Corresponding author: Ladislav Bocak (
Academic editor: L. Penev
The Oriental neotenic net-winged beetles attracted attention of biologists due to conspicuous large-bodied females; nevertheless phylogenetic relationships remain contentious and only a few species are known in both the fully metamorphosed males and neotenic females. The phylogenetic analyses and morphology of larvae and adults provide data for investigation of relationships and species delineation.
Masek M, Bocak L (2014) The taxonomy and diversity of
The platerodriline net-winged beetles are one of elateroid lineages with modified female morphology (
The taxonomy of the neotenic lineages has quite short history despite the fact that the trilobite larvae were first time described in the 19th century (
The DNA data represent an independent source of information for species delineation (
Adult males and female larvae were used for morphological descriptions. A part of specimens used for morphological study was sequenced and labelled with the GenBank voucher numbers in the format UPOL + six-letter/number code and the status of all type specimens were designated with red labels (
Taxonomic coverage, locality data and GenBank accession numbers.
Species | Voucher UPOL+ | Local. data |
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000572 | Japan |
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000573 | Japan |
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000570 | Japan |
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000571 | Japan |
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001012 | Japan |
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001002 | Sumatra |
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001008 | Malaysia |
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000L02 | Sabah |
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000801 | Sumatra |
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000802 | Java |
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000805 | Sumatra |
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000358 | Java |
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000574 | Japan |
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VP0016 | India |
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VP0017 | India |
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VP2312 | India |
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VP0003 | Malaysia |
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VP0004 | Malaysia |
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000585 | Malaysia |
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VP0001 | Sumatra |
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VP0006 | Sumatra |
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000L11 | Malaysia |
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VP0002 | Sumatra |
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000584 | Sumatra |
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001043 | Malaysia |
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VP0050 | Malaysia |
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000L21 | Malaysia |
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000M45 | Java |
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VP0009 | Malaysia |
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VP0010 | Sumatra |
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VP0012 | Malaysia |
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VP0030 | India |
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VP0031 | India |
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VP0034 | India |
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001380 | Mindanao |
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001381 | Mindanao |
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001383 | Mindanao |
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VP0014 | Mindanao |
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VP2316 | Mindanao |
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001388 | Sumatra |
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001373 | Sumatra |
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000588 | Borneo |
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000589 | Borneo |
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000586 | Java |
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001379 | Sumatra |
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001387 | Sumatra |
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001374 | Sumatra |
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001377 | Sumatra |
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001386 | Sumatra |
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VP2303 | Sumatra |
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VP2306 | Sumatra |
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001371 | Sumatra |
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VP2308 | Sumatra |
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VP0021 | Singapore |
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000587 | Sumatra |
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001378 | Sumatra |
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001372 | Sumatra |
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001389 | Sumatra |
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000L01 | Sabah |
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VP0044 | Sabah |
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VP2301 | Sabah |
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VP0020 | Malaysia |
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VP0047 | Sumatra |
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VP2302 | Malaysia |
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VP2304 | Thailand |
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VP2307 | Sumatra |
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VP2309 | Malaysia |
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VP2310 | Malaysia |
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VP2311 | Laos |
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MB1382 | Palawan |
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001375 | Sumatra |
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001376 | Sumatra |
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001385 | Sumatra |
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VP2305 | Sumatra |
|
–length of body –width at humeri –length of pronotum –width of pronotum –maximum eye diameter –minimum interocular distance in frontal part of cranium –Voucher collection of the Laboratory of Molecular Systematics, Faculty of Science UP, Olomouc –Natural History Museum, London; MNHP–Museum d'histoire naturelle, Paris –Zoological Reference Collection, Raffles Museum of Biodiversity Research, NUS –Kiyoshi Matsuda Collection, Takarazuka
Total DNA was extracted using Wizard SV96 kit (Promega Inc.) and primers 16a (5'-CGCCTGTTTAACAAAAACAT-3'), 16b (5'-CCGGTCTGAACTCAGATCATGT-3') and ND1A (5'-GGTCCCTTACGAATTTGAATATATCCT-3') were used for PCR amplification of the 530–810 base pairs of
Sequences were edited using the Sequencher 4.8 software package (Gene Codes Corp.). The
The ultrametric tree was produced from the tree depicted in
Phylogenetic hypothesis for
The DNA sequences of
The
The GMYC analysis of the normalized tree was used as an independent test for morphology based delineation of species. The analysis suggested the clusters designated as
The origin of large-bodied neotenic larvae was recovered in three unrelated lineages:
The results confirm that
The subfamiliar and tribal classification of
The other
Adult male, general appearance:
The sister clade to the
The
Male pronotum of
Larvae of
Further two genus-group names were proposed for species now placed in
Male genitalia of
Male genitalia of
Male genitalia of
To sum up, we propose to consider
Rapid morphological divergence in male genitalia is widespread and results largely from sexual selection (
Head small, prognathous to slightly hypognathous, partly retracted in pronotum. Eyes hemispherically prominent, frontal interocular distance longer than maximum eye diameter. Labrum sclerotized, transverse, separated from clypeus, mandibles slender, long, slightly curved, incisor margin simple, without teeth, maxilla tiny, with setose mala, stipes plate-like, cardo vestigial, palpifer short, maxillary palpi 4-segmented, palpomere 1 shortest, about twice longer than palpifer, palpomere 2 longest, slender, palpomere 3 slightly longer than wide, apical palpomere slender, drop-like, with slender apical part. Labium reduced, mentum plate-like, formed by single sclerite, ligula absent, palpi 3-segmented, basal palpomeres subequal, rectangular to slightly longer than wide, apical palpomere twice longer than wide at base, pointed to apex.
