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Communication

First Record of Dioryctria simplicella (Lepidoptera: Pyralidae) in China: Morphology, Molecular Identification, and Phylogenetic Position

by
Niya Jia
†,
Xiyao Ding
†,
Dan Xie
,
Huanwen Chen
,
Defu Chi
* and
Jia Yu
*
The Key Laboratory of Sustainable Forest Ecosystem Management-Ministry of Education, College of Forestry, Northeast Forestry University, Harbin 150040, China
*
Authors to whom correspondence should be addressed.
†
These authors contributed equally to this work.
Insects 2025, 16(7), 664; https://doi.org/10.3390/insects16070664
Submission received: 5 June 2025 / Revised: 22 June 2025 / Accepted: 22 June 2025 / Published: 26 June 2025
(This article belongs to the Section Insect Systematics, Phylogeny and Evolution)
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Simple Summary

The study reports on the occurrence of Diocrytria simplicella Heinemann, 1865, a potential forestry pest, in China for the first time. Based on detailed morphological comparisons, we provide a systematic description of this species, including key diagnostic characters of the adult external features and genitalia structures. Additionally, molecular identification based on the mitochondrial COI gene, supported by phylogenetic analysis, further corroborates its taxonomic placement within the genus Dioryctria. This work expands the known distribution of D. simplicella and contributes to the understanding of its systematics.

Abstract

Dioryctria Zeller, 1846 (Lepidoptera: Pyralidae) is a significant genus whose species primarily infest coniferous trees and are predominantly distributed across the Northern Hemisphere. To date, 17 species within this genus have been recorded in China. This study reports the discovery of Dioryctria simplicella (Heinemann, 1863) in China. During field surveys in forests of Heilongjiang Province, D. simplicella was observed infesting the cones and trunks of Pinus sylvestris var. mongolica Litv. as larvae. Comprehensive morphological descriptions and diagnostic characteristics of the adult, larva, pupa, and egg stages of D. simplicella are provided herein to facilitate accurate species identification within the genus. Molecular phylogenetic analysis based on mitochondrial cytochrome c oxidase subunit I (COI) DNA barcoding sequences was conducted to assess the phylogenetic position of D. simplicella within Dioryctria. These results strongly support its species identity and clarify its phylogenetic relationships with congeners. This discovery not only expands the known diversity of Lepidoptera in China but also provides new data supporting taxonomic and phylogenetic studies of the genus Dioryctria.

1. Introduction

The genus Dioryctria (Lepidoptera: Pyralidae), established by Zeller in 1846, is primarily distributed across the Holarctic region, with a few species extending into the northern tropics [1,2]. The larvae of the Dioryctria genus feed on various coniferous trees and are significant forest pests [3]. They typically bore into the trunks, new shoots, and cones of Pinus, Picea, Abies, and Larix species, causing substantial damage to tree growth, timber quality, seed production, and landscape value [4,5,6,7,8,9]. To date, 86 Dioryctria species have been documented worldwide, with 12 species groups currently recognized based on morphological characteristics [4,10,11,12,13,14,15,16]. There are 17 Dioryctria species recorded in China [13,17,18,19,20,21], including D. abietella [Denis & Schiffermüller, 1775], D. castanea [Bradley, 1969], D. fanjingshana [Li, 2009], D. magnifica [Munroe, 1958], D. auloi [Barbey, 1930], D. kunmingnella [Wang & Sung, 1985], D. mendacella [Staudinger, 1859], D. mongolicella [Wang & Sung, 1982], D. pryeri [Ragonot, 1893), D. reniculelloides [Mutuura & Munroe, 1973], D. resiniphila [Segerer & Pröse, 1997], D. rubella [Hampson, 1901], D. schutzeella [Fuchs, 1899], D. splendidella [Herrich-Schäffer, 1848), D. sylvestrella [Ratzeburg, 1840], D. yiai [Mutuura & Munroe, 1972], and D. yuennanella [Caradja, 1937].
The taxonomic status of Dioryctria simplicella Heinemann, 1865 has been debated for over a century, with the loss of its type specimens exacerbating doubts regarding its species validity and identity. In 1901, Rebel [22] removed D. simplicella from genus Dioryctria and placed the species in synonymy of Salebriopsis albicilla (Herrich-Schäffer, 1849). In 1968, Roesler [23] reinstated its status as a valid species and name within the genus Dioryctria. In 1980, Petersen and Gaedike [24] regarded both D. simplicella and D. mutatella (Fuchs, 1899) [25] as synonymous, but they expressed reservations about the species identity Heinemann originally described. The advent of DNA barcoding using the cytochrome c oxidase subunit I (COI) gene [26,27,28] provided new tools for resolving this taxonomic uncertainty. Through integrated morphological and mitochondrial COI sequence analysis, Knölke [16] conclusively demonstrated that D. simplicella as it is universally understood represents a melanistic form of D. mutatella, thereby confirming their synonymy.
This study reports the first country record of Dioryctria simplicella in China. Its taxonomic status was confirmed through integrated evidence from classical morphological identification and mitochondrial COI gene sequence analysis. We carefully compared diagnostic characteristics of adult external morphology and genitalia, larvae, pupae, and eggs. Molecular identification involved sequencing the COI gene of D. simplicella. Based on genetic distance thresholds and phylogenetic tree topology, its status as a distinct species was validated. D. simplicella was historically subject to taxonomic controversy with D. mutatella due to the loss of type specimens and melanistic phenotypic variation. This study identified stable diagnostic traits in Chinese populations, supporting it as a separate species, and its discovery provides a critical addition to the East Asian fauna.

