Research Article |
Corresponding author: Radim Gabriš ( gabris.radim@gmail.com ) Academic editor: Francesco Vitali
© 2017 Radim Gabriš, Filip Trnka, Rodzay Abdul Wahab, Robin Kundrata.
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:
Gabriš R, Trnka F, Wahab RA, Kundrata R (2017) Taxonomic revision of the endemic Bornean genera Anexodus Pascoe and Pantilema Aurivillius (Coleoptera, Cerambycidae, Lamiinae). ZooKeys 669: 29-51. https://doi.org/10.3897/zookeys.669.12608
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The genera Anexodus Pascoe, 1866 and Pantilema Aurivillius, 1911 (Cerambycidae: Lamiinae: Morimopsini), both endemic to Borneo, are revised. Four species of Anexodus are recognized: A. aquilus Pascoe, 1886 (Malaysia: Sabah), A. sarawakensis Sudre, 1997 (Malaysia: Sarawak), A. syptakovae sp. n. (Malaysia: Sarawak), and A. tufi sp. n. (Brunei). Pantilema is a monotypic genus containing P. angustum Aurivillius, 1911 (Malaysia: Sarawak) which is known only from the holotype. For the first time, genital structures are studied in these genera. An identification key for the species of Anexodus is provided and their intraspecific morphological variability and distributions are discussed.
Brunei, diversity, endemism, hot-spots, longhorn beetles, Malaysia, Morimopsini
Cerambycidae forms one of the largest and most well-known beetle lineages in the world (
To finish a revision of the genera classified in Morimopsini in Borneo, we herein review the species of Anexodus and Pantilema. For the first time, male and female genitalia are investigated and an identification key is provided for the species of Anexodus.
In this study we examined mounted adults of both sexes. Genitalia were briefly submerged in hot 10% KOH, dissected and transferred to glycerol. Main diagnostics were photographed using a Zeiss Discovery.V12 with ZEN software. The line illustrations were derived from the photographs. All dissected parts were mounted on separate cardboards using Dimethyl Hydantoin Formaldehyde (DMHF) resin and pinned under the specimens. The measurements of taxonomically relevant morphological structures were taken with a measuring tool in ZEN software as follows: body length (BL) measured from the fore margin of head to the apex of elytra; body width (BW), pronotal width at the widest part; pronotal length at midline. Data from the locality labels are cited verbatim. A slash (/) is used to separate lines on the same label and a double slash (//) is used to separate different labels on the pin. The morphological terminology is used as in
PCDH personal collection of Daniel J. Heffern, Houston, TX, USA
PCJC personal collection of James S. Cope, San Jose, CA, USA
UBDC Universiti Brunei Darussalam, Brunei
Anexodus Pascoe, 1886: 242.
Anexodus aquilus Pascoe, 1886.
This genus is easily recognizable among the Bornean Morimopsini by its antennae, which are always shorter than body (Figs
Body elongate to elongate-oval, small to medium-sized. Body densely clothed with very short pubescence; coloration either more or less uniformly brown or brown with yellowish stripes ranging from vertex through sides of pronotum to basal part of elytra, mouthparts lighter; in some cases antennae reddish brown or black (Figs
Head about the same width as anterior margin of pronotum; genae sub-parallel at frontal view; frontoclypeus with distinct midline running from interantennal groove to labrum, sparsely covered with large, rounded, deep punctures; antennal tubercles prominent with deep narrow depression in between; antennal cavities opened dorsally; anterior margin of anteclypeus shallowly emarginate, with sparse long yellowish semi-erect setae. Labrum free, transverse, glabrous, either with one row of punctures bearing long setae (Figs
Prothorax subcylindrical, about as long as wide, widest before middle, then gradually narrowed towards posterior margin, laterally with one small more or less distinct tubercle; pronotal disc weakly convex, sparsely covered with deep punctures, with indistinct tubercles, anterior and posterior angles obtuse. Prosternum in front of coxae slightly shorter than diameter of coxal cavity, procoxal cavities circular, with lateral extension, narrowly separated. Scutellum transverse, widely rounded apically, about two times as wide as long. Elytra elongate, 1.6–1.8 times as long as wide at widest part, 1.7–2.1 times as long as pronotum in males and 2.0–2.3 times in females, basally slightly wider than posterior pronotal margin, widest near middle, from middle gradually tapered towards apex, fused along suture; each elytron with three rows of tubercles irregular in size, in some cases inner row forming a distinct ridge basally (Figs
Abdomen with five ventrites (Figs
Anexodus aquilus Pascoe, 1886: 242.