Antennal tubercles present, usually strongly prominent. Antennae 11-segmented, slightly surpassing middle of elytral length, dark coloured, never with apical antennomeres pale, antennae usually weakly serrate, a few species with flabellate antennae. Scapus pear-like, robust, pedicel and antennomere 3 subequal in length, antennomeres of serrate antennae flattened, from antennomere 4 gradually slenderer, apical antennomere long, parallel-sided. Flabellate antennae with lamellae of antennomeres 3–10 longer than body of antennomere; whole antennae with dense erected pubescence.
Pronotum transverse with prominent to obtuse anterior angles (
Sexually mature larviform females observed only by
The information on biology was given by
Syntype. Male (BMNH), Borneo. Syntype. Female (BMNH), Lundu, Sarawak, G. E. Bryant, 6. 1. 14.
Male. Body 7 mm, dark brown, only humeri and elytral suture slightly lighter. Head small, with hemispherically prominent eyes, head with eyes wider than frontal margin of pronotum, eye diameter 1.9 times frontal interocular distance. Antennae compressed, covered with long, erected, dense pubescence. Pronotum flat, without carinae, 1.8 times wider than long at midline, frontal margin widely rounded anteriorly, frontal angles obtuse, lateral margins almost straight, posterior margin bisinuate. Elytra flat, parallel-sided, elytral costae inconspicuous, elytra 2.6 times longer than width at humeri. Legs slender, compressed, densely pubescent. Male genitalia with robust parameres, phallus slightly curved with bulbous tip. Parameres stout with hooked tip, apical half of ventral edge serrate. Phallobase wide, deeply emarginate.
Female larva. Body flat and wide (
Male. BL 6.9 mm, PL 1.0 mm, PW 1.9 mm, HW 2.0 mm, Edist 0.85 mm, Ediam 0.44 mm. Larva. BL 53.0 mm, PL 8.7 mm, PW 13.4 mm.
Malaysia: Sarawak. Known only from the type locality.
Holotype. Male (LMBC, UPOL 000589), Borneo, Central Kalimantan Prov., 60 km SE Muara Teweh,
Male. Body small-sized, brown, head, antennae, legs except bases of femora and apical three fifths of elytra dark brown (
Female larva. Body extremely flat and wide (
Male. BL 6.0 mm, PL 0.9 mm, PW 1.5 mm, HW 1.7 mm, Edist 0.78 mm, Ediam 0.46 mm. Larva. BL 19.3 mm, PL 5.1 mm, PW 11.2 mm.
Indonesia: Kalimantan. Known only from the type locality.
The specific epithet refers to the flat body shape of the female larva.
The males and female larvae were identified as conspecific on the basis of highly similar sequences of
Holotype. Male (LMBC), Sumatra Utara, Brastagi, Gn. Sibayak, 19–23. Feb. 1998, 700–2000 m.
Body brown, head, pronotum, mesoscutellum and elytra yellow (
BL 7.4 mm, PL 1.0 mm, PW 1.6 mm, HW 1.9 mm, Edist 0.91 mm, Ediam 0.38 mm.
Indonesia: Northern Sumatra.
The specific epithet is a patronym in honour of Alvin T. C. Wong.