2. Materials and Methods

2.1. Specimens Collection

The specimens were collected from a 20-year-old Pinus sylvestris var. mongolica Litv. forest at the Xinjiang Experimental Forest Farm (124.46° N, 48.04° E), Qiqihar City, Heilongjiang Province, China, between 20 May and 30 September 2024. Adult and larval stages collected in the field were transported to the laboratory, where eggs and pupae were obtained. Adult were pinned and stored in dry condition. Specimens (larvae, pupae, eggs) were preserved in 95% ethanol at −20 °C. Morphological examination and DNA barcoding were employed for identification. Voucher specimens are deposited in the Northeast Forestry University (NEFU) Entomological Specimen Room (Harbin, China) and are accessible upon request. Related laboratory work was conducted at NEFU.

2.2. Morphological Study

Following immersion in 10% NaOH solution for 12–24 h, adult abdomens were tissue cleared and genitalia were dissected. Stereomicroscopic imaging of genitalia and eggs employed a Canon A620 camera (Canon, Tokyo, Japan), whereas macrophotography of whole specimens utilized a Canon EOS R5 camera (Canon, Tokyo, Japan). All photographs and images were processed in Adobe Photoshop CS6 v.13.0. Morphological descriptions of adults followed the methodology of Kuang and Li [19], while morphological descriptions of larvae followed the method of Piao et al. [29].