Anexodus
kuntzeni
Kriesche, 1924: 291; synonymized by
(A. aquilus). Holotype, male, [Malaysia] “Type [circular label with red margin, printed] // N / Borneo [blue oval label, handwritten] // Anexodus / aquilus / type Pasc. [handwritten] // Anexodus / aquilus / N. Borneo Pa [handwritten] // Pascoe / Coll. / 93-60” (
(A. kuntzeni). Holotype, female, [Malaysia] “N:Borneo / Kina-Balu-Geb. / Waterstradt S. [printed] // Anexodus / küntzeni Kriesche / Typ! [handwritten] // Anexodus / aquilus Pasc. / Breuning dét. // HOLOTYPUS / Anexodus / kuntzeni
Female, “Malaysia, Sabah, / Crocker Range 20- / IV-2007 Cope / collection” (PCJC); female, “Malaysia, Sabah / Crocker Range / III-22-2009 / local coll” (PCDH); female, “Malaysia, Sabah / Sandakan / II-12-2007 / local coll // Anexodus / aquilus Pasc. / det. D. Heffern ‘07” (PCDH); female, “Malaysia, Sabah / Ranau 700 m / IV-23-2015 / local coll” (PCDH); female, “Malaysia, Sabah / Tenom / III-2-2008 / local coll” (PCDH); male, “Malaysia, Sabah, Mt. / Trus Madi 26-IV-2010 / Cope Collection” (PCJC); male, “Malaysia, Sabah, Mt. / Trus Madi 26-V-2012 / Cope Collection” (PCJC); male, “Malaysia, Sabah / Mt. Trus-Madi / IV-15-2005 / local coll // Anexodus / aquilus / Pascoe / det. J. Sudre” (PCDH).
This species is similar to A. tufi sp. n. in general habitus, yellowish stripes on dorsal body surface and a labrum with the entire surface punctured but differs in having apex of scape with a distinct lateral hook-shaped projection (simple in A. tufi sp. n.; Figs
Holotype (male). BL 11.5 mm, BW 3.8 mm. Body brown with yellowish stripes extending from vertex through sides of pronotum to basal part of elytra; mouthparts lighter; antennae reddish brown. Body densely clothed with very short brown pubescence. Head about as wide as anterior margin of pronotum. Labrum transverse, with whole surface moderately sparsely, irregularly punctured (Fig.
Prothorax as long as wide, laterally with one distinct tubercle; pronotal disc with a pair of indistinct tubercles near middle and one median at second half; pronotal tubercles punctate. Prosternum in front of coxae 0.9 times shorter than diameter of coxal cavity. Scutellum transverse, two times as wide as long. Elytra elongate, 1.6 times as long as wide at widest part, 1.7 times as long as pronotum, widest near middle; each elytron with three rows of indistinct tubercles, inner row forming a distinct ridge basally; sparsely covered with large deep punctures arranged in slightly irregular rows. Legs long, slender; relative lengths of metatarsomeres 1.0 : 0.5 : 1.0 : 2.0.
Male genitalia with tegmen elongate, widest near middle, basally with very short strut; parameres elongate, 3.5 times longer than wide, apically with long setae (Fig.
Variability in males. BL 9.1–12.5 mm, BW 3.1–4.2 mm. Antennae are either reddish brown, brown or black. There is a gradual morphological variation in the pronotal and elytral tubercles, ranging from the less distinct tubercles in the holotype (Figs
Female. Most characters same as for males. BL 10.0–13.0 mm, BW 3.2–4.5 mm. Body more convex dorsally, with distinct tubercles on pronotum and elytra. Antennae shorter, 0.6–0.7 times as long as body. Elytra 1.7 times as long as wide, 2.0–2.1 times as long as pronotum. Spermatheca sclerotized, slender, elongate, curved, widened at second half, gradually tapered toward apex; sclerotized part of spermathecal duct simple (Fig.