1 | Parameres short, reaching slightly over half of phallic length | |
– | Parameres reaching almost to the apex of the phallus | 2 |
2 | Male antennae flabellate | |
– | Male antennae serrate | 3 |
3 | Whole elytra yellow ( |
4 |
– | Elytra dark brown or light brown with dark coloured apical part | 5 |
4 | Parameres reaching to five sixths of the phallic length | |
– | Parameres reaching to three fourths of the phallic length | |
5 | Only apical part of elytra dark coloured, pronotum black | |
– | Whole elytra dark brown, at most the narrow humeral part slightly lighter brown | 6 |
6 | Body slender, inner margin of parameres serrate only in its apical half | |
– | Body robust, inner margin of parameres serrate in three quarter of their length |
* We failed to find any distinguishing character between
Holotype. Male (LMBC), Singapore, Bukit Timah and Central Water Catchment, 19.–22. May 2013, 50–100 m, E. Jendek and O. Šauša leg. Paratypes. Female larvae, 6 spec., Malaysia, Pahang, Tioman, Kg. Tekek–Juara trail, 50–300 m,
Male. Body small, dark brown to black, head, prothorax, mesoscutellum and basal two thirds of elytra dark red; whole body with dense, short, pubescence. Head small, including eyes slightly wider than frontal margin of pronotum. Eyes hemispherically prominent, frontal interocular distance 1.8 times eye diameter. Antennae robust at base, compressed, reaching two thirds of elytral length, antennomere 3 0.6 times antennomere 2, antennomere 3 as long as wide at apex. Pronotum flat, 1.7 times wider than long at midline. Anterior margin almost straight, anterior angles sharply marked, posterior margin bisinuate. Elytra slightly wider posteriorly, elytra 2.8 times longer than width at humeri; only slightly widened posteriorly, elytral costae conspicuous. Legs compressed, densely pubescent. Male genitalia with straight phallus and setose parameres, short phallobase slightly shorter than parameres.
Female larva. Body slender, parallel-sided (
Male. BL 6.9 mm, PL 0.9 mm, PW 1.6 mm, HW 2.1 mm, Edist 0.86 mm, Ediam 0.48 mm. Larva. BL 30.2 mm, PL 5.7 mm, PW 5.6 mm.
Singapore, Malaysia: Pahang. Biology and female specimens collected in Singapore were reported by
The species name is a patronym in honour of Peter Ng.
Holotype. Male (LMBC), Java, K. O. Blawan, Ijen Plateau, Jul. 1940, 900–1500 m, H. Lucht coll.
Male. Body medium-sized, dark brown, head, prothorax, mesoscutellum and basal half of elytra testaceous; antennae, legs, apical half of elytra dark brown to black (
BL 10.1 mm, PL 1.1 mm, PW 2.0 mm, HW 2.6 mm, Edist 0.95 mm, Ediam 0.49 mm.
Indonesia: Java.
The specific epithet is a patronym in honour of the late W. Wittmer, who donated the specimen to the senior author in 1992.
1 | Metathorax orange brown | |
– | Metathorax dark brown to black | 2 |
2 | Phallobase shorter than parameres (≤ 0.95 times length of parameres) | |
– | Phallobase longer than parameres (≥ 1.05 times length of parameres | 3 |
3 | Phallus short and stout, about 0.90 the combined length of parameres and phallobase, pronotum similar in colour to humeral part of elytra | 4 |
– | Phallus long and slenderer, about as long as the combined length of parameres and phallobase, pronotum blac | |
4 | Antennomere 4 wide, about as long as wide at apex | |
– | Antennomere 4 at least 1.25 times longer than wide at apex | 5 |
5 | The expanded base of phallus in lateral view mostly hidden by parameres, phallobase deeply emarginate basally, elytra black apically | |
– | The expanded base of phallus in lateral view exposed, considerably widened, phallobase shallowly emarginate basally, elytra testaceous |
Holotype. Male (LMBC, 000586), Java, Ijen N. P., 12 km W of Sodora, 3–5. May 2001, 1000 m.
Body and head dark brown to black, pronotum, mesoscutellum and elytra in humeral half orange, apical part of elytra dark brown to black (
BL 7.2 mm, PL 0.9 mm, PW 1.6 mm, HW 1.8 mm Edist 0.70 mm, Ediam 0.40 mm.
Indonesia: Java.
The specific name refers to the type locality.
Lectotype (hereby designated). Male (BMNH), Sumatra, Korinchi, 4500 ft, N. 1914, K. G. Blair. Paralectotype. Female (BMNH), same locality data.
The male genitalia are missing. Therefore, only information on external morphology can be compared. The orange pronotum and humeral part of elytra resemble
PL 1.3 mm, PW 1.9 mm, HW 2.6 mm, Edist 0.84 mm, Ediam 0.36 mm.
