Taxonomic revision of the Perilampus carolinensis species complex (Hymenoptera: Chalcidoidea: Perilampidae), and the description of five new species Author Yoo, Jeong Jae Department of Natural History, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario, Canada, M 5 S 2 C 6 & Department of Ecology and Evolutionary Biology, University of Toronto, Toronto, Ontario, Canada, M 5 S 3 B 2 Author Darling, D. Christopher 0000-0002-1207-7967 Department of Natural History, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario, Canada, M 5 S 2 C 6 & Department of Ecology and Evolutionary Biology, University of Toronto, Toronto, Ontario, Canada, M 5 S 3 B 2 cdarling@rom.on.ca text Zootaxa 2025 2025-04-15 5621 2 151 195 https://doi.org/10.11646/zootaxa.5621.2.1 journal article 309444 10.11646/zootaxa.5621.2.1 b63f192e-9851-4f23-b885-f6c398a23fa2 1175-5326 15262606 40FC7108-3276-4EC0-92D0-E5BAE41BC096 Perilampus carolinensis Smulyan Figs 3 [ ♀ ], 4[ ♂ ] Perilampus carolinensis Smulyan, 1936: 376 . (Original description, keyed.) Taltonos carolinensis (Smulyan) . Argaman, 1990: 204 . Perilampus carolinensis Smulyan. Darling, 1996 ( Taltonos subjective synonym of Perilampus ). Type Material. Holotype [ ♀ ]. USA : Virginia . "Roslyn, Va. Sept. 3-1923 . Captured on thistle "; USNM , Type No. 49777, USNMENT01559545 , (image examined, http://n 2t .net/ark:/65665/3ce6ad39f-7095-45ec-9746- 86ac74d1c42f). Also 17 female and 2 male paratypes ( Smulyan 1936 ). Paratypes examined [ 4 ♀ ]. USA : North Carolina . Yancey Co. , Valley of Black Mountains ( 4 ♀ : W Beutenmuller, ROME152649 - AMNH ; W Beutenmuller, ROME152650 - AMNH ; W Beutenmuller, ROME152651 - AMNH ; W Beutenmuller, ROME152652 - AMNH ) . Material examined. Brazil : Minas Gerais . P.N. da Serra da Canastra , São Roque de Minas , 20°27'38.9"S , 46°30'48.9"W ( 1 ♂ : ROME143233 - MZSP ) . São Paulo . Campinas ( 1 ♀ : ROME143111 - USNM . 2 ♂ : ROME143112 - USNM ; ROME143113 - USNM ) . Costa Rica : Guanacaste . Area de Conservacion Guanacaste ( ACG )( 15♀ :06-SRNP-58407, DHJPAR0021672 - CNC ; BOLD : AAA4882 ; 99-SRNP-15036, DHJPAR0023903 - CNC ; BOLD : AAA4882 ; 99-SRNP-3495, DHJPAR0023904 - CNC ; BOLD : AAA4882 ; 91-SRNP-3136.1, DHJPAR0023905 - CNC ; 98-SRNP-9534, DHJPAR0023908 - ROME ; BOLD : AAA4882 ; 93-SRNP-7077, DHJPAR0023911 - CNC ; BOLD : AAA4882 ; 02-SRNP-11966, DHJPAR0023913 - CNC ; BOLD : AAA4882 ; 00-SRNP-19905, DHJPAR0023914 - CNC ; BOLD : AAA4882 ; 00-SRNP-19879, DHJPAR0023915 - CNC ; BOLD : AAA4882 ; 00-SRNP-19879, DHJPAR0023916 - CNC ; BOLD : AAA4882 ; 00-SRNP-19879, DHJPAR0023917 - CNC ; BOLD : AAA4882 / ITS2 ; 06-SRNP-58423, DHJPAR0021669 - MZUCR ; BOLD : AAA4882 ; 00-SRNP-19888, DHJPAR0023918 - ROME ; BOLD : AAA4882 / ITS2 ; 12-SRNP-12669, DHJPAR0051468 - ROME ; BOLD : ACJ4137 / ITS2 ; 12-SRNP-12642, DHJPAR0051480 - ROME ; BOLD : ACJ4137 / ITS2 . 12 ♂ : 04-SRNP-13707, DHJPAR0021330 - ROME ; BOLD : AAA4872 ; 04-SRNP-13718, DHJPAR0021331 - ROME ; BOLD : AAA4872 ; 85-SRNP-552, DHJPAR0023777 - CNC ; 85-SRNP-552, DHJPAR0023778 - CNC ; 99-SRNP-3486.06, DHJPAR0023799 - CNC ; BOLD : AAA4882 / ITS2 ; 00-SRNP-19876, DHJPAR0023919 - CNC ; 00-SRNP-19876, DHJPAR0023920 - CNC ; 20-SRNP-27385, DHJPAR0066296 - ROME ; BOLD : AEK0124 / ITS2 ; 06-SRNP-58474, DHJPAR0021670 - CNC ; BOLD : AAA4882 / ITS2 ; 06-SRNP-58391, DHJPAR0021674 - ROME ; BOLD : AAA4882 / ITS2 ; 06-SRNP-58472, DHJPAR0021677 - MZUCR ; BOLD : AAA4882 / ITS2 ; 06-SRNP-58453, DHJPAR0021678 - ROME ; BOLD : AAA4882 / ITS2 ) . Mexico : Nuevo León . Villa del Carmen ( El Carmen ) ( 1 ♂ : ROME181752 - TAMU ) . USA : Arizona . Santa Cruz Co. , Atascosa Mts , Ruby Rd. ~ 4 mi E of Ruby ( 1 ♀ : ROME204128 - ROME ) . Santa Rita Mts , Montosa Canyon ( 1 ♀ : ROME204126 - ROME ) ; Sycamore Canyon , at FR39 , 31°25.8'N , 111°11.4'W ( 1 ♀ : ROME152657 - CNC ; BOLD : AEU5522 / ITS2 ) . Georgia . Rabun Co. , Satolah ( ♂ : ROME204114 - ESUW ) . Illinois . Cook Co. , Chicago ( 1 ♀ : ROME185937 - FMNH . 