Malaysia: Borneo (Sabah; Fig.
Anexodus sarawakensis Sudre, 1997: 253.
Holotype, male, “E. MALAYSIA: Sarawak / confl. Sun Oyan and / Mujong riv., E. Kapit / 50m, 18.V.1994, # 5 / Löbl & Burckhardt // Holotype // Anexodus / sarawakensis Nov sp. / J. Sudre det. 1996” (
Anexodus sarawakensis is similar to A. syptakovae sp. n. in having uniformly colored habitus and labrum with one row of distinct punctures with setae (Fig.
Holotype (male). BL 9.2 mm, BW 3.1 mm. Body uniformly brown, mouthparts lighter. Body densely clothed with very short brown pubescence. Head about as wide as anterior margin of pronotum. Labrum transverse, with one row of punctures bearing long setae (Fig.
Prothorax as long as wide, laterally with one moderately distinct tubercle; pronotal disc with a pair of distinct tubercles near middle and one median at second half; pronotal tubercles punctate. Prosternum in front of coxae 0.9 times shorter than diameter of coxal cavity. Scutellum transverse, two times as wide as long. Elytra elongate, 1.6 times as long as wide at widest part, 1.8 times as long as pronotum, widest near middle; each elytron with three rows of distinct tubercles (Figs
Lateral habitus of Anexodus spp.: 9 Anexodus aquilus Pascoe, holotype male 10 Anexodus aquilus Pascoe, female (= holotype of Anexodus kuntzeni Kriesche) 11 Anexodus aquilus Pascoe, large male from Trus Madi 12 Anexodus syptakovae sp. n., holotype male 13 Anexodus sarawakensis Sudre, holotype male 14 Anexodus sarawakensis Sudre, paratype female 15 Anexodus tufi sp. n., holotype male 16 Anexodus tufi sp. n., paratype female. Scale bar 5 mm.
Male genitalia with tegmen elongate, widest near middle, basally with short strut; parameres elongate, 2.9 times longer than wide, apically with long setae (Fig.
Female. Most characters same as for males. BL 9.9 mm, BW 3.5 mm. Body more convex dorsally. Antennae slightly shorter than in male, 0.7 times as long as body; length ratio of antennomeres I–III: 6.5 : 2.7 : 1.0. Elytra 1.7 times as long as wide, 2.3 times as long as pronotum. Spermatheca only slightly sclerotized, slender, elongate; sclerotized part of spermathecal duct strongly coiled (Fig.
Malaysia: Borneo (Sarawak: Kapit; Fig.
Holotype, male, [Malaysia] “SARAWAK: / 5th Division / Gn. Mulu NP // Camp 5 / Kerangas // Pitfall / trap // iv. 78, N. M. Collins / B.M.1978-11 // Opsies sp.” (
This species is similar to A. sarawakensis in having uniformly colored habitus and labrum with a row of distinct punctures with setae (Fig.
Holotype (male). BL 8.0 mm, BW 2.5 mm. Body uniformly brown, mouthparts lighter. Body densely clothed with very short brown pubescence. Head slightly wider than anterior margin of pronotum. Labrum transverse, with one row of punctures bearing long setae (Fig.
Prothorax as long as wide, laterally with one indistinct tubercle; pronotal disc with a pair of very indistinct tubercles near middle and one median at second half; pronotal tubercles punctate. Prosternum in front of coxae 0.9 times shorter than diameter of coxal cavity. Scutellum transverse, about two times as wide as long. Elytra elongate, 1.8 times as long as wide at widest part, 2.1 times as long as pronotum, widest near middle; each elytron with three rows of only slightly elevated tubercles (Fig.
Male genitalia with tegmen elongate, widest near middle, basally with short strut; parameres elongate, 2.3 times longer than wide, apically with tufts of short setae (Fig.