2.3. Molecular and Phylogenetic Study

Molecular analysis was performed using the standard mtDNA COI barcoding region on two larvae and four adults (two males and two females). DNA extraction from adult legs and intact larvae was performed with the TIANamp Micro DNA Kit (TIANGEN, Beijing, China) following standardized procedures. PCR amplification of the mitochondrial cytochrome c oxidase subunit I (COI) gene utilized universal primers as follows: LCO1490: 5′-GGTCAACAAATCATAAAGATATTGG-3′, HCO2198: 5′-TAAACTTCAGGGTGACCAAAAAATCA-3′ [30]. The PCR reaction was performed in a total 25 µL volume: 0.5 µL DNA, 1 µL of each primer (100 ng/µL), 12.5 µL of Takara Ex Taq DNA polymerase (Takara Biomedical Technology, Beijing, China), and 10 µL nuclease-free water. PCR conditions were as follows: initial denaturation at 95 °C for 2 min, 35 cycles of denaturation at 95 °C for 30 s, annealing at 49 °C for 30 s, extension at 72 °C for 30 s, with a final extension at 72 °C for 5 min. All PCR products were sequenced by Sangon Biotech Company (Shanghai, China).
The six D. simplicella sequences, evaluated using Chromas V2.6.5, were compared with homologous sequences retrieved from the NCBI GenBank database; all sequences were submitted to GenBank, and their accession numbers were obtained (www.ncbi.nlm.nih.gov/GenBank accessed on 22 May 2025). Additionally, 32 Dioryctria COI sequences were downloaded from GenBank, and Pyralis lienigialis and Orybina regalis were used as outgroups. GenBank accession numbers for all sequences analyzed in this study are listed in Table 1. The sequence genetic distances were estimated by the Kimura 2 parameter model (K2P) implemented on MEGA v7.0. To determine the number of haplotypes using DnaSP v6.12, phylogenetic analyses were performed using PhyloSuite v1.2.3. Sequences were aligned using MAFFT v7.526, resulting in a final aligned dataset that included 40 COI sequences of 682 bp. The optimal nucleotide substitution model was determined using ModelFinder. The concatenated sequence matrix was subjected to phylogenetic analysis using both maximum likelihood (ML) and Bayesian inference (BI) methods. The ML analysis was conducted under the TIM + F + G4 nucleotide model with 1000 bootstrap replicates. The BI analysis was performed using Markov chain Monte Carlo (MCMC) sampling algorithm under the GTR + F + G4 nucleotide model and was run for 1,000,000 generations, sampling every 1000 generations, with four chains. The first 25% of trees were discarded as burn-in, and the remaining trees were used to calculate posterior probabilities (PP). The resulting phylogenetic trees were visualized and edited using FigTree v1.4.2.

3. Results

3.1. Subsection Species Descriptions

  • Dioryctria simplicella Heinemann
Dioryctria simplicella Heinemann, 1865: 148–149
Type locality: Germany, Frankfurt am Main
=Dior. [yctria] abietella var. mutatella Fuchs, 1903: 233
Type locality: Germany—Brandenburg
  • Material examined.
11 males, 16 females, three larvae, Qiqihar, Heilongjiang Province, China, 124.46° N, 48.04° E, 20.V-30VIIII.2024, leg. Niya Jia, laboratory, in P. sylvestris var. mongholica Litv. cones.
  • Diagnosis. The genital morphology provides reliable diagnostic characters for distinguishing D. simplicella. In male genitalia, valva relatively narrow, costa of valva extending distally into a beak-shaped apex bearing a minute spine.
  • Description.
Head (Figure 1A). Head black covered with greyish white scales. Antennae grayish brown, filiform, scape black dorsally, flagellum with several basal segments black, the remainders densely short dark brownish cilia. Ocellus glossy yellow. Labial palpus brown, mixed with grayish white; labial palpus 3rd segment ⅓ of length of 2nd segment.
Thorax (Figure 1A). Wingspan 22–30 mm. Thorax blackish mixed with brown white; patagium grayish brown. Forewing grayish brown, suffused with dark brown and grayish white, whitish brown at base and on costa; antemedial line brownish white, broad, sinuate slightly, nearly obsolete near the costa; discal stigma large, brownish white, elliptical; postmedial line brownish white, with two inward angles and one outward angle; terminal line blackish brown, diffused with brownish white along its inside; cilia grayish brown. Hindwing grayish brown; cilia brownish white.
Abdomen (Figure 1A). Abdomen dark brown, covered with grayish brown scales.
Male genitalia (Figure 1B,D,E). Uncus overall appearance egg-shaped; apex rounded; broad at base. Gnathos small, conical, about 1/4 length of uncus. Valva relatively narrow; costa of valva extending distally into a beak-shaped apex bearing a minute spine. Sacculus thin, about 1/3 length of valva. Clasper somewhat triangular. Juxta U-shaped, medially bearing a scutiform sclerite, lateral arms fingerlike, curved inward. Vincullum V-shaped. Aedeagus elongate-cylindrical, internally armed with short spines.
Female genitalia (Figure 1C). Anal papillae short and small, sparsely with setae. Anterior apophyses equal in length to the posterior apophyses, but slightly thicker and with a slightly swollen base. Ductus bursae long and straight, sclerotized, length greater than 4× width, with sclerotized longitudinal stripe in the central third. Corpus bursae membranous, oval. One spine clusters at junction of ductus bursae and corpus bursae, and two groups of spines in upper half of corpus bursae. Ductus seminalis from the 1/3 of ductus bursae.
Larva (Figure 2C–E). Late instar larvae approximately 16–25 mm long. Head reddish brown. Pronotum dark brown, featuring a median reddish-brown maculation. Body color pale reddish-brown, dorsal surface of abdomen bearing three parallel dark brown longitudinal stripes extending from the cephalic to the caudal end, cuticle smooth. Chaetotaxy as shown in Figure 2E.
Thorax Segment (T1–T3). T1: D1 closer to XD1 than to D2; XD2, SD1, and SD2 are approximately equilateral triangles; L group bisetose; SV1 and SV2 are present above the leg; T2 and T3: D1 and D2 on a pinaculum; L group trisetose, L1 and L2 are clustered on a single pinaculum; SV group unisetose; SD1 and SD2 are present on T2, a dark ring around seta SD1 and SD2.
Abdomen Segment (A1–A10). On A1–A8, D group bisetose; SD group unisetose; L group trisetose, L1 and L2 are clustered on a single pinaculum; SV group trisetose on A1–A6; SV group bisetose on A7 and A8; a dark ring around seta SD1 and the center is pale on A8. On A9, D group unisetose; SD group bisetose; L group trisetose and on a single pinaculum; SV group bisetose. On A10, D1 and D2, SD2 and SD1, all coalesced on a shared pinaculum; L and SV group quadrisetose; L2–L4 are clustered on one pinaculum; SV3 and SV4 share a pinaculum.
Egg (Figure 2A). The egg is elliptical in shape and approximately 0.8 mm in length, pale yellow when freshly laid, with scattered red spots gradually appearing on the surface as embryonic development progresses.
Pupa (Figure 2B). The obtect pupa is spindle-shaped, with a body length of 8–12 mm, yellowish-brown in color, turning dark brown prior to eclosion, with six caudal spines at the posterior end of the pupa.
Distribution. China (Heilongjiang, new record), Germany, Bulgarian, Belarus, England, Austria, Belgium, Denmark, Czechia Republic, Estonia, Finland, France, Hungary, Latvia, Lithuania, Luxembourg, Netherlands, Norway, Romania, Slovakia, Sweden, and Switzerland.
Ecology. D. simplicella has one generation per year in Heilongjiang, China. Adults emerge from the end of May to the middle of September.
Host (Figure 3). P. sylvestris var. mongholica Litv. (New to China), Pinus sylvestris [24,31], and Picea glauca [24].

