Research Article |
Corresponding author: Cheng-Qi Zhu ( zhuchengqi@westlake.edu.cn ) Corresponding author: Ying Zhen ( zhenying@westlake.edu.cn ) Academic editor: Hume Douglas
© 2022 Cheng-Qi Zhu, Xiao-Dong Xu, Ying Zhen.
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:
Zhu C-Q, Xu X-D, Zhen Y (2022) Systematic review of the firefly genus Emeia Fu, Ballantyne & Lambkin, 2012 (Coleoptera, Lampyridae) from China. ZooKeys 1113: 153-166. https://doi.org/10.3897/zookeys.1113.79721
|
The Luciolinae genus Emeia Fu, Ballantyne & Lambkin, 2012 is reviewed. Phylogenetic relationships based on cox1 DNA barcoding sequences from 42 fireflies and 2 outgroup species are reconstructed. The dataset included three main Lampyridae subfamilies: Luciolinae, Photurinae and Lampyrinae, and Emeia was recovered within Luciolinae. A new species, Emeia pulchra Zhu & Zhen sp. nov., is described from the wetland of Lishui, Zhejiang, China. Emeia pulchra is sister species to E. pseudosauteri from Sichuan, which is supported by morphological characters and a phylogeny based on DNA barcoding sequences. The two species are separated geographically as shown on the distribution map. A key to species of Emeia using males is provided.
Cytochrome c oxidase subunit I, DNA barcoding, Emeia, firefly, Lampyridae
Emeia Fu, Ballantyne & Lambkin, 2012 (Luciolinae) was established as a monotypic genus (
In this study, based on specimens collected from Lishui, Zhejiang, China, we describe adults of Emeia pulchra Zhu & Zhen sp. nov. based on morphological and molecular data. We compare it with the previously described E. pseudosauteri. We also provide new information on the adult male hind wing venation of the type species E. pseudosauteri. With our detailed examination of both species, we present a systematic review of the genus Emeia and a key to species.
EL elytral length;
EW elytral width;
PL pronotal length;
BL body length (the sum of PL, EL and the length of the exposed portions of the head from the pronotum);
BW body width (the greatest distance across the elytra, BW=2EW);
T7, 8 abdominal tergite numbers;
V6, 7 abdominal ventrite numbers.
Adult males of Emeia pulchra Zhu & Zhen sp. nov. were collected from Jiulong National Wetland Park, Lishui, Zhejiang Province in April, 2020. The holotype and paratypes of the new species are stored at School of Life Sciences, Westlake University, Hangzhou, Zhejiang. Samples of both male and female Emeia pseudosauteri were collected from Mt. Tian Tai, Sichuan Province in April, 2021.
Habitus images were taken using a Nikon D7500 camera. Images of genitalia were taken using a Nikon D7500 camera mounted on an SZ650 microscope (Chongqing Optec Instrument Co., Ltd.) under reflection or transmission light. Images were edited using Adobe Photoshop CS6. Morphological terminology and measurements follow those decribed in
We sequenced the cox1 gene barcode fragment from Emeia pseudosauteri and E. pulchra. Specifically, total DNA of the two Emeia species was isolated using the DNeasy Blood and Tissue Kit (Zhejiang Easy-Do Biotech CO., Ltd.), according to the manufacturer’s protocol. The primers LCO 1490 and HCO 2198 (
MEGA6 (
Genbank accession numbers for cox1 sequences used for the phylogenetic analysis.
