Research Article |
Corresponding author: Alexander Riedel ( riedel@smnk.de ) Academic editor: Miguel Alonso-Zarazaga
© 2021 Raden Pramesa Narakusumo, Alexander Riedel.
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:
Narakusumo RP, Riedel A (2021) Twenty-eight new species of Trigonopterus Fauvel (Coleoptera, Curculionidae) from Central Sulawesi. ZooKeys 1065: 29-79. https://doi.org/10.3897/zookeys.1065.71680
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Here we present 28 new species of Trigonopterus from Central Sulawesi, mostly from Mt Dako and Mt Pompangeo: Trigonopterus acutus sp. nov., T. ancora sp. nov., T. arcanus sp. nov., T. corona sp. nov., T. dakoensis sp. nov., T. daun sp. nov., T. ewok sp. nov., T. gundala sp. nov., T. hoppla sp. nov., T. kakimerah sp. nov., T. katopasensis sp. nov., T. matakensis sp. nov., T. moduai sp. nov., T. mons sp. nov., T. paramoduai sp. nov. T. pomberimbensis sp. nov., T. pompangeensis sp. nov., T. puspoi sp. nov., T. rosichoni sp. nov., T. rubidus sp. nov., T. sarinoi sp. nov., T. sutrisnoi sp. nov., T. tanah sp. nov., T. tejokusumoi sp. nov., T. toboliensis sp. nov., T. tolitoliensis sp. nov., T. tounaensis sp. nov., T. unyil sp. nov. This fills important areas of distribution and brings the number of Trigonopterus species recorded from Sulawesi to 132.
Celebes, conservation, cox1, Cryptorhynchinae, DNA barcoding, endemism, hyperdiverse, integrative taxonomy, morphology, Southeast Asia, turbo-taxonomy, Wallacea
Trigonopterus is a hyperdiverse genus of flightless hidden-snout weevils (Cryptorhynchinae) ranging over the Indo-Australian-Melanesian archipelago. It originated in Northern Australia and rapidly diversified in New Guinea (
Sulawesi is geologically complex (
This study is based on 866 specimens from Central Sulawesi Province. Holotypes were selected from 197 specimens for which the cox1 gene had been sequenced. DNA was extracted nondestructively as described by
The methods applied for DNA sequencing and sequence analysis are the same as described by
The closest relatives of Central Sulawesi species were identified by creating an alignment of 773 cox1 sequences representing ca. 185 species and generating a maximum likelihood reconstruction using the program IQTREE (
Trigonopterus Fauvel, 1862 Type species: Trigonopterus insignis Fauvel, 1862, by monotypy.
Fully apterous genus of Cryptorhynchinae. Length 1.5–6.0 mm. Rostrum in repose not reaching center of mesocoxa. Scutellar shield completely absent externally. Mesothoracic receptacle deep, posteriorly closed. Metanepisternum completely absent externally. Elytra with nine striae (sometimes superficially effaced). Tarsal claws minute. Usually body largely unclothed, without dense vestiture. For additional information, see http://species-id.net/wiki/Trigonopterus.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation ca. 1700–1800 m.
On foliage in montane forest.
The species name is the Latin adjective acutus -a -um (pointed, acute) and refers both to the elytral shape and the apex of the penis.
Trigonopterus acutus sp. nov. was coded as “Trigonopterus sp. 1207”. This species belongs to the T. tatorensis-group. It is closely related to T. daun sp. nov., from which it can be distinguished by the pointed apex of the penis.
Holotype, male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation 970–1400 m.
On foliage and in leaf litter in montane forests.
This epithet is the Latin noun ancora (anchor) in apposition and refers to the sclerite in the male transfer apparatus.
Trigonopterus ancora sp. nov. was coded as “Trigonopterus sp. 1114” (
Holotype, male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation 830–1400 m.
On foliage in montane forest.
This epithet is based on the Latin adjective arcanus -a, -um (hidden) and it refers to its close similarity to its sibling species T. ovatulus Riedel and T. pseudovatulus Riedel.
Trigonopterus arcanus sp. nov. was coded as “Trigonopterus sp. 1187”. The species belongs to the T. ovatulus-group. It is closely related to T. pseudovatulus Riedel from which it can be distinguished by the absence of a metatibial supra-uncal tooth. Furthermore, it differs by 12.7–13.8% p-distances of its cox1 sequence.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation ca. 1100–1200 m.
On foliage in montane forest.
