Research Article |
Corresponding author: Kôichi Arimoto ( kou.arimoto@gmail.com ) Academic editor: Hume Douglas
© 2017 Kôichi Arimoto, Hisayuki Arimoto.
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:
Arimoto K, Arimoto H (2017) The genus Alaolacon Candèze, a senior synonym of the genus Eumoeus Candèze (Coleoptera, Elateridae, Agrypninae). ZooKeys 656: 85-110. https://doi.org/10.3897/zookeys.656.8914
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Alaolacon Candèze, 1865 is found to be a senior synonym of Eumoeus Candèze, 1874, Luzonicus Fleutiaux, 1916 and Tharopsides Fleutiaux, 1918. Alaolacon is represented by A. bakeri (Fleutiaux, 1916), comb. n., A. candezei Fleutiaux, 1928, A. cyanipennis Candèze, 1865, A. fujiokai sp. n., A. griseus Candèze, 1874, A. megalopus sp. n., A. murrayi (Candèze, 1874), comb. n., and A. philippinensis nom. n. This genus is redescribed based on the descriptions of three species, A. candezei, A. fujiokai, and A. megalopus as well as the examination of the holotypes of A. cyanipennis and A. murrayi comb. n. Males of the genus Alaolacon exhibit 12-segmented and biflabellate antennae, and the females exhibit 11-segmented and subpectinate antennae. A key to species is provided.
Eumaeus , Hemirhipini , Luzonicus , new species, Oriental region, replacement name, taxonomy, Tharopsides
Depositories of the type specimens and non-type specimens examined are as follows:
BMNH
The
ELKU
Entomological Laboratory,
A generic description of Alaolacon was made from the study of the type specimens of A. cyanipennis Candèze, 1865, A. candezei Fleutiaux, 1928, A. murrayi (Candèze, 1874) comb. n. (= Eumoeus murrayi) and two new species described here. Species descriptions of A. cyanipennis and A. murrayi are not provided as they are adequately described in
We could not find the types of two species, A. bakeri (Fleutiaux, 1916), comb. n. (= Luzonicus bakeri) and A. philippinensis comb. n. (= Tharopsides bakeri Fleutiaux, 1918) in the collections of BMNH,
Photographs of specimens were taken by a single-lens reflex camera (Canon EOS 70D) with a macro lens (Canon macro photo lens MP-E 65mm), and then images taken in a series of focal planes were combined using CombineZM 1.0.0 software (Alan Hadley, United Kingdom). Micrographs were prepared using a scanning electron microscope (SEM: Hitachi S-3000N) without gold coating.
Most structures were observed under a stereo microscope (Olympus-SZX9). Measurements are in millimeters and were made with a micro ruler (MR-2, Kenis Limited, Ôsaka; minimum scale value: 0.05 mm). Specimens were softened in warm water. The pregenital segments and genitalia extracted from the abdomen were soaked in 10% KOH solution (room temperature, male: 2 hours, female: 48 hours). The parts were cleaned in 30% ethanol (10 min) and dehydrated in 99.5% ethanol (5 min) and then mounted in glycerin on microscope slides, except the female genitalia, which were examined in water and then mounted in glycerin. A transmission microscope (Nikon Y-IDT) with a camera lucida was used to examine slides and for drawing. Morphological terminology follows
The following abbreviations are used:
BL body length from head to elytral apices
BW the maximum body width
MIE the minimum distance between the eyes
MAE the maximum distance across the eyes
OI Ocular index: MIE/MAE × 100
PL the maximum pronotum length including posterior angles
PML length of the midline of pronotum
PW the maximum pronotum width including posterior angles
PI Pronotam index: PL/PW × 100
EL the maximum elytra length
EW the maximum elytra width
EI Elytra index: EL/EW × 100
Alaolacon
Candèze, 1865: 13 (original description; type species: Alaolacon cyanipennis Candèze, 1865; by monotypy; in Mélanactides);
Eumoeus
Candèze, 1874: 113 (original description; type species: Eumoeus murrayi Candèze, 1874; by monotypy; in tribe Alaites), 214 (as “Eumaeus”; index);
Luzonicus
Fleutiaux, 1916: 232 (original description; type species: Luzonicus bakeri Fleutiaux, 1916; by monotypy; in Corymbitinae);
Tharopsides
Fleutiaux, 1918: 235 (original description; type species: Tharopsides harmandi Fleutiaux, 1918);
Setae flat, wider at midlength than base, with longitudinal micro carinae (Figs
Adult. Body (Figs
Head (Figs
Alaolacon candezei Fleutiaux, 1928, female, holotype. 4 head and pronotum, anterolateral view 5 right antenna, anterior side (dotted line: apical half part of antennomere XI thinner than its basal half part) 6 mouth-parts 7 prosternal process, lateral view 8 anterior angle of hypomeron (arrow: mesal edge carinate anterolaterad) 9 posterior part of hypomeron and mesothorax, ventral view (arrows: posterior margin with two angles) 10 scutellum 11 right hind wing (arrow: cross vein between veins MP4 and CuA2 located at contact point between veins MP3 and MP4).
