Ninety-eight new species of Trigonopterus weevils from Sundaland and the Lesser Sunda Islands

Alexander Riedel; Rene Tänzler; Michael Balke; Cahyo Rahmadi; Yayuk R Suhardjono

doi:10.3897/zookeys.467.8206

. 2014 Dec 22;(467):1–162. doi: 10.3897/zookeys.467.8206

Ninety-eight new species of Trigonopterus weevils from Sundaland and the Lesser Sunda Islands

Alexander Riedel ¹, Rene Tänzler ², Michael Balke ^2,³, Cahyo Rahmadi ⁴, Yayuk R Suhardjono ⁴

PMCID: PMC4296478 PMID: 25610340

Abstract Abstract

The genus Trigonopterus Fauvel, 1862 is highly diverse in Melanesia. Only one species, Trigonopterus amphoralis Marshall, 1925 was so far recorded West of Wallace’s Line (Eastern Sumatra). Based on focused field-work the fauna from Sundaland (Sumatra, Java, Bali, Palawan) and the Lesser Sunda Islands (Lombok, Sumbawa, Flores) is here revised. We redescribe Trigonopterus amphoralis Marshall and describe an additional 98 new species: Trigonopterus acuminatus sp. n., Trigonopterus aeneomicans sp. n., Trigonopterus alaspurwensis sp. n., Trigonopterus allopatricus sp. n., Trigonopterus allotopus sp. n., Trigonopterus angulicollis sp. n., Trigonopterus argopurensis sp. n., Trigonopterus arjunensis sp. n., Trigonopterus asper sp. n., Trigonopterus attenboroughi sp. n., Trigonopterus baliensis sp. n., Trigonopterus batukarensis sp. n., Trigonopterus bawangensis sp. n., Trigonopterus binodulus sp. n., Trigonopterus bornensis sp. n., Trigonopterus cahyoi sp. n., Trigonopterus costipennis sp. n., Trigonopterus cuprescens sp. n., Trigonopterus cupreus sp. n., Trigonopterus dacrycarpi sp. n., Trigonopterus delapan sp. n., Trigonopterus dentipes sp. n., Trigonopterus diengensis sp. n., Trigonopterus dimorphus sp. n., Trigonopterus disruptus sp. n., Trigonopterus dua sp. n., Trigonopterus duabelas sp. n., Trigonopterus echinatus sp. n., Trigonopterus empat sp. n., Trigonopterus enam sp. n., Trigonopterus fissitarsis sp. n., Trigonopterus florensis sp. n., Trigonopterus foveatus sp. n., Trigonopterus fulgidus sp. n., Trigonopterus gedensis sp. n., Trigonopterus halimunensis sp. n., Trigonopterus honjensis sp. n., Trigonopterus ijensis sp. n., Trigonopterus javensis sp. n., Trigonopterus kalimantanensis sp. n., Trigonopterus kintamanensis sp. n., Trigonopterus klatakanensis sp. n., Trigonopterus lampungensis sp. n., Trigonopterus latipes sp. n., Trigonopterus lima sp. n., Trigonopterus lombokensis sp. n., Trigonopterus merubetirensis sp. n., Trigonopterus mesehensis sp. n., Trigonopterus micans sp. n., Trigonopterus misellus sp. n., Trigonopterus palawanensis sp. n., Trigonopterus pangandaranensis sp. n., Trigonopterus paraflorensis sp. n., Trigonopterus pararugosus sp. n., Trigonopterus parasumbawensis sp. n., Trigonopterus pauxillus sp. n., Trigonopterus payungensis sp. n., Trigonopterus porcatus sp. n., Trigonopterus pseudoflorensis sp. n., Trigonopterus pseudosumbawensis sp. n., Trigonopterus punctatoseriatus sp. n., Trigonopterus ranakensis sp. n., Trigonopterus relictus sp. n., Trigonopterus rinjaniensis sp. n., Trigonopterus roensis sp. n., Trigonopterus rugosostriatus sp. n., Trigonopterus rugosus sp. n., Trigonopterus rutengensis sp. n., Trigonopterus saltator sp. n., Trigonopterus santubongensis sp. n., Trigonopterus sasak sp. n., Trigonopterus satu sp. n., Trigonopterus schulzi sp. n., Trigonopterus sebelas sp. n., Trigonopterus sembilan sp. n., Trigonopterus sepuluh sp. n., Trigonopterus seriatus sp. n., Trigonopterus serratifemur sp. n., Trigonopterus setifer sp. n., Trigonopterus silvestris sp. n., Trigonopterus singkawangensis sp. n., Trigonopterus singularis sp. n., Trigonopterus sinuatus sp. n., Trigonopterus squalidus sp. n., Trigonopterus sumatrensis sp. n., Trigonopterus sumbawensis sp. n., Trigonopterus sundaicus sp. n., Trigonopterus suturalis sp. n., Trigonopterus syarbis sp. n., Trigonopterus telagensis sp. n., Trigonopterus tepalensis sp. n., Trigonopterus tiga sp. n., Trigonopterus trigonopterus sp. n., Trigonopterus tujuh sp. n., Trigonopterus ujungkulonensis sp. n., Trigonopterus variolosus sp. n., Trigonopterus vulcanicus sp. n., Trigonopterus wallacei sp. n.. All new species are authored by the taxonomist-in-charge, Alexander Riedel. Most species belong to the litter fauna of primary wet evergreen forests. This habitat has become highly fragmented in the study area and many of its remnants harbor endemic species. Conservation measures should be intensified, especially in smaller and less famous sites to minimize the number of species threatened by extinction.