Species | Family | Sub-family | GenBank id |
---|---|---|---|
Pyrocoelia pectoralis | Lampyridae | Lampyrinae | KP763467.1 |
Pyrocoelia rufa | Lampyridae | Lampyrinae | AF452048.1 |
Pyrocoelia abdominalis | Lampyridae | Lampyrinae | AB608766.1 |
Pyrocoelia atripennis | Lampyridae | Lampyrinae | AB608767.1 |
Pyrocoelia discicollis | Lampyridae | Lampyrinae | AB608768.1 |
Pyrocoelia fumosa | Lampyridae | Lampyrinae | AB608769.1 |
Pyrocoelia matsumurai | Lampyridae | Lampyrinae | AB608770.1 |
Diaphanes nubilus | Lampyridae | Lampyrinae | MG200080.1 |
Diaphanes pectinealis | Lampyridae | Lampyrinae | NC_044793.1 |
Photinus pyralis | Lampyridae | Lampyrinae | KY778696.1 |
Ellychnia corrusca | Lampyridae | Lampyrinae | KR483038.1 |
Ellychnia hatchi | Lampyridae | Lampyrinae | JF887410.1 |
Pyractomena lucifera | Lampyridae | Lampyrinae | MF640134.1 |
Pyractomena borealis | Lampyridae | Lampyrinae | HQ928227.1 |
Pyractomena angulata | Lampyridae | Lampyrinae | JN290381.1 |
Aspisoma sp. | Lampyridae | Lampyrinae | EU009322.1 |
Lucidina accensa | Lampyridae | Lampyrinae | AB608771.1 |
Lucidina kotbandia | Lampyridae | Lampyrinae | FJ462784.1 |
Lucidota atra | Lampyridae | Lampyrinae | HQ984304.1 |
Photuris pensylvanica | Lampyridae | Photurinae | MF634963.1 |
Photuris quadrifulgens | Lampyridae | Photurinae | HM433520.1 |
Bicellonycha lividipennis | Lampyridae | Photurinae | KJ922151.1 |
Bicellonycha wickershamorum | Lampyridae | Photurinae | EU009302.1 |
Pristolycus sp. | Lampyridae | Luciolinae | MK292099.1 |
Sclerotia flavida | Lampyridae | Luciolinae | KP763460.1 |
Sclerotia aquatilis | Lampyridae | Luciolinae | KP763466.1 |
Pygoluciola dunguna | Lampyridae | Luciolinae | MT106243.1 |
Pygoluciola qingyu | Lampyridae | Luciolinae | MK292093.1 |
Curtos bilineatus | Lampyridae | Luciolinae | NC_044789.1 |
Curtos costipennis | Lampyridae | Luciolinae | AB608764.1 |
Abscondita terminalis | Lampyridae | Luciolinae | NC_044776.1 |
Abscondita anceyi | Lampyridae | Luciolinae | NC_039706.1 |
Emeia pseudosauteri 1 | Lampyridae | Luciolinae | MN722654.1 |
Emeia pseudosauteri 2 | Lampyridae | Luciolinae | OK103803 |
Emeia pulchra | Lampyridae | Luciolinae | OK144132 |
Luciola italica | Lampyridae | Luciolinae | KM448530.1 |
Asymmetricata circumdata | Lampyridae | Luciolinae | NC_032062.1 |
Drilaster axillaris | Lampyridae | Ototretinae | AB608756.1 |
Drilaster okinawensis | Lampyridae | Ototretinae | AB608758.1 |
Stenocladius yoshikawai | Lampyridae | Ototretinae | AB608759.1 |
Lamprigera yunnana | Lampyridae | incertae_sedis | MG200082.1 |
Cyphonocerus marginatus | Lampyridae | Cyphonocerinae | AB608754.1 |
Rhagophthalmus lufengensis | Rhagophthalmidae | – | DQ888607.1 |
Rhagophthalmus ohbai | Rhagophthalmidae | – | AB608775.1 |
The cox1 barcode sequences of E. pseudosauteri and E. pulchra share a 94% sequence identity over the 658 bp segment. The phylogeny constructed from cox1 of fireflies showed three main clades corresponding to Lampyrinae, Photurinae and Luciolinae (Fig.
Maximum likelihood cox1 gene tree of Emeia and related genera. The star highlights the new species, E. pulchra Zhu & Zhen sp. nov. Emeia pseudosauteri_1 was downloaded from GenBank (MN722654.1). Emeia pseudosauteri_2 was sequenced during this study. Bootstrap values greater than 0.7 from 1000 replicates are shown.
Emeia pseudosauteri Geisthardt, 2004 (designated by Fu, Ballantyne and Lambkin 2012).
(based on adult male). Emeia belongs to a group of Luciolinae in which the males have aedeagal parameres widely visible beside the phallus (
(based on adult male). Body length 6.5–10.5 mm. Body width 2.7–4.0 mm. Integument black or dark brown, with a narrow (e.g., in E. pulchra, see Fig.
Head. Hypognathous; head depressed between eyes, eyes exposed in front of pronotum; antennae filiform, with 11 antennomeres (Figs
Thorax. Pronotum in dorsal view appearing pink-red or orange-red, with a black median stripe, lateral margins almost parallel (Figs
Abdomen. V2–V5 dark brown or black. Light organs present in V6 and V7, entirely occupying V6; V7 semitransparent (Figs
Male genitalia. Trilobate, parameres extending ~0.14 mm (n = 3) beyond phallus; both parameres equal in length (Figs
(based on adult male). The new species can be differentiated from E. pseudosauteri Fu, Ballantyne & Lambkinby the elytron, hindwing venation and aedeagus. In fresh specimens, the elytral apices are black in E. pulchra (Fig.
Male: BL 10.0–10.4 mm; BW 3.5–3.7 mm (three individuals).
Head. Antennae filiform, black, almost 2/3 as long as body length; antennomere 1 cone-shaped; 2 short and cylindrical; 3 to 10 compressed, not bifurcate; 11th antennomere almost 1.5 times longer than 10th, slightly dilated from base to apex. Concave between eyes dorsally in cross section, both eyes occupying about 2/3 width of whole head in ventral view. Eyes spherical, so that head cannot fully contract into pronotum. Mouthparts fully developed, clypeolabral suture flexible, outer edges of labrum reaching inner edges of closed mandibles.