Prothorax shorter to longer than wide, widest posteriorly or at mid-length. Pronotum with anterior angle bisinuate (Figs
Mesothorax. Scutellum longer than wide; anterior margin straight, well defined by wrinkled band; sides concave or straight, widest posteriorly (Figs
Abdomen. Male. Terigite VIII shorter than wide (Fig.
Genitalia. Male. Aedeagus (Figs
Unknown.
Oriental Region: India, Thailand, Vietnam, Indonesia (Sumatra, Java), Malaysia (Peninsular Malaysia, Borneo), the Philippines (Mindanao Is., Luzon Is.).
Nothing is known of the life history and ecology.
Alaolacon candezei Fleutiaux, 1928: 177 (original description; type locality: Malaysia, East Malaysia (Sabah), Banggi Island).
Holotype. Female, Banggi Island (located off the northern coast of Borneo), Sabah, Malaysia, Waterstradt leg. [
Body black, elytra blue and with metallic luster, legs red-black; setae white; frontal depression moderate; eye small; female antennomere III subpectinate, 1.2 × as long as wide; prothorax almost as long as wide, widest posteriorly; pronotum with anterior angles bisinuate, median longitudinal depression shallow, not reaching anterior margin or base, punctate; anterior angles of hypomeron rounded; prosternal spine inclined weakly behind procoxae; scutellum concave laterally, widest near posterior 2/5; hind wings without wedge cell, with cross vein between veins MP4 and CuA2 located at contact point between veins MP3 and MP4; female sternite VIII with apex concave.
BL: 24.0, BW: 8.35, MIE: 2.56, MAE: 3.47, OI: 74, PL: 7.64, PML: 6.67; PW: 7.70, PI: 99, EL: 15.7, EW: 8.35, EI: 188.
Body (Figs
Head. Frontal depression moderate (Fig.
Prothorax almost as long as wide, widest posteriorly; hind angles straight posteriorly. Pronotum with anterior angle bisinuate; median longitudinal depression shallow, not reaching anterior margin or base, punctate. Hypomeron with anterior angles rounded (Fig.
Abdomen. Ventrite V 0.59 × as long as wide. Tergite VIII (Fig.
Genitalia (Fig.
Unknown.
Malaysia: Sabah: Banggi Island.
This species is similar to Alaolacon cyanipennis Candèze, 1865 in large body size (24.0 mm), black body and elytra with metallic luster, but is distinguished by the following contrasting characters (A. cyanipennis in parentheses): female antennomere III pectinate (Fig.
This species are known only from the female holotype. We predict that the males also exhibit blue elytra and with metallic luster, scutellum widest near posterior 2/5, hind wings without wedge cell and with cross vein between veins MP4 and CuA2 located at contact point between veins MP3 and MP4.