Keywords: Southeast Asia, integrative taxonomy, turbo-taxonomy, weevils, hyperdiverse, morphology, cox1, DNA barcoding, conservation, endemism, Coleoptera, Curculionidae, Cryptorhynchinae

Introduction

Trigonopterus Fauvel, a genus of flightless weevils placed in the subfamily Cryptorhynchinae of Curculionidae (Alonso-Zarazaga and Lyal 1999), is hyperdiverse in the Papuan region (Riedel et al. 2010; Tänzler et al. 2012). However, only a few species have been described from west of the Moluccas to date. Besides three species from the Philippines [Trigonopterus bakeri (Hustache, 1925), Trigonopterus paucisquamosus (Heller, 1915), Trigonopterus semirubrus (Hustache, 1925)] and one from Sulawesi [Trigonopterus fulvicornis (Pascoe, 1885)], there is only one more species described from East Sumatra [Trigonopterus amphoralis (Marshall)]. The conspicuous gap in distribution is here revealed as a sampling artifact and will be remedied with the present publication.

The area covered herein includes parts of Sundaland (i.e. Sumatra, Java, Borneo, and the Philippine island of Palawan) and the Lesser Sunda Islands. No species of Trigonopterus is known to us from the Malay Peninsula and the western and central parts of Sumatra. The intensive field work for this study largely focused on East Sumatra, Java and the Lesser Sunda Islands, while material available from only a few localities in Borneo and Palawan could be included. Thus, many more species can be expected from these islands. The fauna of Sulawesi will be treated in a separate publication.

We have previously established that Trigonopterus weevils are suitable for accelerated taxonomic study combining morphology and the DNA barcoding approach using mitochondrial cox1 data (Riedel et al. 2013a, b; Tänzler et al. 2012); herein we will use the same strategy to provide “faces and names” to an additional 98 undescribed species. While in our earlier study (Riedel et al. 2013b) we tried to name only selected representatives covering all species groups, we now revise all species of a focal region. This newly established taxonomic foundation will be of immediate use to our studies on the evolution and biogeography of Trigonopterus (e.g. Tänzler et al. 2014).

Materials and methods

This study is based on > 4,000 specimens of Trigonopterus, collected specifically for a project on this genus. Unless otherwise stated, specimens were collected by sifting the litter of primary forests with subsequent extraction by hand or, more efficiently, using eclectors (Besuchet et al. 1987). Holotypes were selected from the 703 sequenced specimens; their DNA had been extracted nondestructively as described by Riedel et al. (2010). The genitalia of most specimens did not require maceration after DNA-extraction; they could be directly stained with an alcoholic Chlorazol Black solution and stored in glycerol in microvials attached to the pin of the specimens. Genitalia of collection specimens or specimens whose abdominal muscle tissue was not sufficiently digested after DNA extraction were macerated with 10% KOH and rinsed in diluted acetic acid before staining. Illustrations of habitus and genitalia were prepared from holotypes. Finally, type series were supplemented with specimens stored in ethanol and older material from the dry collection. As always the case in paratypes, there is a chance that some of these are incorrectly assigned; this is especially true for specimens without sequence data as an identification based on external morphological characters is more prone to error than an identification based on a cox1 sequence (Tänzler et al. 2012). Several different species belonging to the Trigonopterus relictus-group occur sympatrically in Sumbawa and Flores. Since it was not possible to separate these sibling species without the extraction of male genitalia and / or their sequence data, a substantial number of specimens were not included in any type series. Type depositories are cited using the following codens:

ARC.

MZB.

SMNK.

ZSM.

The methods applied for DNA sequencing and sequence analysis are described by Riedel et al. (2010) and Tänzler et al. (2012). Morphological descriptions are limited to major diagnostic characters as outlined by Riedel et al. (2013a, b). Negative character states (i.e. the absence of a character) are only mentioned explicitly where it appears appropriate. For example, there are some species with scale-bearing punctures in which the insertion of the scale has shifted from the center to the margin of the puncture. In these cases, the character is described, but for the majority of species where it is placed in the center it is not mentioned. Common practice would require stating explicitly “scale inserted in center of puncture”. Although formally accurate, in groups comprising hundreds of species this leads to inflated descriptions that distract the reader from the important information by enumerating the absence of rare character states.

Morphological terminology follows Beutel and Leschen (2005) and Leschen et al. (2009), i.e. the terms “mesoventrite” / “metaventrite” are used instead of “mesosternite” / “metasternite”, and “mesanepisternum” / “metanepisternum” instead of “mesepisternum” / “metepisternum”; “penis” is used instead of “aedeagus” as the tegmen is usually without useful characters in Trigonopterus and therefore omitted from species descriptions. Descriptions were prepared using a Leica MZ16 dissecting microscope and a fluorescent desk lamp for illumination. Measurements were taken with the help of an ocular grid. The length of the body was measured in dorsal aspect from the elytral apex to the front of the pronotum. Width of elytra was measured between the humeri at their greatest extent and across both elytra. Legs were described in an idealized laterally extended position; there is a dorsal / ventral and an anterior / posterior surface. Habitus illustrations were compiled using the Automontage© software (Syncroscopy, Cambridge, UK) with a JVC KY70 camera (JVC Professional Products) adapted to a Leica Z6 APO (Leica Microsystems, Wetzlar, Germany). Photographic illustrations of genitalia were made using the same software / camera combination adapted to a Zeiss Imager microscope, and for this purpose the genitalia were embedded in glycerol gelatin as described by Riedel (2005). Genitalia were photographed with their longitudinal axis somewhat lifted anteriorly, to adequately illustrate structures of the curved down apex. All photographs were enhanced using Adobe Photoshop CS2. However, care was taken not to obscure or alter any features of the specimens illustrated. Sequence data were submitted to the European Molecular Biology Laboratory (EMBL), and the accession numbers are provided under each species e.g. as “(EMBL # FN429236)”.

Systematics

Trigonopterus

Fauvel, 1862

Type species.

Trigonopterus insignis Fauvel, 1862, by monotypy.

Diagnosis.

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 9 striae (sometimes superficially effaced). Tarsal claws minute. Usually body largely unclothed, without dense vestiture. For additional information, see http://species-id.net/wiki/Trigonopterus.

Trigonopterus angulicollis Riedel, sp. n. was coded as “Trigonopterus sp. 314” by Tänzler et al. (2014).

7. Trigonopterus amphoralis

Marshall

Trigonopterus amphoralis Marshall (1925): p. 217–218.

Diagnostic description.