3.2. Molecular and Phylogenetic Analyses

We successfully obtained and sequenced 682 bp COI fragments from six D. simplicella specimens (two larval, two male adults, and two female adults). The final dataset comprised 40 COI sequences representing 20 species across three genera within the Pyralidae family (Table 1). No intraspecific variation (genetic distance = 0) was observed among the 10 D. simplicella sequences analyzed in this study. The smallest interspecific distance involving D. simplicella was found with D. mendacella (0.021). Genetic distances between D. simplicella and other Dioryctria species ranged from 0.021 to 0.109, with a mean distance of 0.068. The detailed pairwise genetic distance matrix is presented in the Supplementary Materials: Table S1. Haplotype analysis conducted in DnaSP v6.12 revealed a single haplotype among the 10 D. simplicella sequences. Both maximum likelihood (ML) (Figure 4) and Bayesian inference (BI) (Supplementary Materials: Figure S1) analyses of the COI dataset yielded congruent topologies, strongly supporting the monophyly of D. simplicella. The phylogenetic trees placed D. simplicella within the abietella-group. Consistent with the 2% threshold proposed by Hebert et al. [26] for species delimitation using COI barcode sequences, the observed genetic distances support the distinct status of D. simplicella. Overall, the results of the phylogenetic analysis corroborate the morphological identification of D. simplicella.