Alaolacon
cyanipennis
Candèze, 1865: 13 (original description: type locality: Peninsular Malaysia);
Lectotype. Female, Malacca, West Malaysia (Peninsular Malaysia), Malaysia, Janson coll. [BMNH] (Fig.
Body black, elytra blue-black and with metallic luster; setae white; female antennomere III trapezoidal, 1.4 × as long as wide; prothorax as long as wide, widest at mid-length except for posterior angles; pronotal anterior angles bisinuate and rounded; anterior angles of hypomeron rounded; prosternal spine inclined weakly behind procoxae; scutellum concave laterally, widest near posterior 1/3; hind wings with wedge cell, with cross vein between veins MP4 and CuA2 located at contact point between veins MP3 and MP4; female sternite VIII with rounded apex.
See
Malaysia: the Peninsular Malaysia. Indonesia: Sumatra.
We could not locate other syntypes including at
Only female specimens are known (
The name of this species honors Mr. Masahiro Fujioka for providing the material.
Holotype. Male, Tawau, East Malaysia (Sabah), Malaysia, V 2014 [ELKU].
Body black, elytra blue and with metallic luster, legs black; setae black on dorsal side and white on ventral side; frontal depression deep; eye small; prothorax almost as long as wide, widest posteriorly; pronotum with anterior angles bisinuate and rounded, medina longitudinal depression deep, extending from before pronotal anterior margin to base, punctate; prosternal process inclined strongly behind procoxae; anterior angles of hypomeron acute; scutellum concave laterally, widest near posterior 1/3; hind wings with cross vein between veins MP4 and CuA2 located anterad to contact point between veins MP3 and MP4, without wedge cell; median lobe of male aedeagus stout.
BL: 18.9, BW: 6.11, MIE: 2.08, MAE: 3.05, OI: 68, PL: 5.95, PML: 5.23; PW: 5.91, PI: 101, EL: 12.1, EW: 6.11, EI: 197.
Body (Figs
Head (Fig.
Alaolacon fujiokai sp. n., male, holotype. 22 head and pronotum, anterolateral view 23 right antenna, dorsal view 24 mouth-parts 25 prosternal process, lateral view 26 anterior angle of hypomeron (arrow: mesal edge carinate anterolaterad) 27 posterior part of hypomeron and mesothorax, ventral view (arrows: posterior margin with two angles) 28 scutellum 29 right hind wing (arrow: cross vein between veins MP4 and CuA2 located anterad to contact point between veins MP3 and MP4).
Prothorax almost as long as wide, widest posteriorly; sides rounded anteriorly, liner posteriorly. Pronotum with anterior angles bisinuate and rounded; median longitudinal depression deep, extending from before pronotal anterior margin to base, punctate. Prosternal spine inclined strongly (at 18 degrees) behind procoxae (Fig.
Abdomen. Ventrite V 0.67 × times as long as wide. Tergite VIII (Fig.
Unknown.
Malaysia: Sabah: Tawau.
This species is distinct by black body, blue elytra with metallic luster, black setae on dorsal side, white setae on ventral side and strong antennomeres III-XI flabellation. It is similar to Alaolacon candezei Fleutiaux, 1928 in having a black body, blue elytra with metallic luster, pronotum anterior angles bisinuate and rounded, and scutellum concave laterally, except for drastic sexual differences of antennae, but is distinguished by the following contrasting characters (A. candezei in parentheses): legs black (Fig.
Alaolacon fujiokai and A. candezei are similar species from the same island, but we recognized they are different species by the setal color and the hind wing venation. We believe that setal complementary color difference probably is not caused by sexual dimorphism because such dimorphism has not previously been observed in species of the Agrypninae. We also believe that differences in hind wing venation are unlikely to be caused by sexual dimorphism because such dimorphism has not previously been observed in species with flying females.
Eumoeus
murrayi
Candèze, 1874;
A combination of the Greek megalos, meaning great, and the Greek ops, meaning eye, refer to the large compound eyes.