This epithet refers to the Corona virus (Sars-Cov2). The global pandemic led to the cancellation of field work, and a focus on this and other manuscripts. It is a noun in apposition.
Trigonopterus corona sp. nov. was coded as “Trigonopterus sp. 1235”. This species belongs to the T. fulvicornis-group and is related to T. seticnemis Riedel, from which it can be easily distinguished by the ferruginous elytra.
Holotype, male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation 1030–1200 m.
In leaf litter of mountain forest.
This epithet is a Latinized adjective based on Mt Dako.
Trigonopterus dakoensis sp. nov. was coded as “Trigonopterus sp. 1189”. The species belongs to the T. manadensis-group. It is related to T. manadensis Riedel, from which it differs by the peculiar morphology of the penis and by the longitudinal rows of coarse punctures on the pronotum. The p-distance of their cox1 sequences is 14.5–14.8%.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation 970–1200 m.
On foliage in montane forest.
This epithet is the Indonesian word for “leaf” and a noun in apposition. It refers to the species´ lifestyle on foliage.
Trigonopterus daun sp. nov. was coded as “Trigonopterus sp. 1116” (
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Pompangeo). Elevation ca. 1900–2000 m.
In leaf litter of montane forest.
This epithet is a noun in apposition based on the fictional character of small bear-like creatures from Star Wars VI movie.
Trigonopterus ewok sp. nov. was coded as “Trigonopterus sp. 1188” (
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation ca. 1900–2200 m.
On foliage in montane forest.
This epithet is a noun in apposition based on the fictional character of Indonesian comic superhero “Gundala, Son of Thunder”. The black and ferruginous colors of this species resemble Gundala’s movie costume.
Trigonopterus gundala sp. nov. was coded as “Trigonopterus sp. 1194”. It belongs to the T. satyrus-group and is closely related to T. mons sp. nov., but differs by the deeply striate elytra and a 8.2–8.7% cox1 p-distance.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation 1300–1700 m.
On foliage in montane forest.
This epithet is based on the German word “Hoppla”, an exclamation of surprise, comparable to the English “whoops”. It is to be treated as a noun in apposition.
Trigonopterus hoppla sp. nov. was coded as “Trigonopterus sp. 1232”. This species belongs to the T. tatorensis-group. It is closely related to T. daun sp. nov., from which it can be distinguished by the pointed apex of the penis and a cox1 p-distance of 7.8%. The marked difference in sculpture between holotype and the single paratype is remarkable, and would usually indicate a separate species. However, genital morphology and cox1 sequence of both specimens are almost identical, so either the coarse sculpture of the holotype, or the smooth sculpture of the paratype may be an aberration. Additional specimens are needed to clarify this matter.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Pompangeo). Elevation 1800–1900 m.
On foliage in montane forest.
This epithet is the Indonesian term for “red legs”. It is a noun in apposition.
Trigonopterus kakimerah sp. nov. was coded as “Trigonopterus sp. 1202”. This species may belong to the T. fulvicornis-group. The cox1 p-distance to other known species is above 14%.
Holotype. Male (Fig.
C-Sulawesi Prov. (Mt Katopasa). Elevation ca. 1380 m.
On foliage in montane forest.
This epithet is a Latinized adjective based on Mt Katopasa.
Trigonopterus katopasensis sp. nov. was coded as “Trigonopterus sp. 1190”. This species presumably belongs to the T. barbipes-group. From T. barbipes Riedel it differs by less distinct body sculpture, the structure of the penis, and a 18.5% cox1 p-distance.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Pompangeo). Elevation 1800–2000 m.
On foliage and in leaf litter in montane forests.
This epithet is a Latinized adjective based on Matako village.
Trigonopterus matakensis sp. nov. was coded as “Trigonopterus sp. 1197”. This species belongs to the T. ovalipunctatus-group. It is closely related to T. ovalipunctatus Riedel, but differs by the subrotund shape of the supporting sclerites of the transfer apparatus and a 9.6–10.3% cox1 p-distance.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation ca. 1700–1800 m.
On foliage in montane forest.
This epithet is a noun in apposition based on “Moduai”, a folk dance of people from Toli-Toli Regency.
Trigonopterus moduai, sp. nov. was coded as “Trigonopterus sp. 1201” and belongs to the T. arachnobas-group. It is very close to T. paramoduai sp. nov. (3.20–4.11% cox1 p-distance) but can be distinguished by the darker elytral color, a slightly shorter rostrum, and the much longer flagellum of the male genital.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation 1700–1900 m.
On foliage in montane forest.