Holotype, male (Fig. 95a). Length 2.27 mm. Color of antennae and legs ferruginous; remainder black. Body subovate, in dorsal aspect and in profile with weak constriction between pronotum and elytron. Rostrum with median ridge and pair of indistinct submedian ridges, intervening furrows each with sparse row of mesad directed setae; anteriorly scabrous; epistome with transverse ridge. Pronotum coarsely punctate, interspaces longitudinally rugose, with median costa; with sparse, subrecumbent scales. Elytra with striae deeply impressed; each puncture containing small suberect seta; subbasally with sparse, slender, suberect scales; intervals weakly costate, almost flat, subglabrous, sutural interval with sparse coarse punctures; in basal half incised suture bordered by row of punctures, partly overgrown by sutural interval. Femora with simple, crenate anteroventral ridge. Metafemur subapically with stridulatory patch. Metatibia apically with uncus, without premucro. Abdominal ventrite 5 coarsely punctate, except apical third subglabrous. Penis (Fig. 96b) with sides of body subparallel, weakly converging; apex rounded, sparsely setose; transfer apparatus small, compact, symmetrical; apodemes 2.0 × as long as body; ductus ejaculatorius without bulbus. Intraspecific variation. Length 2.27–2.36 mm. Female unknown.

Figure 95. — *Trigonopterus* *tujuh* Riedel, sp. n., holotype; a Habitus b Penis.

Figure 96. — *Trigonopterus* *ujungkulonensis* Riedel, sp. n., holotype; a Habitus b Penis.

Material examined.

Holotype (MZB): ARC2231 (EMBL # LM655687), East Nusa Tenggara Prov., Flores Isl., Labuhan Bajo, Roe, sample 3, S08°36.259', E120°01.539', 975 m, 13-III-2011 (MZB). Paratypes (MZB, SMNK): 4 exx, ARC2590 (EMBL # LM655899), same data as holotype.

Distribution.

East Nusa Tenggara Prov., Flores (Labuhan Bajo). Elevation: 975 m.

Etymology.

This epithet is based on the Indonesian word for “seven” and is treated as a noun in apposition.

Notes.

Trigonopterus tujuh Riedel, sp. n. was coded as “Trigonopterus sp. 325” by Tänzler et al. (2014).

96. Trigonopterus ujungkulonensis

Riedel sp. n.

http://zoobank.org/3C7F602D-DEEA-4CC6-AA5B-B854D33BD761

Diagnostic description.

Figure 99. — *Trigonopterus* *wallacei* Riedel, sp. n., holotype; a Habitus b Penis.

Material examined.

Holotype (SMNK): ARC2525 (EMBL # LM655862), MALAYSIA, Sarawak, Kuching, Mt. Santubong, sample 3, N01°43.884', E110°19.727', 496 m, 06-XII-2011. Paratypes (ARC, SMNK): MALAYSIA, Sarawak, Kuching: 2 exx, ARC2526 (EMBL # LM655863), same data as holotype; 1 ex, ARC2529 (EMBL # LM655866), Mt. Santubong, sample 4, N01°43.884', E110°19.715', 487 m, 06-XII-2011; 2 exx, sample 5, N01°43.843', E110°19.672', 478 m, 06-XII-2011; 2 exx, sample 6, N01°43.830', E110°19.601', 375 m, 06-XII-2011; 1 ex, Mt. Santubong, 700 m, 25-III-1990; 2 exx, Bako N.P., 27-III-1990.

Distribution.

Sarawak (Santubong). Elevation: 487–700 m.

Etymology.

This species is named in honour of Alfred Russel Wallace (1823–1913) for laying the foundation of research on the Natural History of the Indo-Australian Archipelago. He spent some time at this species´ type locality where he prepared the draft of his “Sarawak Law”.

Notes.

Trigonopterus wallacei Riedel, sp. n. was coded as “Trigonopterus sp. 362” by Tänzler et al. (2014).