Indonesia: Sumatra, Jambi, Kerinci massif.
The lectotype is damaged (the apical half of elytra and abdomen are missing).
Holotype. Male (LMBC, UPOL 001379), Sumatra, Jambi Kersik Tua, Gn. Kerinci, 19.–22. Jan. 2005, 1600–2200 m.
Body black, head dark brown, pronotum and elytra orange; antennae, and legs dark brown to black (
BL 10.1 mm, PL 1.1 mm, PW 1.8 mm, HW 2.3 mm, Edist 0.83 mm, Ediam 0.45 mm.
Indonesia: Sumatra.
The specific epithet refers to yellow colouration of the body.
Holotype. Male (LMBC, UPOL 001386), Sumatra, Barat Lake Maninjau, E coast, 12.–23. Jan. 2005, 800 m. Paratype. Male (LMBC, UPOL 001374), Sumatra, Barat Lake Maninjau, E coast, 12.–23. Jan. 2005, 800 m. Paratypes. Male, 2 females (LMBC, UPOL 001377), Sumatra, Barat, Pasaman, Gn. Talamau, 14.–15. Jan. 2005, 1000 m; female larva (LMBC, UPOL VP2303), Sumatra, Barat Lake Maninjau, E coast, 12.–23. Jan. 2005, 800 m.
Male. Body medium-sized, dark brown; head, prothorax, mesoscutellum and basal three fifths of elytra orange; antennae, legs, apical two fifths of elytra dark brown to black (
Female larva. Body flat, wide (
BL 7.0 mm, PL 0.9 mm, PW 1.6 mm, HW 1.7 mm, Edist 0.84 mm, Ediam 0.38 mm. Larva. BL 24.1 mm, PL 5.4 mm, PW 10.3 mm.
Indonesia: Sumatra.
The specific epithet refers to the type locality of the holotype.
Holotype. Male (LMBC, UPOL 001371), Sumatra Utara, Brastagi, Gn. Sibayak, 26. Jan.–1. Feb. 2005, 1600–2200 m. Paratype. Female larva (LMBC, UPOL VP2308), Sumatra Utara, Brastagi, Gn. Sinabung, 29.–30. Jan. 2005, 1400–2000 m.
Body medium-sized, dark brown to black, only basal half of elytra orange and pronotum with irregular light coloured patches in disc (
Female larva. Body flat, wide (
BL 8.2 mm, PL 0.9 mm, PW 1.8 mm, HW 2.1 mm, Edist 0.94 mm, Ediam 0.39 mm. Larva. BL 32.3 mm, PL 7.7 mm, PW 13.0 mm.
Indonesia: Sumatra, North Sumatra Province, Gn. Sibayak.
The specific epithet is derived from the Latin adjective
Holotype. Male (LMBC, UPOL 001382), Philippines, Palawan, Tanabak river, 150 m,
Body medium-sized, dark brown to black, only pronotum, mesoscutellum and humeral two thirds of elytra orange; apical third of elytra and two thirds of elytral suture dark brown to black. Head small, including eyes apparently narrower than frontal margin of pronotum, antennal tubercles robust, deeply separated. Eyes hemispherically prominent, eye diameter 1.9 times frontal interocular distance. Antennae flabellate, antennomere 3 with long process, pubescent, length of antennomere 3 1.6 times antennomere 2. Pronotum strongly transverse, 1.5 times wider than long at midline, anterior margin straight, with prominent anterior angles, lateral margins almost straight, posterior margin bisinuate, surface of disc mat, finely punctuate, with dense short pubescence. Elytra 2.9 times longer than width at humeri, elytra almost parallel-sided; slightly widened posteriorly, widest at apical fourth, elytral costae inconspicuous. Legs compressed, with dense pubescence. Male genitalia with curved phallus, short rounded parameres bearing slender membranous processes; phallobase wide, deeply emarginate (
BL 7.1 mm, PL 1.0 mm, PW 1.6 mm, HW 1.9 mm, Edist 0.66 mm, Ediam 0.34 mm.
Philippines: Palawan.
The specific epithet refers to the type locality of the holotype.
Holotype. Male (LMBC, UPOL 000587), Sumatra, SW coast of Ranau Lake, 1–4. Jun. 2001, 1200 m.
Body medium-sized, dark brown to black, only pronotum testaceous to brown and basal quarter of elytra orange (
BL 6.0 mm, PL 0.8 mm, PW 1.3 mm, HW 1.7 mm, Edist 0.79 mm, Ediam 0.35 mm.
Indonesia: Sumatra.