This epithet is the Latin noun mons (mountain) in apposition.
Trigonopterus mons sp. nov. was coded as “Trigonopterus sp. 1195”. It belongs to the T. satyrus-group and is closely related to T. gundala sp. nov., which differs by the deeply striate elytra and a 8.2–8.7% cox1 p-distance.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation 1900–2200 m.
On foliage in montane forest.
This epithet is based on the combination of the Greek prefix para- (next to; near by) and the sibling species Trigonopterus moduai, sp. nov..
Trigonopterus paramoduai sp. nov. was coded as “Trigonopterus sp. 1233” and belongs to the T. arachnobas-group. It is very close to T. moduai sp. nov. (3.20–4.11% cox1 p-distance) from which it can be distinguished by the ferruginous elytral color, a slightly longer rostrum, and the shorter flagellum of the male genital.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Pompangeo). Elevation 1900–2000 m.
In leaf litter of montane forest.
This epithet is a Latinized adjective based on Pomberimbe hill.
Trigonopterus pomberimbensis sp. nov. was coded as “Trigonopterus sp. 1191”. This species belongs to the T. barbipes-group. It is closely related to T. viduus Riedel, which differs by weakly impressed elytral striae and 19.7–21.2% cox1 p-distance.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Pompangeo). Elevation 1800–2000 m.
On foliage in montane forest.
This epithet is a Latinized adjective based on Mt Pompangeo.
Trigonopterus pompangeensis sp. nov. was coded as “Trigonopterus sp. 1198”. This species belongs to the T. ovalipunctatus-group. It is closely related to T. ovalipunctatus Riedel, but differs by a more densely punctate pronotum, the apically extended and converging penis, and 7.2–9.7% cox1 p-distance.
Holotype
(
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation 830–1250 m.
On foliage in montane forest.
This species is named in honor of Saleh Poespo, grandfather of the first author, and for his pioneering animal husbandry science in Indonesia. An invariable genitive.
Trigonopterus puspoi sp. nov. was coded as “Trigonopterus sp. 1113” (
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation ca. 1400–1750 m.
On foliage in montane forest.
This species is named in honor of Rosichon Ubaidillah, curator and researcher of Hymenoptera at
Trigonopterus rosichoni sp. nov. was coded as “Trigonopterus sp. 1193”. It belongs to the T. satyrus-group and is closely related to T. ancora sp. nov. from which it differs by the shape of the transfer apparatus and 9.4–9.6% p-distance of its cox1 sequence.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation 1700–1900 m.
On foliage in montane forest.
This epithet is the Latin adjective rubidus, -a, -um (reddish) referring to the elytral color.
Trigonopterus rubidus sp. nov. was coded as “Trigonopterus sp. 1199”. This species belongs to the T. tatorensis-group. It is closely related to T. tatorensis Riedel, from which it differs by denser pronotal punctures, its reddish elytral color and 9.5–9.9% cox1 p-distance.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Gn. Torompupu). Elevation ca. 800 m.
In leaf litter.
This species is named in honor of Sarino, technician working at the Coleoptera collection of LIPI-
Trigonopterus sarinoi sp. nov. was coded as “Trigonopterus sp. 1208”. This species belongs to the T. lampros-group. It is closely related to T. yoda Riedel, which differs by its black-bronze elytral color and a 19.3% cox1 p-distance.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation ca. 720–970 m.
On foliage in lower montane forest.
This epithet is named in honor of Hari Sutrisno, curator of moths and researcher at
Trigonopterus sutrisnoi sp. nov. was coded as “Trigonopterus sp. 1206”. This species belongs to the T. toraja-group. It is related to T. toboliensis sp. nov., which differs by its lateral extensions of the penis and a cox1 p-distance of 15.3–15.5%.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation ca. 1100–1200 m.
In leaf litter of montane forest.
This epithet is the Indonesian word for “soil” and a noun in apposition. It refers to the species´ lifestyle on the ground among leaf litter.
Trigonopterus tanah sp. nov. was coded as “Trigonopterus sp. 1234”. It is closely related to T. darwini Riedel, from which it can be distinguished by its coarser sculpture, and the subparallel body of the penis. The cox1 p-distance of both species is 10.8%.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Pompangeo). Elevation 1800–2000 m.
On foliage in montane forest.
This species is named in honor of Slamet Tedjokoesoemo, pioneer of veterinary science in Indonesia and grandfather of the first author. An invariable genitive.