Key to species from Java and Sumatra

1	Metafemur subapically simple, without stridulatory patch. Length 1.90–2.65 mm. (subgenus Mimidotasia)	2
1'	Metafemur subapically with stridulatory patch. Length 2.35–3.80 mm	9
2 (1)	Elytra (Figs 18a, 51a) elongate, 1.36–1.46 × longer than wide	3
2'	Elytra (e.g. Figs 24a, 29a, 38a) compact, 1.15–1.35 × longer than wide	4
3 (2)	Sutural interval of elytra swollen subapically, projecting in dorsal aspect. Penis (Fig. 18b), with long asymmetrical apical extension slightly upcurved and weakly directed to the left; E-Java	Trigonopterus costipennis Riedel, sp. n.
3'	Elytral apex with sutural interval costate but not distinctly projecting in dorsal aspect. Penis (Fig. 51b), with long asymmetrical apical extension markedly directed to the left; E-Sumatra	Trigonopterus misellus Riedel, sp. n.
4 (2')	Elytral striae with rows of elongate-claviform, rather slender scales	5
4'	Elytral striae with rows of claviform, wider scales	6
5 (4)	Elytral intervals costate-tuberculate. Penis (Fig. 24b) with medium-sized asymmetrical apical extension to the left	Trigonopterus diengensis Riedel, sp. n.
5'	Elytral intervals costate. Penis (Fig. 29b) with long asymmetrical apical extension directed to the right	Trigonopterus echinatus Riedel, sp. n.
6 (4')	Elytral intervals with relatively low costae. Penis (Figs 38b, 78b) with apical extension short, subangulate	7
6'	Elytral intervals with costae more distinct. Penis (Figs 34b, 73b) with apical extension acute, directed to the left	8
7 (6)	Color largely ferruginous. Penis (Fig. 38b). W-Java	Trigonopterus honjensi s Riedel, sp. n.
7'	Color largely black. Penis (Fig. 78b). E-Sumatra	Trigonopterus seriatus Riedel, sp. n.
8 (6')	Penis with endophallic structures as in Fig. 73b. E-Java	Trigonopterus satu Riedel, sp. n.
8'	Penis with endophallic structures as in Fig. 34b. W-Java	Trigonopterus foveatus Riedel, sp. n.
9 (1')	Elytra with striae deeply impressed; intervals costate to carinate. Femora each with small tooth. Penis (Fig. 64b) subapically forming blade-like processes. E-Java	Trigonopterus relictus Riedel, sp. n.
9'	If elytral intervals costate, at least profemur edentate. Penis different, without pair of subapical blade-like processes	10
10	Anteroventral ridge of femora forming blunt tooth, especially in meso- and metafemur	11
10'	All femora without tooth, or if species from Sumatra, meso- and metafemur with small denticle	16
11 (10)	Sutural interval of elytra at apex markedly swollen, forming pair of distinct apical protrusions. Species from W-Java	12
11'	Sutural interval of elytra at apex simple or weakly swollen. Species from E-Java	14
12 (11)	Sutural interval of elytra at apex forming median, apically bifid protrusion. Penis (Fig. 98b) with transfer apparatus minute	Trigonopterus vulcanicus Riedel, sp. n.
12'	Sutural interval of elytra at apex markedly swollen, forming pair of rounded apical protrusions. Penis (Figs 15b, 25b) basally with distinct transfer apparatus	13
13 (12')	Transfer apparatus of penis (Fig. 25b) compact, apically blunt	Trigonopterus dimorphus Riedel, sp. n.
13'	Transfer apparatus of penis (Fig. 15b) apically with spine	Trigonopterus binodulus Riedel, sp. n.
14 (11')	Disk of pronotum (Fig. 48a) densely, uniformly punctate. Protibia subbasally with distinct dorsoposterior tooth. Apex of penis (Fig. 48b) bidentate, with median notch	Trigonopterus merubetirensis Riedel, sp. n.
14'	Punctation of pronotum more coarsely, forming more or less distinct pattern (Figs 8a, 39a). Protibia subbasally with simple angulation, at most with minute denticle. Apex of penis (Figs 8b, 39b) simple or with small median extension	15
15 (14')	Disk of pronotum (Fig. 39a) with punctures partly elongate and / or arranged in confluent rows forming rhombus-like pattern. Transfer apparatus of penis in profile ventrally with subangular extension	Trigonopterus ijensis Riedel, sp. n.
15'	Disk of pronotum (Fig. 8a) with punctures less confluent; rhombus-like pattern less distinct. Transfer apparatus of penis in profile with ventral contour evenly convex	Trigonopterus argopurensis Riedel, sp. n.
16 (10')	Elytral intervals each with secondary rows of punctures resembling striae; at middle with transverse subglabrous band	Trigonopterus alaspurwensis Riedel, sp. n.
16'	Elytra with sculpture different; if sculpture coarse, never with transverse subglabrous band at middle	17
17 (16')	Elytra with striae indistinct, marked by fine lines and rows of small punctures; intervals flat, more or less coriaceous	18
17'	Elytra with striae deeply impressed and / or intervals elevated	20
18 (17)	Tarsomere 3 of protarsus 2.8–3.0 × as wide as tarsomere 1, markedly larger than of mesotarsus, (Fig. 45a)	Trigonopterus latipes Riedel, sp. n.
18'	Tarsomere 3 of protarsus 2.4–2.7 × as wide as tarsomere 1, at most slightly larger than of mesotarsus (Figs 1a, 9a)	19
19 (18')	Body (Fig. 9a) slender; elytra 1.25–1.34 × longer than wide; sculpture more distinct. Transfer apparatus of penis (Fig. 9b) flagelliform, 4.0 × longer than body; apodemes ca. 3 × as long as body	Trigonopterus arjunensis Riedel, sp. n.
19'	Body (Fig. 1a) across humeri wider; elytra 1.11–1.23 × longer than wide; sculpture rather subglabrous. Transfer apparatus of penis (Fig. 1b) thick flagelliform, 2.0 × longer than body; apodemes ca. 2 × as long as body	Trigonopterus acuminatus Riedel, sp. n.
20 (17')	Pronotum laterally without distinct preapical constriction. Usually larger species, length of elytra+pronotum 2.05–3.44 mm	21
20'	Pronotum laterally with distinct preapical constriction. Usually smaller species, length of elytra+pronotum 1.76–3.22 mm	26
21 (20)	Pronotum in apical half slightly rounded apicad. Species from East Sumatra	22
21'	Pronotum anterolaterally subangularly projecting. Species from Java	23
22 (21)	Elytra 2.62–2.65 × as long as pronotum. Penis (Fig. 83b) with more distinct lateral constriction. Mt. Rajabasa	Trigonopterus singularis Riedel, sp. n.
22'	Elytra 2.50–2.56 × as long as pronotum. Sides of penis (Fig. 7b) gently diverging towards apex. Pedada Bay and Bukit Barisan N.P	Trigonopterus amphoralis Marshall
23 (21')	Elytral intervals carinate; striae with rows of long erect piliform scales	24
23'	Elytral intervals costate; striae with rows of shorter subrecumbent scales	25
24 (23)	Rostrum apically without distinct median denticle. Apex of penis (Fig. 81a) subangulate, without median extension	Trigonopterus silvestris Riedel, sp. n.
24'	Rostrum apically with distinct median denticle. Apex of penis (Fig. 17b) medially with small rounded extension	Trigonopterus cahyoi Riedel, sp. n.
25 (23')	Elytral apex (Fig. 53a) wider and with sutural interval only weakly swollen. Elytral intervals almost flat, without punctures	Trigonopterus pangandaranensis Riedel, sp. n.
25'	Elytral apex (Fig. 96a) narrow and with swollen sutural interval. Irregular elytral intervals sparsely punctate	Trigonopterus ujungkulonensis Riedel, sp. n.
26 (20')	Sides of pronotum behind preapical constriction rounded, without distinct angular projections	27
26'	Sides of pronotum behind preapical constriction angularly projecting	31
27 (26)	Sutural intervals of elytra extended into median apical tooth. Mt. Gede	Trigonopterus gedensis Riedel, sp. n.
27'	Sutural intervals of elytra at most swollen, but without distinct tooth	28
28 (27')	Elytral intervals costate-tuberculate; striae each with sparse row of suberect bristles	29
28'	Elytral intervals carinate; striae each with row of suberect piliform scales or without conspicuous pubescence	30
29 (28)	Penis (Fig. 10b) with flagellum 1.5 × longer than body of penis	Trigonopterus asper Riedel, sp. n.
29'	Penis (Fig. 97b) with flagellum 1.1 × longer than body of penis	Trigonopterus variolosus Riedel, sp. n.
30 (28')	Elytral striae each with row of minute recumbent scales. E-Sumatra	Trigonopterus lampungensis Riedel, sp. n.
30'	Elytral striae with rows of suberect piliform scales. W-Java	Trigonopterus setifer Riedel, sp. n.
31 (26')	Elytral intervals irregularly costate-tuberculate; especially towards sides costae laterally corrugate	32
31'	Elytral intervals costate-carinate	36
32 (31)	Rostrum apically with denticle	33
32'	Rostrum apically without denticle	34
33 (32)	Metatibia ventrally with sparse row of long erect setae. Penis (Fig. 37b) with flagellum ca. 2.4 × longer than body of penis	Trigonopterus halimunensis Riedel, sp. n.
33'	Metatibia ventrally simple, without row of long setae. Penis (Fig. 28b) with flagellum ca. 1.9 × longer than body of penis	Trigonopterus duabelas Riedel, sp. n.
34 (32')	Elytral intervals partly flat, partly costate-tuberculate; interval 5 subbasally swollen. E-Sumatra	Trigonopterus squalidus Riedel, sp. n.
34'	Elytral intervals irregularly costate-tuberculate; interval 5 subbasally simple. W-Java	35
35 (34')	Penis (Fig. 4b) in dorsal aspect with body slender, sides subparallel	Trigonopterus allopatricus Riedel, sp. n.
35'	Penis (Fig. 97b) with body flattened, in dorsal aspect wider, lateral contours weakly diverging apicad	Trigonopterus variolosus Riedel, sp. n.
36 (32')	Elytra subnude, with sparse small recumbent setae	37
36'	Elytral striae each with row of suberect piliform scales	38
37 (36)	Anterolateral angular projections of pronotum distinct. Elytra setae recumbent but distinct. Java	Trigonopterus javensis Riedel, sp. n.
37'	Anterolateral angular projections of pronotum indistinct. Elytral setae minute, easily overlooked. E-Sumatra	Trigonopterus lampungensis Riedel, sp. n.
38 (36')	Profemur edentate, meso- and metafemur ventrally with small denticle. E-Sumatra	Trigonopterus sumatrensis Riedel, sp. n.
38'	All femora edentate. Java	39
39 (38')	Sutural interval of elytra basally swollen, widened, raised above level of other intervals	Trigonopterus suturalis Riedel, sp. n.
39'	Sutural interval of elytra surmounted by other intervals	40
40 (39')	Rostrum subapically with median denticle. Abdominal ventrite 5 swollen, at middle with shallow impression	41
40'	Rostrum subapically simple. Abdominal ventrite 5 flat or slightly swollen in apical half	42
41 (40)	Profile of abdominal ventrite 2 projecting angularly. Penis (Fig. 88b) apically with rounded extension; body with ca. 4 symmetrical pairs of elongate endophallic sclerites	Trigonopterus sundaicus Riedel, sp. n.
41'	Profile of abdominal ventrite 2 simple. Penis (Fig. 6b) apically with pointed extension; endophallic sclerites asymmetrical	Trigonopterus angulicollis Riedel, sp. n.
42 (40')	Body (Fig. 59a) slightly more elongate; elytra 1.31–1.52 × longer than wide. Penis (Fig. 59b) asymmetrical, appearing twisted	Trigonopterus porcatus Riedel, sp. n.
42'	Body (Fig. 58a, 67a) slightly more compact; elytra 1.18–1.29 × longer than wide. Penis (Figs 58, 67b) symmetrical	43
43 (42')	Penis (Fig. 58b) apically with short, rounded extension; flagellum 2 × longer than body of penis	Trigonopterus payungensis Riedel, sp. n.
43'	Penis (Fig. 67b) apically with acute triangular extension; flagellum slightly shorter than body of penis	Trigonopterus rugosostriatus Riedel, sp. n.

