Research Article |
Corresponding author: Wan F. A. Jusoh ( wfawan@gmail.com ) Academic editor: Marianna Simões
© 2024 Wan F. A. Jusoh, Lesley Ballantyne.
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:
Jusoh WFA, Ballantyne L (2024) A catalogue and redescription of type specimens of fireflies (Coleoptera, Lampyridae, Luciolinae) deposited in Naturalis Biodiversity Center, Leiden. Contributions to Entomology 74(1): 63-80. https://doi.org/10.3897/contrib.entomol.74.e107520
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Type specimens of flashing fireflies (Coleoptera, Lampyridae, Luciolinae) in the Naturalis Biodiversity Center, Leiden (
Atyphella, Curtos, Indonesia, Luciola sensu stricto, name-bearing type, Philippines, Pteroptyx decolor, Southeast Asia, Sri Lanka
Firefly beetles, or Lampyridae, are diverse insects with more than 2,200 species (
Identifying species in the Luciolinae taxonomy is a difficult task for taxonomists due to inadequate descriptions and difficulties in locating types (
In late 2019, one of us (WFAJ) had the opportunity to visit the Naturalis Biodiversity Center in Leiden, the Netherlands and search collections to locate and digitally catalogue type specimens of Lampyridae. Naturalis Biodiversity Center (
In this study, we examined Luciolinae specimens explicitly marked or indicated as type material, collected from major islands in Indonesia (including parts of Borneo), Sri Lanka, and the Philippines, currently housed in
To ensure consistency, we compared the external morphology of the type specimen with the original descriptions. Each specimen and its labels was photographed using a high-resolution imaging system, and a museum barcode catalogue number was assigned. We also clarified the taxonomic status of each named species and designated lectotypes for selected species.
Images were captured using digital imaging systems.
Species are addressed in alphabetical order under a generic heading with full generic synonymic tables as given in
Type localities are cited in their original spelling with current interpretation of cited locations in Table
Interpretation by the authors, of the verbatim used to describe locality.
Verbatim location | Interpretation of locality (Country) |
---|---|
“Buru” | Buru is one of the islands within Maluku Islands (Indonesia) |
“L. Petak” | Long Petak in northern Kalimantan on island of Borneo (Indonesia) |
“SW Celebes” / “Paloppo” / “Todjamboe” | Southwest West Sulawesi / Palopo / Tojambu. Tojambu is a locality in Palopo in the province of South Sulawesi (Indonesia) |
“Ceylan” / “Talgaswella” | Sri Lanka / Talgaswella (Sri Lanka) |
“Bodjonegoro” | Bojonegoro is in East Java (Indonesia) |
“Koetoer” | Kutur (?) in Sumatra (Indonesia) |
“Rawas” | Rawas could possibly refer to Rawas area of the upper Musi River on the island of Sumatra (Indonesia) |
“Lebong” | Lebong in Bengkulu Province, Indonesia, on the island of Sumatra (Indonesia) |
“Palembang bovenland” | Palembang in South Sumatra (Indonesia) |
“Atjeh” | Aceh or Aceh Province on the northwest tip of Sumatra Island (Indonesia) |
“Borneo” / “occ” / “Sambas” | Borneo / occidental(?) / Sambas. Sambas is one of the regencies of West Kalimantan province (Indonesia) |
“H. pg” | It could be interpreted as an abbreviation of “Highlands of Palembang” based on Gorham’s original description of Luciola picea (Indonesia) |
“Alahan pandjang” | Alahan Panjang is in West Sumatra (Indonesia) |
“Nueva Vizcaya” / “Imugan” | Imugan is in the municipality of Santa Fe, province of Nueva Vizcaya (Philippines) |
Abbreviations follow
ASD distance between antennal sockets;
BP basal piece;
FS antennal flagellar segments;
GHW greatest head width (across eyes, measured parallel to ASD);
L length;
LL lateral lobes;
LO light organ;
ML median lobe;
MN mesonotal plates;
MPP median posterior projection ventrite 7 male only;
MS mesoscutellum;
W width;
V abdominal ventrites referred to by actual number e.g., V2, V6;
T abdominal tergites.
Elytral interstitial lines are numbered from 1, nearest the suture, to 4, nearest the lateral margin. Condition of the specimens precluded any attempt to investigate further features of the mouthparts including the nature of the apical labial palpomere. All figures have anterior end to top of page unless otherwise indicated.
