sonora Male A habitus, lateral view B lateral panel of pronotum, parascutal carina, and axilla, posterior oblique view C head, anterior view D head, dorsal view E lower face F lateral lobe of mesoscutum along notaulus G, H scape. [A–F, G Paratype, ROME182754; H Paratype, ROME198215]. Scale bars: 1 mm (A); 100 μm (G, H).
This species is distinguished by the strong cupreous iridescence on the mesonotum of both females and males (Figs
(Fig.
Type specimens of the previously described species of the Perilampus hyalinus species complex A P. entellus Walker (synonym of P. hyalinus Say), lectotype, female B P. aciculatus Provancher (synonym of P. hyalinus Say), lectotype, male C P. sirsiris (Argaman), holotype, female D P. americanus Girault, holotype, female E, F P. nigriviridis Girault, lectotype, male. Photo credits: A ©The Trustees of the Natural History Museum, London (Licensed under CC BY 4.0) B Joseph Moisan-De Serres (Ministry of Agriculture, Fisheries and Food in Québec) C National Museum of Natural History, Paris D, E, F Museum of Natural History, Berlin.
Distribution of the Nearctic species of the Perilampus hyalinus species complex A P. hyalinus (green circles) and P. crassus (red squares) B P. neodiprioni (cyan circles) P. monocteni (red stars) C P. sirsiris D P. seneca E P. arcus F P. pilosus (yellow stars) P. ute (blue triangles) P. sonora (red circles).
Perilampus sonora is a hyperparasitoid, parasitizing dipteran and hymenopteran parasitoids of Lepidoptera. Hosts: Probably Tachinidae (Diptera) and/or Ichneumonoidea (Hymenoptera) from Utetheisa ornatrix (Linnaeus) (Erebidae). The potential hosts are: Gymnosoma sp. (Tachinidae), Lespesia sp. (Tachinidae), and Cotesia sp. (Braconidae) (G. R. Buckingham, personal communication).
A female from Arizona (ROME152671) has an iridescent green head and weakly iridescent green midlobe of mesoscutum and scutellum. COI and ITS2 suggest that this specimen is a morphological variant of P. sonora. A female from Sonora, Mexico (ROME189103), has a strongly transverse clypeus with CW/CH about 1.7.
Cocoons, pupa, and host remains associated with reared specimens of Perilampus. A–C, E Neodiprion lecontei cocoons A, B P. neodiprioni as primary parasitoid C, E P. neodiprioni as hyperparasitoid, parasitoid of Ichneumonidae D Neodiprion virginianus cocoon, P. neodiprioni as hyperparasitoid, parasitoid of Tachinidae [insert: tachinid puparium spiracles] F Lepidoptera pupa, P. seneca as hyperparasitoid, parasitoid of Tachinidae G, H Monoctenus suffusus cocoons G P. monocteni as primary parasitoid H P. monocteni as hyperparasitoid, parasitoid of Tachinidae [insert: tachinid puparium spiracles]. Abbreviations: eh, emergence hole; ex, Perilampus pupa exuvia; Ic, ichneumonid cocoon; Lp, lepidopteran pupa; M, Monoctenus remains; Mc, Monoctenus cocoon; N, Neodiprion remains; Nc, Neodiprion cocoon; Tp, tachinid puparium. [associated Perilampus: A, B ROME207376 C ROME207377 D ROME198223 E ROME162273 F ROME202000 G ROME207330 H ROME207328]. Scale bars: 1 mm.
The distribution ranges of P. sonora and its undescribed Neotropical sister species, P. hyalinus 17 (Fig.