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Key to species from Bali

1	Body in dorsal aspect with marked constriction between pronotum and elytron. Pronotum in basal half with pair of sublateral, kidney-shaped impressions (Fig. 12a)	Trigonopterus baliensis Riedel, sp. n.
1'	Body subovate, in dorsal aspect with weak constriction between pronotum and elytron. Pronotum without pair of distinct sublateral impressions	2
2	Profile with distinct constriction between pronotum and elytron. Pronotum (Figs 55a, 68a) coarsely sculptured with irregular ridges and tubercles	Trigonopterus rugosus Riedel, sp. n. and Trigonopterus pararugosus Riedel, sp. n.
2'	Profile dorsally convex. Pronotum coarsely punctate, reticulate	3
3 (2')	Elytral intervals (Figs 43a, 49a) weakly costate or flat	4
3'	Elytral intervals (Figs 13a, 42a, 91a) dorsally markedly costate	5
4 (3)	Median rod of transfer apparatus thick, straight, 2.0 × as long as body of penis (Fig. 43b)	Trigonopterus klatakanensis Riedel, sp. n.
4'	Median rod of transfer apparatus flagelliform, forming almost full coil, ca 2.7 × as long as body of penis (Fig. 49b)	Trigonopterus mesehensis Riedel, sp. n.
5 (3')	Median rod of male transfer apparatus thin, shorter (1.4 × as long as body of penis), projecting basad (Fig. 13b)	Trigonopterus batukarensis Riedel, sp. n.
5'	Median rod of male transfer apparatus thick, longer (1.6–1.7 × as long as body of penis), projecting basad further than apicad (Figs 42b, 91b)	6
6 (5')	Body of penis (Fig. 42b) with sides subparallel, before apex weakly converging in straight line	Trigonopterus kintamanensis Riedel, sp. n.
6'	Body of penis (Fig. 91b) with shallow constriction at middle, before apex sides slightly swollen	Trigonopterus telagensis Riedel, sp. n.