We examined 15 specimens of lucioline fireflies in
An incomplete specimen of Atyphella testaceolineata Pic, 1939 (abdomen only) was redescribed and figured; no dissections were made.
Two specimens of Luciola cerea Gorham, 1882 were designated as lectotype and paralectotype and redescribed, and this species confirmed as a species of Curtos Motschulsky, 1845; a further two specimens labelled L. cerea were not considered to be types.
Luciola laticollis Gorham, 1883 was redescribed from a male lectotype and female paralectotype designation and confirmed to belong to the genus Luciola sensu stricto. Luciola nicollieri Bugnion, 1922 was redescribed from a lectotype (intact male) and paralectotype (male without head and prothorax) designation and confirmed to belong to the genus Luciola sensu stricto.
A single male of Pteroptyx decolor Olivier, 1911 from the type locality was designated a lectotype and redescribed. Coloured pictures of the holotype male and labels of Pygoluciola stylifer Wittmer, 1939, were provided to supplement previous depictions of only line figures and to correct label data.
The taxonomic status Luciola picea Gorham, 1882 was discussed. Existing taxonomic categories are not suitable for accommodating Luciola picea, which is considered a species inquirenda. A lectotype male and two paralectotype males, along with a female paralectotype were designated.
The species addressed here are listed with current, and original combinations (Table
Nomenclature of species addressed in this study in two forms: currently accepted and original names.
Current nomenclature | Original name |
---|---|
Atyphella testaceolineata Pic, 1939 | Atyphella testaceolineata Pic, 1939 |
Curtos cerea (Gorham, 1882) | Luciola cerea Gorham, 1882 |
Luciola laticollis Gorham, 1883 | Luciola laticollis Gorham, 1883 |
Luciola nicollieri Bugnion, 1922 | Luciola nicollieri Bugnion, 1922 |
Pteroptyx decolor Olivier, 1911a | Pteroptyx decolor Olivier, 1911a |
Pygoluciola stylifer Wittmer, 1939 | Pygoluciola stylifer Wittmer, 1939 |
Luciola picea Gorham, 1882 species inquirenda | Luciola picea Gorham, 1882 |
Atyphella
Luciola (Luciola)
Laporte. Sensu
Luciola (Atyphella)
(Olliff).
Atyphella lychnus Olliff, 1890.
Atyphella testaceolineata
Pic, 1939: 370.
Luciola (Luciola) testaceolineata
(Pic).
1 ♂ (incomplete, abdomen only; by original description and determined by authors in this study).
“Buru”.
(1♂ specimen). Holotype: INDONESIA ● ♂; (1) “L.J.TOXOPEUS / Buru,Station 7 / alt.Sept. 1921”; (2) “TYPE / Atyphella / testaceolineata / 1928 Pic”; (3) “?astyphella / testaceolineata / n. sp.”; (4) “Atyphella / testaceolineata /
Pronotum with median dark marking (from original description); elytra brown with suture, lateral margin and three longitudinal pale stripes corresponding to interstitial lines 1–3 (Fig.
Colour
(Fig.
Elytra
(Fig.
Abdomen
(Fig.
We consider this specimen a holotype because it corresponds with the original description outlined in Pic’s publication from 1939, especially in terms of its size and type locality. We can confirm features of colouration of the hind body as described by
Curtos
Curtos mongolicus Motschulsky by original designation.
Luciola cerea
Gorham, 1882: 103–104; 1887: 70.
2 ♂ (herein designated).
“Koetoer”.
(2♂ specimens). Lectotype (herein designated): INDONESIA ● ♂; (1) “cerea”; (2) “Sum. Exp. / Koetoer / 6/78”; (3) “Koetoer / 6.78”; (4) “
Luciola cerea Gorham lectotype male (A, E, G–V.
(2 ♂ non types). INDONESIA ● possibly ♂, abdomen missing; (1) “Rawas / 5.78”; (2)
The lectotype male,
Body length. 6.0 mm long. L/W 2.6.
Colour. Specimen with the accession number
Pronotum
(Fig.
Elytra
(Fig.
Head
(Fig.
Abdomen
(Fig.
Aedeagal sheath
(Fig.
Aedeagus
(Fig.