There is considerable complexity in the host associations of the P. hyalinus species complex and the host associations and modes of parasitism are congruent with the newly delimited species. Perilampus hyalinus Say is a hyperparasitoid that parasitizes Tachinidae and Sarcophagidae (Diptera) parasitoids of Orthoptera or a parasitoid of dipteran kleptoparasites of Crabronidae and Sphecidae (Hymenoptera) that provision their nests with Orthoptera. Two new species are associated with pine sawflies or cedar sawflies, (Diprionidae) and are able to develop as both primary parasitoids or as hyperparasitoids that parasitize the tachinid and ichneumonid parasitoids of diprionid sawflies. And four new species are described for hyperparasitoids, associated with Lepidoptera, parasitizing Tachinidae or Ichneumonoidea.
The molecular analyses and outgroup comparison with the P. platigaster species group suggests that hyperparasitoids are basal in the P. hyalinus species group and development as a primary parasitoid is derived. These and other host shifts may have been facilitated by parasitism involving the mobile planidial first-instar larva. Planidia that successfully make contact and burrow into a novel but suitable host (e.g. sawfly larva) could successfully develop which may ultimately have resulted in the evolution of the species capable of developing as primary parasitoids, e.g., P. neodiprioni and P. monocteni. Planidia that make contact with a novel but unsuitable host (e.g. Orthoptera) may be “rescued” by subsequent parasitism by primary parasitoids (e.g. Diptera or Hymenoptera) on which the planidia can develop (
The morphological characters used to distinguish the species in the P. hyalinus species complex are often subtle (e.g. widely vs deeply curved parascutal carina) and can show interspecific variation (e.g. the sculpture of axillula in P. neodiprioni varies between punctate and smooth). Despite the fine distinctions, these morphological characters should allow the identification of most specimens in the absence of host information or molecular data, particularly series of specimens with males and females. It is noteworthy that eight of ten Nearctic species were successfully delimitated by COI and ITS and that each species has a unique BIN number and is monophyletic on the maximum likelihood and Bayesian trees of the P. hyalinus species group (Fig.
The phylogenetic analyses of COI and ITS2 suggest a Neotropical origin of the P. hyalinus species complex (Fig.
This revision provides resolution to 100 years of the confusion surrounding P. hyalinus in the Nearctic region. However, the phylogenetic relationships and species diversity of the P. hyalinus species complex in the Mexican transition zone and Neotropical region needs further study (
We would like to thank Andrew Bennett and James O’Hara from
Bayesian inference 50% majority consensus tree for Perilampus hyalinus species group based on concatenated dataset of COI and ITS2 (
Data type: tif
Explanation note: Bayesian inference 50% majority consensus tree for Perilampus hyalinus species group based on concatenated dataset of COI and ITS2 (
Maximum likelihood trees of Perilampus hyalinus species group retrieved from the separate analyses of COI and ITS2, and the results of morphological analyses (M) and molecular species delimitation (ABGD, BIN, and PTP) (modified from
Data type: zip
Explanation note: Maximum likelihood trees of Perilampus hyalinus species group retrieved from the separate analyses of COI and ITS2, and the results of morphological analyses (M) and molecular species delimitation (ABGD, BIN, and PTP) (modified from
Bayesian inference 50% majority consensus tree for Perilampus hyalinus species group based on COI (
Data type: tiff
Explanation note: Bayesian inference 50% majority consensus tree for Perilampus hyalinus species group based on COI (
Bayesian inference 50% majority consensus tree for Perilampus hyalinus species group based on ITS2 (
Data type: tiff
Explanation note: Bayesian inference 50% majority consensus tree for Perilampus hyalinus species group based on ITS2 (
The summary of congruence between Perilampus hyalinus species group morphospecies, phylogenetic analyses, the molecular species delimitation methods (ABGD, BIN, PTP) (modified from
Data type: xlsx
Explanation note: The summary of congruence between Perilampus hyalinus species group morphospecies, phylogenetic analyses, the molecular species delimitation methods (ABGD, BIN, PTP) (modified from
The type specimens of species previously described in the Perilampus hyalinus species group
Data type: xlsx
Explanation note: The type specimens of species previously described in the Perilampus hyalinus species group.
Material examined
Data type: docx
Explanation note: Material examined for each species other than the types and additional material.