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Key to species from Lombok, Sumbawa and Flores

1	Body in dorsal aspect with marked constriction between pronotum and elytron (e.g. Figs 20a, 87a)	2
1'	Body subovate, in dorsal aspect with weak constriction between pronotum and elytron (e.g. Figs 2a, 27a)	20
2 (1)	Length 1.44–2.20 mm. Pronotum in basal half with pair of sublateral impressions. Coloration of body dark ferruginous to black	3
2'	Length 2.20–4.04 mm. Pronotum without pair of distinct impressions. Coloration of body partly ferruginous, coppery, bronze, or black	11
3 (2)	Flagelliform transfer apparatus (Fig. 47b) subequal to body of penis. Lombok	Trigonopterus lombokensis Riedel, sp. n.
3'	Flagelliform transfer apparatus (Figs 33b, 54b, 56b, 57b, 60b, 61b, 66b, 87b) longer than body of penis	4
4 (3')	Penis (Figs 61b, 87b) subapically with sides angularly projecting, abruptly converging to slightly extended apex. Sumbawa	5
4'	Penis with weakly subangulate or rounded apex never extended medially	6
5 (4)	Flagelliform transfer apparatus (Fig. 87b) 2.9 × longer than body of penis	Trigonopterus sumbawensis Riedel, sp. n.
5'	Flagelliform transfer apparatus (Fig. 61b) 2.3–2.4 × longer than body of penis	Trigonopterus pseudosumbawensis Riedel, sp. n.
6 (4')	Flagelliform transfer apparatus (Fig. 66b) relatively thick, 2.6 × as long as body of penis; endophallus with sclerite. Flores	Trigonopterus roensis Riedel, sp. n.
6'	Flagelliform transfer apparatus (Figs 33b, 54b, 56b, 57b, 60b) thin, 3–6 × as long as body of penis; endophallus without sclerite	7
7 (6')	Flagelliform transfer apparatus ca. 3.0–3.3 × longer than body of penis; apodemes 2.4–2.5 × as long as body of penis. Sumbawa	8
7'	Flagelliform transfer apparatus ca. 4.0–7.0 × longer than body of penis; apodemes 3.2–4.1 × as long as body of penis. Flores	9
8 (7)	Penis (Fig. 56b) in apical third distinctly widening and rounded to weakly subangulate apex	Trigonopterus parasumbawensis Riedel, sp. n.
8'	Penis (Fig. 57b) with sides parallel before rounded apex	Trigonopterus pauxillus Riedel, sp. n.
9 (7')	Penis (Fig. 54b); flagelliform transfer apparatus 6.0–6.9 × as long as body	Trigonopterus paraflorensis Riedel, sp. n.
9'	Penis (Figs 33b, 60b); flagelliform transfer apparatus 3.9–4.6 × as long as body	10
10 (9')	Elytra more compact (Fig. 33a). Penis (Fig. 33b); flagelliform transfer apparatus 3.9–4.0 × as long as body. Medium elevations of the Mt. Ranaka / Lake Ranamese area	Trigonopterus florensis Riedel, sp. n.
10'	Elytra more slender (Fig. 60a). Penis (Fig. 60b); flagelliform transfer apparatus 4.1–4.6 × as long as body. Summit area of Mt. Ranaka	Trigonopterus pseudoflorensis Riedel, sp. n.
11 (2')	Profemur with large subacute tooth. Color of elytra black, usually basal third ferruginous	Trigonopterus dentipes Riedel, sp. n.
11'	Profemur with blunt tooth or simple	12
12 (11')	Color of elytra with bright metallic lustre, coppery	13
12'	Color of elytra black, sometimes basally ferruginous; at most with bronze lustre, never bright metallic	17
13 (12)	Onychium minute, drop-shaped, with median spine instead of pair of claws; weakly projecting beyond margin of tarsomere 3	Trigonopterus syarbis Riedel, sp. n.
13'	Onychium distinctly projecting beyond margin of tarsomere 3, apically with pair of claws	14
14 (13')	Elytra with humeri markedly swollen, laterally markedly projecting (Figs 35a, 50a)	15
14'	Elytra with humeri simple (Fig. 19a) or weakly swollen (Fig. 20a), not markedly projecting	16
15 (14)	Elytra (Fig. 35a) with interval 4 subbasally swollen, forming dorsal protrusion; sutural interval simple	Trigonopterus fulgidus Riedel, sp. n.
15'	Elytra (Fig. 50a) with sutural interval in basal half distinctly swollen; interval 4 without dorsal protrusion	Trigonopterus micans Riedel, sp. n.
16 (14')	Elytra (Fig. 20a) with interval 5 and 6 behind middle swollen, gently projecting from lateral outline. Penis (Fig. 20b) apically subtruncate, weakly rounded, subglabrous	Trigonopterus cupreus Riedel, sp. n.
16'	Elytra (Fig. 19a) with interval 5 and 6 simple. Penis (Fig. 19b) apically rounded, with fringe of conglutinate flattened setae, interrupted by glabrous median notch	Trigonopterus cuprescens Riedel, sp. n.
17 (12')	Body (Fig. 79a) subovate, slender. Meso- and metafemur anteroventrally with row of large sharp teeth	Trigonopterus serratifemur Riedel, sp. n.
17'	Body (Figs 21a, 72a, 84a) wider. Meso- and metafemur anteroventrally with single tooth	18
18 (17')	Elytra (Fig. 84a) completely black with slight bronze lustre; interval 5 forming dorsal tubercle; interval 6 behind middle swollen, gently projecting from lateral outline	Trigonopterus sinuatus Riedel, sp. n.
18'	Elytra basally ferruginous; intervals 5 and 6 simple	19
19 (18')	Elytral interval 5 in front of middle swollen, dorsally projecting. Penis (Fig. 21b)	Trigonopterus dacrycarpi Riedel, sp. n.
19'	Elytral interval 5 simple. Penis (Fig. 72b)	Trigonopterus sasak Riedel, sp. n.
20 (1')	species from Lombok	21
20'	species from Sumbawa	23
20'	species from Flores	27
21 (20)	Elytral intervals flat, subglabrous. Pronotum longitudinally rugose	Trigonopterus disruptus Riedel, sp. n.
21'	Elytral intervals costate. Pronotum coarsely punctate	22
22 (21')	Elytra black except sutural interval ferruginous; intervals 3 and 5 more distinctly swollen, with row of punctures	Trigonopterus rinjaniensis Riedel, sp. n.
22'	Elytra with bronze lustre; intervals 3 and 5 as other intervals, subglabrous	Trigonopterus aeneomicans Riedel, sp. n.
23 (20')	Elytra with intervals flat, striae marked by regular rows of punctures	Trigonopterus punctatoseriatus Riedel, sp. n.
23'	Elytra with striae deeply impressed, intervals costate	24
24 (23')	Elytra with bronze lustre	Trigonopterus aeneomicans Riedel, sp. n.
24'	Elytra largely black, without distinct metallic lustre	25
25 (24')	Sutural intervals of elytra costate, with row of punctures facing laterad. Abdominal ventrite 5 medially with shallow impression	Trigonopterus sembilan Riedel, sp. n.
25'	Sutural intervals of elytra forming pair of ridges, inner face with dense row of punctures. Abdominal ventrite 5 with median carina	26
26 (25')	Elytral interval 3 with dense row of punctures. Penis as in Fig. 70b, transfer apparatus with long median extension	Trigonopterus saltator Riedel, sp. n.
26'	Elytral interval 3 subglabrous, with few punctures. Penis as in Fig. 92b, transfer apparatus without median extension	Trigonopterus tepalensis Riedel, sp. n.
27 (20')	Tarsomere 3 of protarsus enlarged, medially deeply incised. Penis (Fig. 32b) in basal half with marked constriction; apex medially with distinct spine, with anterolateral subangular flanges	Trigonopterus fissitarsis Riedel, sp. n.
27'	Tarsomere 3 of protarsus of similar shape as tarsomere 3 of mesotarsus. Penis without marked constriction	28
28 (27')	Elytral intervals weakly costate, almost flat; striae consisting of rows of deeply impressed punctures	Trigonopterus tujuh Riedel, sp. n.
28'	Elytral intervals distinctly costate; striae incised	29
29 (28')	Abdominal ventrite 5 with median carina, at least in basal third	30
29'	Abdominal ventrite 5 without median carina	32
30 (29)	Elytral interval black; suture broadly sunk-in. Abdominal ventrite 5 with median carina in basal third	Trigonopterus delapan Riedel, sp. n.
30'	Elytral interval ferruginous. Abdominal ventrite 5 with complete median carina	31
31 (30')	Elytral intervals subglabrous, forming regular costae. Penis (Fig. 93b) with large anchor-shaped endophallic sclerite	Trigonopterus tiga Riedel, sp. n.
31'	Elytral intervals on dorsum punctate, along basal margin corrugate. Penis (Fig. 63b) with sides of body in basal half converging	Trigonopterus ranakensis Riedel, sp. n.
32 (29')	Metatibia apically with uncus and small premucro	33
32'	Metatibia apically with uncus, without premucro	34
33 (32)	Body (Fig. 27a) more slender. Suberect slender scales of elytral striae shorter	Trigonopterus dua Riedel, sp. n.
33'	Body (Fig. 69a) wider. Suberect slender scales of elytral striae longer	Trigonopterus rutengensis Riedel, sp. n.
34 (32')	Body (Fig. 31a) more slender. Suberect slender scales of elytral striae shorter	Trigonopterus enam Riedel, sp. n.
34'	Body wider (Figs 30a, 46a). Suberect slender scales of elytral striae longer	35
35 (34')	Penis (Fig. 46b) with transfer apparatus small, with median spine projecting to apex of endophallus	Trigonopterus lima Riedel, sp. n.
35'	Penis (Fig. 30b) with transfer apparatus compact, without median spine	Trigonopterus empat Riedel, sp. n.