After discovering that the specimens we subsequently assigned to lectotype and paralectotype status had been mixed in with other specimens, we took extra care to verify whether the additional specimens we examined were part of the type material or not. We meticulously examined each specimen to ensure we accurately identified the type-material, and our results were reliable. Our thorough examination allowed us to confidently identify the actual type specimens and exclude any specimens not part of the type material. However, because it is not possible to confidently identify so many similarly coloured Curtos species (see further below) we retained a list above of two further non type specimens which may aid in any future revision.
There is no recent revision of Curtos apart from
In Gorham’s description of Luciola cerea, he noted the specimens he examined were all males (5–6 millimetres). He also listed six locations (see type localities) where the specimens were collected, which he referred to as “(Sum. Exp.)” for Sumatra Expedition. In the
A specimen with the accession number
Little detailed work has been done thus far on species of Curtos and examination of these Leiden types allowed WFAJ not only to dissect them but to expose various new features of their morphology especially in the male genitalia. Clearly, comparisons with other species are not presently possible, and it is very probable that many of these species are synonyms. The hooked posterior lobes attributed here to the sheath tergite have not been seen elsewhere in the Luciolinae. Species of Sclerotia Ballantyne have irregularly shaped sclerites in a band of muscle which surrounds the aedeagal sheath in life. Both may have the same function, that of extra surface area for muscle attachment, but their origins appear to be different.
Luciola
Laporte s. str. (sensu
Luciola
to be determined (see
Luciola (Luciola) Laporte.
Bourgeoisia Olivier.
Type species: Luciola antipodum Bourgeois designated by
See
Luciola laticollis
Gorham, 1883: 4.
“Java: Bodjonegoro”.
1♂, 1♀ (herein designated).
(1♂, 1♀ specimens). Lectotype (herein designated): INDONESIA ● ♂; (1) “♂”; (2) “type”; (3) “Luciola / laticollis / Gorham”; (4) “Piepers / Bodjonegoro / Java”; (5) “
Male with dark brown elytra having narrowly paler (appearing orange) lateral margins, apex and suture, light brown pronotum with a wide median dark brown marking (Fig.
Luciola laticollis Gorham, 1883 lectotype male (C, D, G–R) and paralectotype female (E, F). A, B. Specimen labels; C, D. Male dorsal (C), and ventral habitus (D); E, F. Female dorsal (E) and ventral habitus (F); G, H. Head, prothorax and anterior part of mesothorax, dorsal (G) and ventral (H); I. V7 (left) and V6 dorsal aspects; J. T8 dorsal; K–N. Aedeagal sheath: K. Dorsal view (arrow indicates midanterior margin sheath tergite); L. Dorsal view; M. Left lateral view (arrow indicates midanterior margin sheath tergite); N. Slightly oblique left lateral view; O–R. Aedeagus: O. Dorsal view (anterior left arrow indicates thickened lateral margin of base of ML; anterior right arrow indicates posterior extension of ML towards inner base of LL; posterior single left arrow indicates divergence of inner dorsal margins of LL); P. Ventral view (leafy lobes on inner margins LL arrowed); Q. Right lateral view (left arrow indicates base of LL in area of attachment of the ML; lower right arrow indicates posterior margin of BP); R. Slightly oblique left dorsolateral. All images are to scale, except specimen labels.
Body length. 5.5–6.0 mm long (Fig.
Colour
(Fig.
Pronotum
(Fig.
Elytra
(Fig.
Head. Not able to be retracted into prothoracic cavity; head width subequal to width across cavity; mouthparts well developed; antennal sockets not contiguous; antennal segments elongate slender, length antenna/GHW 1.5.
Abdomen
(Fig.
Aedeagal sheath
(Fig.
Aedeagus
(Fig.
Body length
(Fig.
Colour
(Fig.
Pronotum
(Fig.
Elytra
(Fig.
Head
(Fig.
Abdomen
(Fig.
This species is assigned to Luciola s. str. because of the distinctive features of the male aedeagus.
Luciola nicollieri
Bugnion, 1922: 1–14.
Luciola nicolleri.
2♂ (herein designated).
“Ceylan, Talgaswella, district d’Elpitiya”.