1 ♂ : ROME152654 - FMNH ) . Indiana . Starke Co. , Knox ( 1 ♀ : ROME185971 - FMNH ) . Maryland . Prince George’s Co. , Bowie , Patuxent Research Refuge ( 1 ♂ : ROME152656 - FMNH ) . New Jersey . Ocean Co. , Manahawkin ( 2 ♀ : ROME199587 - CUIC ; ROME199588 - CUIC ) . Additional material examined. Mexico : Michoacán . Uruapan ( 1 ♀ : ROME201102 - CNC ) . Description. FEMALE ( Fig. 3 ). Length: 3.8–4.7 mm . Color: head iridescent greenish blue or violet, with or without black coloration on vertex ( Fig. 3F ); mesosoma and metasoma iridescent greenish blue or violet; clypeus ventral margin black; antenna with scape and pedicel weakly iridescent greenish blue or violet, flagellum dark brown or black, lighter ventrad and distad ( Fig. 3I ). Head ( Fig. 3E–I ): in dorsal view transverse, width equal to or slightly greater than twice length, HW/HL 2.0–2.1. Frontal carina: in anterior view weakly to strongly sinuate below midlevel of eye; in dorsal view gradually narrowed V shape around median ocellus, FC/MOD 1.5–1.8 ( Fig. 3E ); distance from lateral ocellus short, FCLO/LOD 0.6– 0.7 ( Fig. 3E ). Scrobal cavity ( Fig. 3G ): in anterior view narrow, SW/HW about 0.4. Ocelli ( Fig. 3E ): a line between anterior margin of lateral ocelli reaching anterior margin of median ocellus. POL/OOL 2.1–2.5. Ocellar ratios LOD: POL: OOL: LOL: 1, 3.2–3.6, 1.3–1.6, 1.0–1.3. Vertex: smooth to wrinkled or with weak transverse striations, with or without large piliferous punctures. Parascrobal area: in lateral view abruptly narrowed slightly below mid-eye height, about 0.4 EH above lower eye margin ( Fig. 3I ); width narrow to wide, PSW/EL 0.3–0.4; sculpture smooth or wrinkled, with or without large piliferous punctures. Gena: with wide and long smooth area along outer eye margin, striate behind. Malar space: MSL/EH about 0.2. Lower face: with setae sparse laterad torulus, and sparse below. Clypeus ( Fig. 3H ): CW/ CH 1.3–1.4; ventral margin concave; setae evenly distributed. Mesosoma ( Fig. 3B–D ): Lateral panel of pronotum: slightly narrower than prepectus, LPP/PPT 0.6–0.8; without flange or with small, rounded flange below level of mesothoracic spiracle in posterior oblique view ( Fig. 3C ). Mesofemoral depression: smooth. Mesoscutum: punctures weakly angulate to rounded, with slightly wide and weakly coriarious interspaces ( Fig. 3B ); lateral lobe weakly punctate with coriarious or smooth interspaces, or smooth along notaulus; parascutal carina broadly curved, acuminate. Mesoscutellum: apex with inner margins gradually diverging; punctures weakly angulate or rounded, with narrow to slightly wide and weakly coriarious interspaces. Axilla ( Fig. 3D ): in lateral view imbricate dorsad, and carinate ventrad. Axillula: smooth dorsad. Fore wing: stigma small, 2.0–2.5× as wide as postmarginal vein width. MALE ( Fig. 4 ). Length: usually smaller, 2.8–4.3 mm . As in female, except: Color: mesonotum nearly black or with cupreous iridescence. Frontal carina ( Fig. 4B ): distance from lateral ocellus shorter, FCLO/LOD 0.4–0.5. Scape ( Fig. 4F, G ): pits sparse, covering 0.3–0.4× scape length. Scrobal cavity in anterior view narrow or wide, SW/HW 0.4–0.5. Diagnosis. Perilampus carolinensis can be recognized by a parascrobal area that is abruptly narrowed slightly below the midlevel of the eye ( Figs 3I , 4E ), a wide frontal carina around the median ocellus ( Fig. 3E , 4B ), a narrow female scrobal cavity in anterior view ( Fig. 3G ), and a male scape with a long pitted area (about 0.3–0.4× scape length, Figs 4F, G ). Distribution. Previously known only from the Nearctic region, here newly recorded for the Neotropical region: USA . ( Arizona , Georgia , Illinois , Indiana , Maryland , New Jersey , North Carolina , Virginia ), Mexico ( Michoacán , Nuevo Leon ), Costa Rica (ACG), and Brazil ( Minas Gerais , São Paulo ). Host association. Perilampus carolinensis is a hyperparasitoid, parasitizing dipteran and hymenopteran parasitoids of Lepidoptera . Hosts: Tachinidae ( Diptera ). Calolydella inflatipalpis Fleming and Wood from Dione juno (Cramer) ( Nymphalidae ) on Erblichia odorata Seem ( Passifloraceae ). Chetogena scutellaris (Wulp) from Orgyia povera Schaus ( Erebidae ) on Byrsonima crassifolia (L.) ( Malpighiaceae ), and Hylesia umbrata Schaus ( Saturniidae ) on Anacardium occidentale L. ( Anacardiaceae ). Hyphantrophaga edwinapui Fleming & Wood from Doa sp. ( Doidae )on Euphorbia colletioides Benth. ( Euphorbiaceae ). Leschenaultia sp. from Ammalo helops (Cramer) ( Erebidae ) on Ficus ovalis (Liebm) ( Moraceae ). Lespesia sp. from Trosia obsolecens Dyar ( Megalopygidae ), and from Parachabora sp. ( Erebidae ) on Ateleia hebert-smithii Pittier ( Fabaceae ). Lespesia aletiae (Riley) from Apatelodes pudefacta (Smith) , and D. juno (Cramer) on E. odorata . Winthemia picea (Walker) from Pseudosphinx tetrio L. ( Sphingidae ) on Plumeria rubra L. ( Apocynaceae ). Winthemia subpicea (Walker) from Arsenura arianae Brechlin on Guazuma ulmifolia Lam. ( Malvaceae ) and Pochota fendleri (Seem.) ( Malvaceae ). Braconidae ( Hymenoptera ). Triraphis bradzlotnicki Sharkey from Megalopygidae on B. crassifolia . Ichneumonidae ( Hymenoptera ). Enicospilus chiriquensis (Cameron) from H. umbrata on A. occidentale . FIGURE 3. Perilampus carolinensis Female. A , habitus, lateral view; B , habitus, dorsal view; C , lateral panel of pronotum, posterior oblique view; D , axilla, lateral view; E , vertex, dorsal view; F , head, dorsal view; G , head, anterior view; H , clypeus; I , head, lateral view. [ A–C , G , H : Paratype, ROME152649; D, I : Paratype, ROME152652; E , F : ROME199587]. Scale bar: 1mm. FIGURE 4. Perilampus carolinensis Male. A , habitus, lateral view; B , head, dorsal view; C , head, anterior view; D , clypeus; E , head, lateral view; F, G , scape. [ A–C : ROME152654; D : DHJPAR0021678; E, F : ROME152656; G : DHJPAR0021330]. Scale bar: 1mm. The following records of lepidopteran associates lack information on the associated parasitoid hosts but likely represent hyperparasitoids Lepidoptera . Anisota senatoria (Smith) ( Saturniidae ) ( Smulyan 1936 ), Anisota virginiensis (Drury) ( Saturniidae ), Datana integerrima Grote & Robinson ( Notodontidae ) ( Smulyan 1936 ). Remarks. Perilampus carolinensis is similar to P. cabecar in having a wide frontal carina around the median ocellus ( Figs 3E , 4B , 5E , 6B cf. Figs 7E , 8B ). The only character distinguishing female P. carolinensis from P. cabecar is the scrobal cavity width in the anterior view of the head, which is narrower in P. carolinensis ( Fig. 3G cf . Fig. 5G ). This character cannot discriminate the males of these two species, as it varies from wide to narrow in both species. But the males can be separated by the length of the pitted area on the scape, which is longer in P. carolinensis ( Figs 4F, G ) and shorter in P. cabecar ( Figs 6F, G ). The two species are also differentiated in COI and ITS2 ( Figs 1, S1 ). There is variation in both morphology and COI in our species concept of P. carolinensis . Piliferous punctures on the lateral lobes of the mesoscutum in Nearctic specimens tend to be larger than those in the Neotropical specimens, and there is a divergence of 2.0–3.7% in COI between the Nearctic and Costa Rican specimens (ACG). However, the sculpture of mesoscutal lateral lobes is variable in some species of the P. hyalinus species group ( Yoo and Darling 2024 ), and therefore not a reliable diagnostic character by