Trigonopterus tejokusumoi sp. nov. was coded as “Trigonopterus sp. 1200”. This species belongs to the T. barbipes-group. It is most closely related to T. barbipes Riedel, but differs by smaller and more irregular elytral punctures, a peculiar obtuse apex of the penis and a 16% cox1 p-distance.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Palu, Palolo). Elevation 700–850 m.
On foliage in montane forest.
This epithet is a Latinized adjective based on Toboli village.
Trigonopterus toboliensis sp. nov. was coded as “Trigonopterus sp. 1192”. This species belongs to the T. toraja-group. It is closely related to T. ampanensis Riedel from which it differs by the shape and position of the lateral extensions of the penis and a cox1 p-distance of 14.5–14.7%.
Holotype. Male (Fig.
Holotype
(
C-Sulawesi Prov. (Mt Dako). Elevation 830–1250 m.
On foliage in montane forests.
This epithet is a Latinized adjective based on Toli-Toli regency.
Trigonopterus tolitoliensis sp. nov. was coded as “Trigonopterus sp. 1186”. The species belongs to the T. palopensis-group. It is closely related to T. puspoi sp. nov. from which it differs by the setose brushes on the dorsal surface of the penis, the scaling of the metatibia, and a 16.4–18.1% p-distance of its cox1 sequence.
Holotype. Male (Fig.
C-Sulawesi Prov. (Mt Katopasa). Elevation ca. 1380 m.
On foliage in montane forest.
This epithet is a Latinized adjective based on the Indonesian abbreviation of Tojo Una-Una “Touna” and refers to the type locality.
Trigonopterus tounensis, sp. nov. was coded as “Trigonopterus sp. 1115” (
Holotype. Male (Fig.
Map of Trigonopterus records across Sulawesi and adjacent islands. Blue dots from
Holotype
(
C-Sulawesi Prov. (Mt Pompangeo). Elevation ca. 1900–2000 m.
In leaf litter of montane forest.
This epithet is a noun in apposition based on the Indonesian hand puppet character from “Si Unyil” TV series.
Trigonopterus unyil sp. nov. was coded as “Trigonopterus sp. 1196”. Presumably, this species belongs to the T. nanus-group.
Relatively short field trips to two mountains in Sulawesi (i.e. five days at Mt Pompangeo and 14 days at Mt Dako) resulted in the discovery of 25 out of 28 Trigonopterus species newly described herein. Only a single species, T. toboliensis sp. nov. from Palu, respectively Palolo had been collected earlier. This exemplifies what great positive impact dedicated field work can have to species-discovery of neglected arthropod taxa, especially in montane areas. While the more conspicuous species of birds, mammals, butterflies, larger beetles or plants may have been collected on earlier surveys (
It is noteworthy that in some localities pairs of sister species have been discovered: T. moduai sp. nov., and T. paramoduai sp. nov. are both found in narrowly separated elevation zones of Mt Dako. Although genetically very close (3.20–4.11% cox1 p-distance) they are very distinct morphologically. Other such species pairs are morphologically very hard or impossible to separate, but very divergent genetically, i.e. T. matakensis sp. nov. and T. pompangeensis sp. nov. from Mt Pompangeo or the allopatric T. ovatulus Riedel and T. pseudovatulus Riedel from mountains north of Lake Poso (
Mt Dako is a nature reserve, and forests above 800 m are largely intact. On the other hand, Mt Pompangeo is without any conservation status and has been logged extensively between 1970 m and 2000 m. Patches of rainforest remaining in the steeper areas are still at risk being affected by regular forest fires. Both Mt Dako and Mt Pompangeo harbour endemic Trigonopterus species, and presumably additional ones could be discovered if longer field trips are conducted in the remaining forest patches. These forest patches among the Sulawesi rainforest still hold the largely unknown diversity of Trigonopterus and other arthropod species. They should be of greater concern to conservation despite or rather because of their fragmentation.
subgenus Mimidotasia Voss: T. pauper Riedel, T. luwukensis Riedel, T. selayarensis Riedel, T. wangiwangiensis Riedel.
T. abnormis-group: T. abnormis Riedel, T. kolakensis Riedel
T. analis-group: T. analis Riedel, T. pagaranganensis Riedel, T. pseudanalis Riedel
T. arachnobas-group: T. arachnobas Riedel, T. darwini Riedel, T. gracilipes Riedel, T. idefix Riedel, T. moduai sp. nov., T. paramoduai sp. nov., T. pumilus Riedel, T. rudis Riedel, T. tanah sp. nov., T. tenuipes Riedel.