Holotype. Male, Java, Indonesia [
Body brown, without metallic luster; setae yellow-brown; frontal depression moderate; eye very large; prothorax wider than long, widest posteriorly; pronotal anterior angles rounded; median longitudinal depression shallow, located at anterior half, punctate; anterior angles of hypomeron acute; prosternal spine inclined strongly behind procoxae; scutellum 1.5 × as long as wide; sides of scutellum parallel; hind wings with cross vein between veins MP4 and CuA2 located just anterior to contact point between veins MP3 and MP4, without wedge cell; male tergite VIII longer than wide; median lobe of male aedeagus elongate.
BL: 11.8, BW: 3.54, MIE: 1.02, MAE: 2.31, OI: 44, PL: 3.11, PML: 2.74, PW: 3.26, PI: 95, EL: 7.70, EW: 3.54, EI: 218.
Body (Figs
Head (Fig.
Prothorax wider than long; sides widest posteriorly, rounded anteriorly, liner posteriorly. Pronotum convex; anterior angles rounded; median longitudinal depression shallow, located at anterior half, punctate (Fig.
Alaolacon megalopus sp. n., male, holotype. 41 head and pronotum, anterolateral view 42 right antenna, dorsal view 43 mouth-parts 44 prosternal process, lateral view 45 anterior angle of hypomeron (arrow: mesal edge carinate anterolaterad) 46 posterior part of hypomeron and mesothorax, ventral view (arrows: posterior margin with two angles) 47 scutellum 48 right hind wing.
Abdomen. Ventrite V 0.65 × as long as wide. Tergite VIII (Fig.
Unknown.
Indonesia: Java.
The holotype is damaged with most appendages lost. The holotype of this species was identified as Eumoeus murrayi (= Alaolacon murrayi comb. n.) by Candèze (Fleutiaux, 1928), but separated from A. murrayi by the following characteristics (the holotype of A. murrayi in parentheses): eye very large (OI: 44) (eye large, OI: 50); anterior angles of pronotum rounded (Fig.
Only this species exhibits parallel sides of scutellum in this genus. The scutellum shape could be a useful specific diagnostic feature for this species including its female.
Eumoeus
murrayi
Candèze, 1874: 113 (original description on male; type locality: Madras, India), 214 (as “Eumaeus murrayi”; index);
Tharopsides
harmandi
Fleutiaux, 1918: 235 (original description on male; type locality: Bangkok, Thailand);
Luzonicus
murrayi
(Candèze, 1874):
Holotype. Male, Madras, India, Murray leg. [
Body red-brown, without metallic luster; setae yellow-brown; frontal depression deep; eye large; prothorax shorter than wide, widest posteriorly; pronotum with anterior angles bisinuate, median longitudinal depression not reaching anterior margin or base and impunctate at posterior half; anterior angles of hypomeron acute; prosternal spine inclined strongly behind procoxae; scutellum 1.3 × as long as wide, with sides straight, widest posteriorly; hind wings without cross vein between veins MP4 and CuA2 and wedge cell; male tergite VIII shorter than wide; median lobe of male aedeagus elongate.
BL: 14.9, BW: 4.85, MIE: 1.43, MAE: 2.85, OI: 50, PL: 4.34, PML: 3.68, PW: 4.64, PI: 94, EL: 10.1, EW: 4.85, EI: 208.
See
India. Thailand. Vietnam.
This species is only known from the male.
Eumoeus and Tharopsides were described from males exhibiting 12-segmented and biflabellate antennae, whereas Luzonicus were described from female exhibiting 11-segmented and filiform to subpectinate antennae.
Alaolacon Candèze, 1865 was only known from female with 11-segmented and pectinate antennae. We determined that a male specimen (the holotype of A. fujiokai sp. n.), in possessing biflabellate antennae, belongs to Alaolacon because of the similarity to Alaolacon cyanipennis and Alaolacon candezei including: black body, blue elytra with metallic luster, pronotum anterior angles bisinuate, scutellum concave laterally. This association demonstrates that Alaolacon also has sexually dimorphic antennae.