The specific epithet refers to the type locality of the holotype.
Holotype. Male (LMBC, UPOL 001389), Sumatra Utara, Brastagi, Gn. Sibayak, 26. Jan.–1. Feb. 2005, 1600–2200 m. Paratypes. 2 males (LBMC), Sumatra, SW of Brastagi, Gn. Sinabung, 22. Feb. 1991, 1300–1800 m. Paratype. Male (LBMC), Sumatra, SW of Brastagi, Gn. Sinabung, 19.–23. Feb. 1991, 1300–1800 m. Paratype. Male (LBMC, UPOL 001372), Sumatra, Utara, Brastagi, Gn. Sibayak, 26. Jan.–1. Feb. 2005, 1600–2200 m. Paratype. Male (LBMC), Sumatra, Sinabung, Mar. 1998.
Body medium-sized, dark brown to black, pronotum testaceous to brown and basal third of elytra orange (
BL 8.5 mm, PL 0.9 mm, PW 1.8 mm, HW 2.0 mm, Edist 0.73 mm, Ediam 0.44 mm.
Indonesia: Sumatra.
The specific epithet refers to the type locality of the holotype.
Holotype. Male (LMBC), Sumatra, SW of Brastagi, Gn. Sinabung, 22. Feb. 1991, 1300–1500 m. Paratypes. 2 males (LBMC), Sumatra, SW of Brastagi, Gn. Sinabung, 22. Feb. 1991, 1400–1900 m.
Body medium-sized, dark brown to black, only basal two fifths of elytra orange testaceous (
BL 8.9 mm, PL 1.0 mm, PW 1.7 mm, HW 2.1 mm, Edist 0.89 mm, Ediam 0.4 mm.
Indonesia: Sumatra.
The specific epithet refers to the type locality of the holotype.
Holotype. Male (LMBC, UPOL 001376), Sumatra Barat, Pasaman, Gn. Talamau, 14.–15. Jan. 2005, 1000 m. Paratype. Male (LMBC, UPOL 001375), Sumatra Barat, Pasaman, Gn. Talamau, 14.–15. Jan. 2005, 1000 m.
Male. Body medium-sized, dark brown to black, only pronotum, mesoscutellum and basal three fifths of elytra orange (
BL 6.5 mm, PL 1.0 mm, PW 1.6 mm, HW 1.9 mm, Edist 0.88 mm, Ediam 0.36 mm.
Indonesia: Sumatra.
The specific epithet refers to the type locality of the holotype.
Holotype. Male (LMBC, UPOL 001385), Sumatra, Jambi Kersik Tua, Gn. Kerinci, 19–22. Jan. 2005, 1600–2200 m. Paratype. Female (LMBC, VP2305), Sumatra, Jambi prov. Kerinci Seblat N. P., 7 km E Kayuaro, Mt. Tujuh,
Body dark brown to black; head, pronotum and elytra yellow to orange, apical margins of elytra infuscate (
Female larva. Body flat, wide, dark brown, only margins of tergites lighter, pronotum triangular (
BL 8.1 mm, PL 0.9 mm, PW 1.6 mm, HW 1.9 mm, Edist 0.81 mm, Ediam 0.31 mm. Larva. BL 32.7 mm, PL 6.6 mm, PW 10.2 mm.
Indonesia: Sumatra.
The specific epithet refers to the locality of the paratype.
Holotype. Sumatra, Sungei Kumbang, Korinchi, 4500 ft, Apr. 1914 (BMNH). Other material examined. Male (LMBC, UPOL 001378), Sumatra, Jambi Kersik Tua, Gn. Kerinci, 19.–22. Jan. 2005, 1600–2200 m.
Body black; head and mesoscutellum dark brown, pronotum and elytra orange yellow; antennae, and legs dark brown to black (
BL 7.8 mm, PL 1.0 mm, PW 1.6 mm, HW 2.0 mm, Edist 0.85 mm, Ediam 0.40 mm.
Indonesia: Sumatra.
Holotype, male, Presqu'ile Malaise (MNHP).
The type series contains just female larvae judging from the illustrations might include two species. The adult is unknown.
We are obliged to P. Ng and A. Wong for providing as with the manuscript of A. Wong's Master Thesis and for permission to publish all information. The authors are grateful to R. Bilkova for technical assistance, to M. Barclay for providing an access to the BMNH collection and to K. Matsuda for an access to additional material. The study was funded by the Grant Agency of the Czech Republic (P506/11/1757) and IGA Project from UP Olomouc. Additionally, this research received support from the Synthesys Project, which is financed by EC Research Infrastructure Action.