Paratype is smaller (BL 7.00 mm, BW 2.3 mm), with reddish brown antennae.
Female unknown.
Malaysia: Borneo (Sarawak: Gn. Mulu NP; Fig.
The specific name is a matronym in honor of Ms. Hana Gabriš Syptáková (Salisov, Czech Republic).
Holotype, male, “BRUNEI, Ulu Temburong NP / Kuala Belalong FSC / 4°32'47.6"N 115°09'27"E / I. H. Tuf leg. II.2013” (UBDC); paratype, female, same data as holotype (
This species is similar to A. aquilus in having yellowish stripes on the dorsal body surface and labrum with the whole surface punctured (Fig.
Anexodus spp.: 17–20 Head, frontal view: 17 Anexodus aquilus Pascoe, holotype male 18 Anexodus sarawakensis Sudre, holotype male 19 Anexodus syptakovae sp. n., holotype male 20 Anexodus tufi sp. n., holotype male 21–27 Antenna: 21 Anexodus aquilus Pascoe, male 22 Anexodus aquilus Pascoe, female 23 Anexodus sarawakensis Sudre, male 24 Anexodus sarawakensis Sudre, female 25 Anexodus syptakovae sp. n., male 26 Anexodus tufi sp. n., male 27 Anexodus tufi sp. n., female. Not to scale.
Holotype (male). BL 7.4 mm, BW 2.3 mm. Body brown with yellowish stripes ranging from vertex through sides of pronotum to basal part of elytra, densely clothed with very short brown pubescence. Head about as wide as anterior margin of pronotum. Labrum transverse, its surface with moderately sparse punctures (Fig.
Prothorax as long as wide, laterally with one distinct tubercle; pronotal disc with a pair of moderately distinct tubercles near middle and one median at second half and one indistinct median at anterior half; pronotal tubercles punctate. Prosternum in front of coxae 0.9 times shorter than diameter of coxal cavity. Scutellum transverse, two times as wide as long. Elytra elongate, 1.8 times as long as wide at widest part, 1.9 times as long as pronotum, widest near middle; each elytron with three rows of distinct, longitudinally elongate tubercles (Figs
Male genitalia with tegmen elongate, widest before middle, basally with short strut; parameres elongate, 3.3 times longer than wide, apically with long setae (Fig.
The male paratype is larger (body length 9.4 mm, body width 2.8 mm).
Female. Most characters same as for males. BL 8.5–10.2 mm, BW 2.7–3.4. Antennae shorter, 0.7 times as long as body, with relatively longer scape and pedicel (length ratio of antennomeres I–III: 3.4–3.6 : 1.7–1.8 : 1.0). Elytra 2.2–2.3 times as long pronotum. Fifth ventrite with apex truncate. Spermatheca sclerotized, slender, elongate, curved, gradually tapered toward apex; sclerotized part of spermathecal duct simple (Fig.
Anexodus species: 28–31 Aedeagus, ventral view: 28 Anexodus aquilus Pascoe, holotype 29 Anexodus sarawakensis Sudre, holotype 30 Anexodus syptakovae sp. n., holotype 31 Anexodus tufi sp. n., holotype 32–34 Spermatheca: 32 Anexodus aquilus Pascoe 33 Anexodus tufi sp. n. 34 Anexodus sarawakensis Sudre. Not to scale.
Brunei (Fig.
This species is named after Mr. I. H. Tuf (
Pantilema Aurivillius, 1911: 196.
Pantilema angustum Aurivillius, 1911.
Pantilema differs from the remaining Bornean Morimopsini by having a slender, narrow, parallel-sided body (body length/width ratio = 3.5), tibial spurs 1-1-2, truncate elytral apex, and tubercles only in the apical half of the elytra (Figs
Body slender, elongate, densely clothed with very short pubescence; coloration brown, with some parts paler, yellowish, antennae and legs reddish-brown (Figs
Head about the same width as anterior margin of pronotum; genae convex at frontal view; frontoclypeus with distinct midline running from interantennal groove to labrum, sparsely punctured; antennal tubercles prominent with narrow, deep depression in between; antennal cavities opened dorsally; anterior margin of anteclypeus shallowly emarginate, with sparse long yellowish semi-erect setae. Labrum free, transverse, glabrous, with a row of distinct punctures and sparsely and irregularly distributed additional less distinct punctures, with sparse long semi-erect setae (Fig.