4. Discussion

This study reports the discovery of D. simplicella in China. We found D. simplicella larvae in the trunks and green cones of P. sylvestris var. mongholica Litv., captured adults through light trapping in the field, and obtained eggs and pupae under laboratory conditions. Both adult and larval specimens were identified to the species level through morphological and molecular biological methods, confirming them as D. simplicella.
Accurate identification of D. simplicella requires comprehensive consideration of geographical distribution and ecological information. According to reports, adult D. simplicella exhibit two morphs: the grey and the black. The black morph can be rapidly distinguished by its black forewing coloration [23,32], while the grey morph more closely resembles the typical wing pattern of most Dioryctria species [10,24]. The D. simplicella specimens discovered in China belong to the grey morph. Recent reports also confirm the grey morph in Belarus [33]. D. simplicella is widely distributed across Europe, where the occurrence frequency of the grey is significantly higher than that of the black. Initially, Hassler and Speidel [34] considered the black D. simplicella endemic to Germany; however, subsequent research revealed its distribution extends far beyond southwestern Germany and is widespread across Europe. In British populations, the black can comprise up to 25% of individuals [35]. Molecular evidence supports that the grey and black morphs of D. simplicella represent a single species. These morphs do not form distinct clusters corresponding to their phenotypes but instead jointly form a terminal polytomy, characteristic of a typical monophyletic group [16]. In the present study, D. simplicella specimens from China clustered closely with European D. simplicella from NCBI on the phylogenetic tree, supporting the conspecificity of Eurasian populations as D. simplicella. The D. simplicella larvae primarily feed on green cones, shoots, and buds of P. sylvestris and P. glauca [24,31]. The population of D. simplicella in China damaged not only these plant parts but also the main trunk, inducing substantial resin accumulation on trunks (Figure 3D).
The present study extends the geographic distribution of D. simplicella to East Asia and provides new data supporting taxonomic and phylogenetic research on the subfamily Phycitinae. This discovery enriches the diversity records of Chinese Lepidoptera. Our surveys confirm the extensive distribution of D. simplicella across P. sylvestris var. mongolica stands in Northeast China. This species poses a latent threat to pine health in Northeast China. Our study establishes fundamental data essential for the monitoring, management, and further taxonomic studies of D. simplicella in China.

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/insects16070664/s1, Figure S1: Bayesian inference (BI) phylogenetic trees constructed based on mtDNA COI; Table S1: Intraspecific and interspecific genetic distances of Dioryctria species in this study.

Author Contributions

Conceptualization, N.J., X.D. and J.Y.; methodology, N.J.; software, N.J.; validation, D.C. and J.Y.; formal analysis, N.J., X.D. and D.X.; investigation, N.J., X.D., D.X. and H.C.; resources, N.J., H.C., D.X., X.D. and D.X.; data curation, N.J. and X.D.; writing—original draft preparation, N.J.; writing—review and editing, D.C.; visualization, N.J. and X.D.; supervision, D.C.; project administration, J.Y.; funding acquisition, D.C. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the National Key R&D Program of China [2022YFD1401000] and the Key R&D Program of Heilongjiang Province [2023ZX02B05].

Data Availability Statement

The original contributions presented in this study are included in the article/Supplementary Materials. Further inquiries can be directed to the corresponding authors.

Acknowledgments

We greatly appreciate the Xinjiang Experimental Forest Farm, which supported us with the field investigation work.

Conflicts of Interest

The authors declare no conflicts of interest.