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Key to species from Borneo and Palawan

1	Body subovate. Elytra with striae each bearing distinct row of suberect scales, intervals costate	2
1'	Body subovate or subrhomboid. Elytra with striae simple, punctures containing small scale or seta; intervals flat. IF elytra with sparse rows of suberect scales and costate intervals, then body subrhomboid	4
2	Epistome with dorsoposteriad directed horn and rostrum at middle with dorsal protrusion	3
2'	Epistome with denticle; rostrum with median and pair of submedian ridges, without dorsal protrusion between antennal insertions	Trigonopterus palawanensis Riedel, sp. n.
3 (2)	Length 1.52–1.79 mm. Elytral intervals narrow, weakly costate	Trigonopterus wallacei Riedel, sp. n.
3'	Length 2.32–2.45 mm. Elytral intervals subcarinate	Trigonopterus schulzi Riedel, sp. n.
4 (1')	Elytra with humeri swollen, laterally markedly projecting	5
4'	Elytra with humeri simple, not markedly projecting	6
5 (4)	Pronotum anteriorly simple. Elytral striae deeply impressed; intervals costate. Anteroventral ridge of meso- and metafemur simple	Trigonopterus trigonopterus Riedel, sp. n.
5'	Pronotum anteriorly with flanges projecting laterally. Elytral striae marked by small punctures; intervals flat. Anteroventral ridge of meso- and metafemur denticulate	Trigonopterus sebelas Riedel, sp. n.
6 (4')	Profemur in basal half posteroventrally with slightly bifid tooth. Male metatibia ventrally with dense fringe of long, stiff setae	Trigonopterus santubongensis Riedel, sp. n.
6'	Profemur in basal half posteroventrally without tooth. Male metatibia with sparse rows of setae	7
7 (6')	Elytral apex with suture distinctly incised. Abdominal ventrite 2 in profile simple.
7'	Elytral apex with suture simple, not incised. Abdominal ventrite 2 in profile projecting dentiform	9
8 (7)	Penis as in Fig. 16b; endophallus containing numerous coarse denticles, apically with pair of relatively small sclerites	Trigonopterus bornensis Riedel, sp. n.
8'	Penis as in Fig. 41b; endophallus with complex structures, several sclerites, containing few denticles	Trigonopterus kalimantanensis Riedel, sp. n.
9 (7')	In front of forehead median ridge of rostrum swollen, projecting subangularly from profile	10
9'	Base of rostrum in profile simple, evenly confluent with forehead
10 (9)	Penis as in Fig. 14b, near middle with shallow constriction; transfer apparatus complex, wider than long	Trigonopterus bawangensis Riedel, sp. n.
10'	Penis as in Fig. 82b, near middle widened; transfer apparatus complex, small	Trigonopterus singkawangensis Riedel, sp. n.
11 (9')	Color largely black. Elytral intervals flat. Penis as in Fig. 77b, apex rounded; body without distinct sclerites; transfer apparatus small, complex, wider than long	Trigonopterus sepuluh Riedel, sp. n.
11	Color largely ferruginous. Elytral intervals weakly costate. Penis as in Fig. 11b, with pair of large orificial sclerites, slightly more basad with pair of smaller, darker sclerites; apex with distinct median incision	Trigonopterus attenboroughi Riedel, sp. n.