(2♂ specimens). Lectotype (herein designated): SRI LANKA ● ♂: (1) “E.BUGNION / Hiver 1906–7 / Ceylan / Talgaswella”; (2) “don.E.BUGNION’22”; (3) “COTYPE: Luciola / Nicollieri / Bugnion / ♂”; (4) “Luciola. / nicollieri / Bugnion, 1922 / ZMAN type COLE.0929.1”; (5) “
Male with orange pronotum, black elytra with narrow pale orange lateral and sutural margins, elytral apex appearing more widely pale due in part to an accumulation of fat body. Venter black except for yellowish creamy LO in V6, 7. The only Luciola s. str. so far recorded with pale coloured pronotum without darker markings, and dark brown to black elytra with all margins pale except at the base.
Body length
(Fig.
Luciola nicollieri Bugnion, 1922 lectotype male (A, C, E, F, H–Q) and paralectotype male (B, D, G. Without head and prothorax). A, B. Specimen labels; C. Ventral mesothorax – end of abdomen; D. Dorsal habitus; E. Dorsal head prothorax and anterior area of mesothorax; F. Anterior head; G. Ventral, V5–7 and elytral apices; H. V7 ventral; I, J. Tergite 8 – T8 dorsal view (I) and ventral view (J); K–N. Aedeagal sheath: K. Dorsal with aedeagus ventral surface to right; L. Ventral view; M. Dorsal view; N. Left lateral; O–Q. Aedeagus: O. Dorsal view (arrow top left indicates thickened left margin of ML, lower arrow thickened lobe from inner margin of LL, lower arrow right side indicates area of attachment to inner surface of base of LL); P. Ventral view (upper oblique arrows left and right indicate leafy lobes from inner margins of LL, lower left arrow lateral expansion of ML margins); Q. Left lateral (upper arrow indicates junction between lobes from ML to left and lobes from LL to right; lower right arrow indicates area of attachment to inner base of LL). All images are to scale, except specimen labels.
Colour
(Fig.
Pronotum
(Fig.
Elytra
(Fig.
Head
(Fig.
Abdomen
(Fig.
Aedeagal sheath
(Fig.
Aedeagus
(Fig.
Pteroptyx
Luciola testacea Motschulsky designated by
Poluninius Ballantyne, 2013: 100.
Type species: Poluninius selangoriensis Ballantyne 2013: 101.
Pteroptyx decolor
Olivier, 1911a: 17. Olivier 1913: 55.
1♂ (herein designated).
“Atjeh”.
(1♂ specimen). Lectotype (herein designated): INDONESIA ● ♂: (1) “Atjeh”; (2) “Coll. Veth”; (3) “Pteroptyx / decolor / Ern. Oliv.”; (4) “
Very pale yellowish brown dorsal colouration, with elytra semi-transparent and globules of fat body showing beneath pronotal cuticle; ventral surface very pale brown, with globular fat bodies visible beneath abdominal ventrites, LO in V6, 7 cream; labrum (partly obscured) antennae, palpi and tarsi dark brown, head between eyes yellowish brown. P. decolor belongs to that group of Pteroptyx which do not have paired lobes to either side of T8 emargination, the deflexed elytral apices are elongated and not dimpled on the posterior margin, the posterolateral corners of V7 are rounded and scarcely produced, and the MPP of V7 has a flat dorsal surface with a short posterior prolongation. Most similar to P. valida from which it is distinguished by the paler colour (P. valida has black elytral apices) and its occurrence in Indonesia.
Body length
(Fig.
Colour
(Fig.
Pteroptyx decolor Olivier, 1911 lectotype male. A. Specimen labels; B. Dorsal habitus; C. Ventral anterior body including head; D. Ventral abdominal apex and apex left elytron; E–G. V7 ventral view (E), dorsal view (F), and G posterior with ventral surface uppermost (projection of tip of MPP of V7 arrowed in E, G). H–I. T8 dorsal and ventral (oblique arrows in I indicate position of flanges); J–K. Aedeagal sheath: J. Ventral view; K. Dorsal view (paraprocts arrowed in J; widest margin of sternite arrowed in K); L–N. Aedeagus: L. Ventral view; M. Dorsal view; N. Right lateral (upper arrows M, N anterior dorsal margin of LL; lower arrows M, N attachment of ML to inner surface of LL). All images are to scale, except specimen labels.
Pronotum. With lateral margins subparallel-sided, posterior angles almost 90°, anterolateral corners rounded obtuse.
Elytra
(Fig.
Abdomen
(Fig.
Aedeagal sheath
(Fig.
Aedeagus
(Fig.