Prothorax about as long as wide, subparallel-sided at anterior half, widest slightly medially, then gradually narrowed towards posterior margin, laterally with one very weakly developed tubercle; pronotal disc weakly convex, sparsely covered with deep punctures, not smooth, without tubercles (Fig.
Abdomen with five visible ventrites, first ventrite (excluding intercoxal process) almost 1.5 times longer than second; intercoxal process subparallel-sided basally, narrowed and broadly rounded apically. Fifth ventrite with apex truncate, margin with sparse semi-erect pubescence. Male genitalia with tegmen elongate, widest at apical third, basally with long strut; parameres moderately long, setose apically. Penis relatively long, with dorsal struts diverged from about two fifths of penis length (Fig.
Pantilema angustum Aurivillius, 1911: 196.
Holotype, male, “Samarahan / June 1906 [handwritten] // Type. //
Holotype (male). BL 10.6 mm, BW 3.1 mm. Body brown, with antennae and legs reddish-brown, elytral apex and two median spots near anterior and posterior pronotal margins yellowish, mouthparts lighter (Figs
Prothorax 1.1 times as long as wide, laterally with one very weakly developed tubercle; pronotal disc without tubercles (Fig.
Male genitalia with tegmen elongate, widest at apical third, basally with long strut; parameres moderately long. Penis long, apically broadly rounded; dorsal struts diverged from about two fifths of penis length (Fig.
Malaysia: Borneo (Sarawak: Samarahan; Fig.
1 | Body uniformly brown (Figs |
2 |
– | Body brown with yellowish stripes dorsally (Figs |
3 |
2 | Body length 7.0–8.0 mm; length ratio of antennomeres I–III 4.4–4.6 : 1.7–1.9 : 1.0 (Fig. |
A. syptakovae sp. n. |
– | Body length 9.2–9.9 mm; length ratio of antennomeres I–III 6.3–6.5 : 2.5–2.7 : 1.0 (Figs |
A. sarawakensis Sudre |
3 | Apex of the scape with distinct lateral hook-shaped projection (Figs |
A. aquilus Pascoe |
– | Apex of the scape simple (Figs |
A. tufi sp. n. |
Genus Anexodus Pascoe, 1886
A. aquilus Pascoe, 1886 (Malaysia: Sabah; Fig.
A. sarawakensis Sudre, 1997 (Malaysia: Sarawak; Fig.
A. syptakovae sp. n. (Malaysia: Sarawak; Fig.
A. tufi sp. n. (Brunei; Fig.
Genus Borneostyrax Gabriš, Kundrata & Trnka, 2016
B. cristatus Gabriš, Kundrata & Trnka, 2016 (Malaysia: Sabah; Fig.
Genus Dolichostyrax Aurivillius, 1911
D. longipes Aurivillius, 1913 (Malaysia: Sabah; Fig.
D. moultoni Aurivillius, 1911 (Malaysia: Sarawak; Fig.
Genus Eurystyrax Gabriš, Kundrata & Trnka, 2016
E. nemethi Gabriš, Kundrata & Trnka, 2016 (Malaysia: Sabah; Fig.
Genus Microdolichostyrax Gabriš, Kundrata & Trnka, 2016
M. hefferni Gabriš, Kundrata & Trnka, 2016 (Malaysia: Sabah; Fig.
M. minutus Gabriš, Kundrata & Trnka, 2016 (Malaysia: Sabah; Fig.
Genus Pantilema Aurivillius, 1911
P. angustum Aurivillius, 1911 (Malaysia: Sarawak; Fig.
In 2013, two Czech universities (Palacky University in Olomouc, University of Ostrava) and the Universiti Brunei Darussalam established a collaboration which resulted in the biodiversity survey of the Ulu Temburong National Park in Brunei (
We thank M. Barclay, M. Geiser (