In the tribe Hemirhipini, only four genera, Alaolacon, Eumoeus, Mocquerysia Fleutiaux, 1899 and Eleuphemus Hyslop, 1921 have strongly sexually dimorphic antennae. Their males exhibit 12-segmented and biflabellate antenna, and females exhibit 11-segmented and subpectinate antennae. Eleuphemus is separated from Alaolacon, Eumoeus, Mocquerysia (the latter three genera in parentheses) by the supra-antennal carinae continuous across frons (supra-antennal carina not continuous across frons) and mandible without subapical tooth (mandible with subapical tooth). Mocquerysia is separated from Alaolacon and Eumoeus (the latter two genera in parentheses), prosternal suture shortly grooved (prosternal suture not grooved), scutellum narrowed apically and with straight side (scutellum widest apically and concave laterally or with parallel sides in A. megalopus sp. n.), elytral intervals flat (elytral intervals convex).
Luzonicus bakeri Fleutiaux, 1916 and T. bakeri Fleutiaux, 1918 are eventual homonyms since Luzonicus and Tharopsides are junior synonyms of Alaolacon. We propose A. philippinensis nom. n., as the replacement name for A. bakeri (Fleutiaux, 1916) comb. n. Alaolacon currently contains eight species, 1, A. bakeri, 2, A. candezei, 3, A. cyanipennis, 4, A. fujiokai, 5, A. griseus Candèze, 1874, 6, A. megalopus, 7, A. murrayi and 8, A. philippinensis.
We could not find the types of two species, A. bakeri and A. philippinensis, and have not examined these species. Further effort to find the localities of the types of the two species are needed in order to understand the complete picture of these species.
1 | Head and pronotum brown to red-brown | 2 |
– | Head and pronotum black | 4 |
2 | Prothorax longer than wide, elytra red-brown but brown-black on posterior half | A. bakeri (Fleutiaux, 1916) |
– | Prothorax shorter than wide, elytra brown to red-brown | 3 |
3 | Scutellum widest posteriorly | A. murrayi (Candèze, 1874) comb. n. |
– | Scutellum with parallel sides | A. megalopus sp. n. |
4 | Elytra blue or blue-black, and with metallic luster | 5 |
– | Elytra black and without metallic luster | 7 |
5 | Setae black dorsally and white ventrally | A. fujiokai sp. n. |
– | All setae white | 6 |
6 | Prothorax widest posteriorly, wedge cell of hind wings absent | A. candezei Fleutiaux, 1928 |
– | Prothorax widest at mid-length except for posterior angles, wedge cell of hind wings present | A. cyanipennis Candèze, 1865 |
7 | Ventral surface red-brown | A. griseus Candèze, 1874 |
– | Ventral surface black | A. philippinensis nom. n. |
We appreciate the efforts of Dr. Roger Booth (Natural History Museum, London) and Mr. Antoine Mantilleri (Muséum national d’Histoire naturelle, Paris) for providing access to the facilities and type material, Mr. Masahiro Fujioka (Tokyo) for providing the valuable specimen, Mr. Katsumi Akita (Mie, Japan) for his sincere cooperation, and Drs. Keita Matsumoto and Michael Geiser (Natural History Museum, London) for providing images of A. cyanipennis. We greatly thank Dr. Hume Douglas (Agriculture and Agri-Food Canada, Ottawa) and an anonymous reviewer. Their comments and advices helped improve the manuscript. We are also grateful to Dr. Munetoshi Maruyama (Kyushu University Museum, Fukuoka, Japan) and Prof. Toshiya Hirowatari (Entomological laboratory, Faculty of Agriculture, Kyushu University, Fukuoka, Japan) for reading through an earlier draft of this paper. This is a contribution from the Entomological Laboratory, Kyushu University, Fukuoka (Ser.7, No. 50).