In Olivier’s original description, he described Pteroptyx decolor based on a male from “Atjeh” and a female from “Borneo” (
In
It is very probable that the following references to P. decolor are a presently undescribed species:
We herein designated a lectotype for Pteroptyx decolor to reduce the potential for confusion, especially considering previous misidentifications.
Pygoluciola
Luciola subgenus Pygoluciola
(Wittmer).
Pygoluciola stylifer
Pygoluciola stylifer
Wittmer, 1939: 21–32.
Luciola (Pygoluciola) stylifer.
1 ♂ (by original description, as indicated by Wittmer’s original type label and determined by authors in this study).
“Long Petah” (from page 3, “Fundort: “M.O.-Borneo Expedition, Long Petah, 450 in, IX–X.1925 (leg. H C. Siebers)”).
(1♂ specimen). Holotype: INDONESIA ● ♂; (1) “Holotype”; (2) “H. C. Siebers / M.O. Borneo Exp. / 1200 M. bij L. Petak / 15–20 X. 1925 VIII–IX”; (3) Pygoluciola / stylifer / Wittm. / det. W. Wittmer”; (4) “T.Y.P.US.” (Fig.
P. stylifer belongs to that group of Pygoluciola where males have the posterior margin of V7 and T8 narrowly prolonged and curving (Fig.
Here, we have the first opportunity to present coloured pictures of the type male and verify the original labels that come with the specimen.
Luciola picea
Gorham, 1882: 104–105; 1887: 71.
4 ♂ (herein designated).
“Palembang bovenland”.
(4♂ specimens). Lectotype (herein designated): INDONESIA ● ♂; (1) “Luciola / picea, Gorh:”; (2) “Sum. Exp. / Palembang / bovenland / 5 of 6.78”; (5) “
We can confirm only that this species does not conform to Luciola s. str. in features of the aedeagus (see Fig.
In Gorham’s original description, he mentioned the specimens are all males from four localities, suggesting that there could be at least another male syntype (
While the unique holotype remains the pinnacle for species identification, locating it can be a daunting task. Additionally, while museums may indicate they possess a holotype, further investigation often reveals that this supposed unique specimen is part of a syntype series. We have been able to establish the status of the specimens standing in this collection by detailed examination of the specimen labels, and corroboration by similar examination of how the literature was worded when the specimen was first described.
The holotype itself may be so old and discoloured that it conveys little. Fortunately, all but one of these specimens have retained all sections and only display the inevitable loss of some colour over the original description. Luciolinae taxonomy has come to rely more and more on features of the male genitalia, including that of the last abdominal segments, which are retracted within the abdomen (the aedeagal sheath of
Here, we have had the good fortune to address seven name-bearing type specimens which fulfil most of the conditions we outlined above. They are either identified as types (though they may be syntypes), are from the original locality specified in the description and conform to the original description. The museum was generous in its practices and not only loaned but permitted dissection.
Additionally, the detailed dissections allow us to present descriptions of certain features of the male not previously addressed. The Curtos species are shown to have the posterior area of their narrow aedeagal sheath tergite with paired asymmetrical hooks. The means of attachment of the dorsal surface of the aedeagal median lobe to the inner surface of the lateral lobes is investigated and certain generic distinctions are described.
Overall, this study demonstrates that despite the numerous challenges taxonomists face in identifying specimens, a detailed examination of the specimen labels and literature, along with delicate dissections of male specimens, can help overcome some of these challenges and shed light on previously unknown taxonomic features.
This study was funded by Vereniging Nederland-Maleisië (Netherlands-Malaysia Association) through the VNM Fellowship 2019 to Wan F. A. Jusoh. Menno Schilthuizen was instrumental in hosting Wan Jusoh in Leiden in late November 2019 and making sure the visit was successful. At Naturalis, Hans Huijbregts (former curator of Coleoptera collection) provided Wan with his expert knowledge to assist with tracking specimens and organise them for loan. Due to a series of COVID-19 ‘circuit breaker’ between 2020 and 2021, the dissection of material and high-resolution imaging was done in several stages at Lee Kong Chian Natural History Museum, Singapore (where Wan was a Research Fellow) and finally completed at the end of 2022. Wan and Lesley undertook the final examination of the specimens between Feb and March in 2023 for which they like to thank Dr Robert Ballantyne for his tremendous support in making sure nothing gets in the way until the final manuscript is ready. The final preparation of the manuscript could not be continued and completed without a seed funding from Monash University Malaysia to Wan (SED000135).