Thorax. Scutellum black and slightly emarginate distally. Elytra elongated, dark brown to black, apices not deflexed in dorsal view, sides slightly convex. Hind wing well developed, r3 half the length of r4 (Fig.
Abdomen. Dark brown, ventrites gradually diminishing in length posterad. Light organs yellow-white, occupying almost all of V6 and half of V7, not reaching to posterior edges of V7. V6 and V7 rounded laterally (Fig.
Male genitalia
(Fig.
The specific name pulchra refers to the bright pronotum coloration.
China • 1♂; Zhejiang, Lishui; 28°37.56'N, 119°49.7'E; H: 60 m, 2. IV. 2020; Chengqi Zhu leg.; ‘HOLOTYPE (red), ♂, Emeia pulchra sp. nov., det. Zhu, Zhen, 2021’ (Westlake University).
China • 1♂; Zhejiang, Lishui; 28°37.56'N, 119°49.7'E; H: 60 m, 2. IV. 2020; Chengqi Zhu leg.; ‘PARATYPE (yellow), ♂, Emeia pulchra sp. nov., det. Zhu, Zhen, 2021’ (Westlake University).
China: Zhejiang Province.
The males were found in an open forest of mainly Chinese wingnut, of the family Juglandaceae [Pterocarya stenoptera C. DC.] (Fig.
There are many terrestrial snails and slugs in this habitat, which may be potential food for Emeia pulchra larvae. Combining descriptions from local people and our field observations, adult fireflies are usually observed mid-March. The protection of fireflies has been supported by the Lishui government and Jiulong National Wetland Park management departments, and this area has been protected as Jiulong National Wetland Park (Fig.
There are two obvious luminous bands at the terminal end of the adult male abdomen. The two bands both emit intermittent bright light during courtship. The male courtship behavior usually starts at 19:00 (approximately 1h after sunset), and peaks at about 20:30. Adult males rest on higher herbs and emit yellow and green flashing light. Males are reluctant flyers; the distance of each flight ranges from 0.5 to 5 m.
Emeia pseudosauteri Fu, Ballantyne & Lambkin, 2012. Male and female 8 habitus of male A dorsal view. Arrow highlights narrow orange stripe on elytral apices. The color appears darker in this photo, but it is orange and easily seen in both dried and fresh samples B ventral view 9 habitus of female. A dorsal view B ventral view 10 right wing of male. Dorsal view. Arrow points to wing venation, which differs between the two Emeia species 11 right wing of female. Dorsal view. Scale bars: 5 mm (8, 9); 2 mm (10); 0.5 mm (11).
Emeia pseudosauteri
(
China: 6♂♂,1♀, Sichuan, Mt. Tian Tai, 3.IV. 2021, Chengquan Cao leg. We herein examined specimens of E. pseudosauteri from Mt. Tian Tai (the type locality), and their identity was further verified using cox1 barcode sequences (Fig.
Distribution map of the genus Emeia in China. The black star indicates E. pulchra Zhu & Chen sp. nov., the black dot E. pseudosauteri (map of China from: http://bzdt.ch.mnr.gov.cn/).
1 | The elytral apices have a narrow orange stripe in both fresh and dried specimens; upper vein of MP3+4 forked and reaching edge of hind wing (Fig. |
E. pseudosauteri Fu, Ballantyne & Lambkin |
– | The elytral apices are black in fresh and preserved specimens (Fig. |
E. pulchra Zhu & Zhen, sp. nov. |
In this study, we summarized the diagnostic features of the genus Emeia. Emeia pulchra Zhu & Zhen, sp. nov. is morphologically similar to E. pseudosauteri Fu, Ballantyne & Lambkin, 2012 from Sichuan Province. However, we found differences in the antennal length and body size between the two species. The body size of a species may vary due to nutrition and environmental factors, so we did not include size in the diagnosis to the new species. The antenna of male E. pulchra (Fig.
The “barcode region” of cox1 is often used as an aid to new species’ identification and distinction from close relatives in the Barcode of Life Data system (
We are obliged to Ms. Dan-Dan Tu for assistance with DNA amplification and Mr. Zhuo-Heng Jiang for helpful comments on our manuscript. We thank Dr. Cheng-Quan Cao and Dr. Fang-Zhou Ma for specimen collection and sampling information. This work was supported by the National Natural Science Foundation of China [grant number 31900315 to Y.Z.], the Zhejiang Provincial Natural Science Foundation of China [grant numbers LR21C030001 to Y.Z.] and the Westlake Education Foundation.