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Insects 16 00664 g001
Figure 1. Adult and genitalia of D. simplicella. (A) adult; (B) male genitalia; (C) female genitalia; (D) male culcita; (E) male aedeagus. Scale bars: 5 mm (A); 1 mm (B–E).
Figure 1. Adult and genitalia of D. simplicella. (A) adult; (B) male genitalia; (C) female genitalia; (D) male culcita; (E) male aedeagus. Scale bars: 5 mm (A); 1 mm (B–E).
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Figure 2. Egg, pupa and larva of D. simplicella. (A) egg; (B) pupa; (C,D) larva; (E) chaetotaxy. Scale bars: 0.1 mm (A); 3 mm (B–D).
Figure 2. Egg, pupa and larva of D. simplicella. (A) egg; (B) pupa; (C,D) larva; (E) chaetotaxy. Scale bars: 0.1 mm (A); 3 mm (B–D).
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Figure 3. P. sylvestris var. mongholica Litv. cone and trunk damaged by D. simplicella larvae. (A) external view of an infested cone; (B) larva inside the cone; (C) cone damaged by a mature larva in October; (D) infested trunk.
Figure 3. P. sylvestris var. mongholica Litv. cone and trunk damaged by D. simplicella larvae. (A) external view of an infested cone; (B) larva inside the cone; (C) cone damaged by a mature larva in October; (D) infested trunk.
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Figure 4. Maximum likelihood (ML) phylogenetic trees constructed based on mtDNA COI.
Figure 4. Maximum likelihood (ML) phylogenetic trees constructed based on mtDNA COI.
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Table 1. Samples of COI genes used in this study.
Table 1. Samples of COI genes used in this study.
Species GroupSpeciesGenBank NO.
abietella-groupDioryctria simplicellaPV668953.1
Dioryctria simplicellaPV668954.1
Dioryctria simplicellaPV668955.1
Dioryctria simplicellaPV668956.1
Dioryctria simplicellaPV668957.1
Dioryctria simplicellaPV668958.1
Dioryctria simplicellaHM875123.1
Dioryctria simplicellaHM872148.1
Dioryctria simplicellaHM872031.1
Dioryctria simplicellaGU707355.1
Dioryctria abietellaGU706362.1
Dioryctria abietellaKX048878.1
Dioryctria abietivorellaMG468430.1
Dioryctria abietivorellaJF852743.1
Dioryctria mendacellaOR449782.1
Dioryctria mendacellaOQ563613.1
Dioryctria pineaeHQ563610.1
zimmermani-groupDioryctria amatellaGU801522.1
Dioryctria banksiellaHM869672.1
Dioryctria banksiellaKT147303.1
auranticella-groupDioryctria auranticellaHM868369.1
Dioryctria auranticellaKF492320.1
Dioryctria disclusaKT143314.1
Dioryctria pryeriMF052103.1
Dioryctria pryeriMF049781.1
Dioryctria yiaiMF055629.1
Dioryctria yiaiMF055620.1
ponderosae-groupDioryctria okanaganellaHM867929.1
baumhoferi-groupDioryctria pentictonellaHM426749.1
Dioryctria subtractaHM406370.1
schuetzeella-groupDioryctria pseudotsugellaKM554558.1
Dioryctria pseudotsugellaKM554272.1
Dioryctria reniculelloidesKM553829.1
Dioryctria reniculelloidesKM548677.1
Dioryctria schuetzeellaHM874495.1
Dioryctria schuetzeellaHM871547.1
Dioryctria sylvestrellaHM875194.1
Dioryctria sylvestrellaKC135937.1
outgroupPyralis lienigialisHM874786.1
outgroupOrybina regalisPQ298688.1
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MDPI and ACS Style

Jia, N.; Ding, X.; Xie, D.; Chen, H.; Chi, D.; Yu, J. First Record of Dioryctria simplicella (Lepidoptera: Pyralidae) in China: Morphology, Molecular Identification, and Phylogenetic Position. Insects 2025, 16, 664. https://doi.org/10.3390/insects16070664

AMA Style

Jia N, Ding X, Xie D, Chen H, Chi D, Yu J. First Record of Dioryctria simplicella (Lepidoptera: Pyralidae) in China: Morphology, Molecular Identification, and Phylogenetic Position. Insects. 2025; 16(7):664. https://doi.org/10.3390/insects16070664

Chicago/Turabian Style

Jia, Niya, Xiyao Ding, Dan Xie, Huanwen Chen, Defu Chi, and Jia Yu. 2025. "First Record of Dioryctria simplicella (Lepidoptera: Pyralidae) in China: Morphology, Molecular Identification, and Phylogenetic Position" Insects 16, no. 7: 664. https://doi.org/10.3390/insects16070664

APA Style

Jia, N., Ding, X., Xie, D., Chen, H., Chi, D., & Yu, J. (2025). First Record of Dioryctria simplicella (Lepidoptera: Pyralidae) in China: Morphology, Molecular Identification, and Phylogenetic Position. Insects, 16(7), 664. https://doi.org/10.3390/insects16070664

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