T. barbipes-group: T. barbipes Riedel, T. ejaculatorius Riedel, T. katopasensis sp. nov., T. pomberimbensis sp. nov., T. tejokusumoi sp. nov., T. vicinus Riedel, T. viduus Riedel.
T. bornensis-group: T. rotundatus Riedel.
T. collaris-group: T. collaris Riedel, T. klabatensis Riedel, T. paracollaris Riedel.
T. crenulatus-group: T. costatulus Riedel, T. crenulatus Riedel, T. humilis Riedel, T. squalidulus Riedel.
T. curtus-group: T. procurtus Riedel, T. pseudosimulans Riedel, T. scitulus Riedel.
T. dimorphus-group: T. reticulatus Riedel, T. scabripes Riedel, T. pendolensis Riedel.
T. fulvicornis-group: T. celebensis Riedel, T. corona sp. nov., T. fulvicornis (Pascoe, 1885), T. kakimerah sp. nov., T. pseudofulvicornis Riedel, T. seticnemis Riedel.
T. honestus-group: T. inhonestus Riedel.
T. incendium-group: T. incendium Riedel.
T. impressicollis-group: T. adspersus Riedel, T. castaneipennis Riedel, T. ewok sp. nov., T. impressicollis Riedel, T. mesai Riedel, T. serripes Riedel, T. suturatus Riedel.
T. laevigatus-group: T. invalidus Riedel, T. jasminae Riedel, T. laevigatus Riedel.
T. lampros-group: T. artemis Riedel, T. carinirostris Riedel, T. curvipes Riedel, T. lampros Riedel, T. sarinoi sp. nov., T. yoda Riedel.
T. manadensis-group: T. ambangensis Riedel, T. armipes Riedel, T. cirripes Riedel, T. dakoensis sp. nov., T. heberti Riedel, T. mahawuensis Riedel, T. manadensis Riedel, T. modoindingensis Riedel, T. pseudomanadensis Riedel, T. volcanorum Riedel.
T. minahassae-group: T. indigenus Riedel, T. minahassae Riedel, T. prismae Riedel, T. sampunensis Riedel, T. silvicola Riedel.
T. nanus-group: T. hirsutus Riedel, T. nanus Riedel, T. unyil sp. nov..
T. nitidulus-group: T. cricki Riedel, T. nitidulus Riedel.
T. ovalipunctatus-group: T. lompobattangensis Riedel, T. matakensis sp. nov., T. ovalipunctatus Riedel, T. pompangeensis sp. nov., T. pseudovalipunctatus Riedel.
T. ovatulus-group: T. arcanus sp. nov., T. ovatulus Riedel, T. pseudovatulus Riedel.
T. palopensis-group: T. asterix Riedel, T. fuscipes Riedel, T. kotamobagensis Riedel, T. latipennis Riedel, T. matalibaruensis Riedel, T. moatensis Riedel, T. palopensis Riedel, T. puspoi sp. nov., T. rhombiformis Riedel, T. rufipes Riedel, T. tolitoliensis sp. nov..
T. politus-group: T. allotopus Riedel, T. pseudallotopus Riedel.
T. posoensis-group: T. obelix Riedel, T. posoensis Riedel, T. tounensis sp. nov..
T. relictus-group: T. mangkutanensis Riedel
T. rotundulus-group: T. rotundulus Riedel, T. watsoni Riedel
T. saltator-group: T. bonthainensis Riedel
T. satyrus-group: T. ancora sp. nov., T. gundala sp. nov., T. mons sp. nov., T. rosichoni sp. nov., T. satyrus Riedel.
T. sampuragensis-group: T. sampuragensis Riedel.
T. tatorensis-group: T. acutus sp. nov., T. daun sp. nov., T. hoppla sp. nov., T. hypocrita Riedel, T. incognitus Riedel, T. rubidus sp. nov., T. sulawesiensis Riedel, T. tatorensis Riedel, T. tomohonensis Riedel.
T. toraja-group: T. ampanensis Riedel, T. rantepao Riedel, T. scaphiformis Riedel, T. sutrisnoi sp. nov., T. toboliensis sp. nov., T. toraja Riedel.
We thank LIPI (Indonesian Institute of Sciences), RISTEK (Ministry of State for Research and Technology, Indonesia) and the Indonesian Department of Forestry for providing relevant permits. The field work in Indonesia would not have been possible without the generous hospitality and help of many local people from Matako, Enrekan, Teluk Bone, Kinapasan villages and we thank all of them very warmly. Thanks to Anang Setiawan Achmadi (