itself. Moreover, even greater intraspecific genetic differences are observed among the ACG specimens (maximum distance of 4.3%) that are rendered paraphyletic by the only sequenced Nearctic specimen (ROME182657, Fig. S1 ). A divergence of 4.3% is shown between two females (DHJPAR0051468, DHJPAR0051480), the only P. hyalinus species group specimens recorded to be associated with Ammalo helops from Ficus , and other ACG specimens. It is unclear whether their genetic divergence in COI represents interspecific ecological differentiation or male-biased dispersal driven by host fidelity ( Henry et al. 2007 ) because other specimens associated with different families of caterpillars and plants show no significant genetic differentiations in COI. Two ACG males ( P. carolinensis 2 in Yoo (2023) ; DHJPAR0021330, DHJPAR0021331) have scapes with pitted areas that are slightly shorter than those of the other ACG and Nearctic specimens ( Fig. 4G cf . 4F). Genetic divergence in COI between these two specimens and the other ACG males with the longer pitted area on a scape is 3.4%. It’s unclear whether these differences indicate distinct species, as a larger genetic distance is observed between one male ACG specimen (DHJPAR0066296, not included in the genetic analyses due to lack of trace file for quality verification), which also has a longer pitted area on the scape, and the rest of its morphological conspecifics (3.7–4.7%). Additionally, there is subtle intraspecific variation in the height of the pitted area of male scapes (0.3 vs 0.4× scape length) shown in some species of the P. hyalinus species group ( Yoo and Darling 2024 ). Highly divergent COI sequences resulted in multiple BINs for P. carolinensis (BOLD: AAA4882, BOLD:AAA4872, BOLD:AEU5522, BOLD:AEK0124, and BOLD:ACJ4137). However, except for two aforementioned males with short punctate area of the scape and DHJPAR0066296, which lacks ITS2 data, all specimens show no divergence in ITS2 ( Figure S1 ). Only three specimens from the southern Neotropical region, all from São Paulo , Brazil , were available for examination. These have stronger wrinkled parascrobal area sculpture than most specimens from ACG and the Nearctic region . However , one Nearctic male ( ROME152654 ) also has similarly strong wrinkled parascrobal area sculpture ( Fig. 4B ). The absence of genetic divergences in ITS2 and the lack of concordance between COI and geographical distribution, host information, and subtle morphological variations prevent a precise taxonomic decision on the species boundaries of P. carolinensis . A conservative approach is taken in this study, tentatively considering the P. carolinensis Smulyan as a single widely distributed species with high intraspecific divergence in COI but monophyletic in both COI and ITS2. More specimens and genes are needed, in particular from Brazil , to further test the species boundaries within P. carolinensis . The possibility of high mitochondrial divergences induced by the multiple strains of Wolbachia infections also needs to be explored (Hurst and Jiggins 2005; Kodandaramaiah et al. 2013). Finally, DNA sequencing of the specimens from the vicinity of the type locality (e.g., Virginia) is needed to confirm their conspecificity with the only sequenced Nearctic specimen (Arizona, ROME152657, CNC). The majority of the rearing records indicate that P. carolinensis is a hyperparasitoid, parasitizing Tachinidae or Ichneumonoidea parasitoids of Lepidoptera that feed on angiosperms. An exception is a specimen from Michoacán , Mexico , labelled "ex. Neodiprion sp. " (ROME201102) that lacks associated host remains for verification. This unlikely host association may be the result of mislabeling or an extremely rare association of a planidium of P. carolinensis with a pine sawfly larva.