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Catalogue of species groups of Trigonopterus Fauvel in Sundaland and the Lesser Sunda Islands

subgenus Mimidotasia Voss:Trigonopterus costipennis Riedel, sp. n., Trigonopterus diengensis Riedel, sp. n., Trigonopterus echinatus Riedel, sp. n., Trigonopterus foveatus Riedel, sp. n., Trigonopterus honjensis Riedel, sp. n., Trigonopterus misellus Riedel, sp. n., Trigonopterus satu Riedel, sp. n., Trigonopterus seriatus Riedel, sp. n.

Trigonopterus attenboroughi-group:Trigonopterus attenboroughi Riedel, sp. n., Trigonopterus bawangensis Riedel, sp. n., Trigonopterus santubongensis Riedel, sp. n., Trigonopterus sebelas Riedel, sp. n., Trigonopterus sepuluh Riedel, sp. n., Trigonopterus singkawangensis Riedel, sp. n.

Trigonopterus bornensis-group:Trigonopterus bornensis Riedel, sp. n., Trigonopterus kalimantanensis Riedel, sp. n.

Trigonopterus dimorphus-group:Trigonopterus acuminatus Riedel, sp. n., Trigonopterus alaspurwensis Riedel, sp. n., Trigonopterus allopatricus Riedel, sp. n., Trigonopterus amphoralis Marshall, Trigonopterus angulicollis Riedel, sp. n., Trigonopterus argopurensis Riedel, sp. n., Trigonopterus arjunensis Riedel, sp. n., Trigonopterus asper Riedel, sp. n., Trigonopterus binodulus Riedel, sp. n., Trigonopterus cahyoi Riedel, sp. n., Trigonopterus cuprescens Riedel, sp. n., Trigonopterus cupreus Riedel, sp. n., Trigonopterus dacrycarpi Riedel, sp. n., Trigonopterus dentipes Riedel, sp. n., Trigonopterus dimorphus Riedel, sp. n., Trigonopterus duabelas Riedel, sp. n., Trigonopterus fulgidus Riedel, sp. n., Trigonopterus gedensis Riedel, sp. n., Trigonopterus halimunensis Riedel, sp. n., Trigonopterus ijensis Riedel, sp. n., Trigonopterus javensis Riedel, sp. n., Trigonopterus lampungensis Riedel, sp. n., Trigonopterus latipes Riedel, sp. n., Trigonopterus merubetirensis Riedel, sp. n., Trigonopterus micans Riedel, sp. n., Trigonopterus pangandaranensis Riedel, sp. n., Trigonopterus payungensis Riedel, sp. n., Trigonopterus porcatus Riedel, sp. n., Trigonopterus rugosostriatus Riedel, sp. n., Trigonopterus sasak Riedel, sp. n., Trigonopterus serratifemur Riedel, sp. n., Trigonopterus setifer Riedel, sp. n., Trigonopterus silvestris Riedel, sp. n., Trigonopterus singularis Riedel, sp. n., Trigonopterus sinuatus Riedel, sp. n., Trigonopterus squalidus Riedel, sp. n., Trigonopterus sumatrensis Riedel, sp. n., Trigonopterus sundaicus Riedel, sp. n., Trigonopterus suturalis Riedel, sp. n., Trigonopterus syarbis Riedel, sp. n., Trigonopterus ujungkulonensis Riedel, sp. n., Trigonopterus variolosus Riedel, sp. n., Trigonopterus vulcanicus Riedel, sp. n.

Trigonopterus politus-group:Trigonopterus allotopus Riedel, sp. n.

Trigonopterus relictus-group:Trigonopterus baliensis Riedel, sp. n., Trigonopterus florensis Riedel, sp. n., Trigonopterus lombokensis Riedel, sp. n., Trigonopterus paraflorensis Riedel, sp. n., Trigonopterus parasumbawensis Riedel, sp. n., Trigonopterus pauxillus Riedel, sp. n., Trigonopterus pseudoflorensis Riedel, sp. n., Trigonopterus pseudosumbawensis Riedel, sp. n., Trigonopterus relictus Riedel, sp. n., Trigonopterus roensis Riedel, sp. n., Trigonopterus sumbawensis Riedel, sp. n.

Trigonopterus saltator-group:Trigonopterus aeneomicans Riedel, sp. n., Trigonopterus batukarensis Riedel, sp. n., Trigonopterus delapan Riedel, sp. n., Trigonopterus disruptus Riedel, sp. n., Trigonopterus dua Riedel, sp. n., Trigonopterus empat Riedel, sp. n., Trigonopterus enam Riedel, sp. n., Trigonopterus fissitarsis Riedel, sp. n., Trigonopterus kintamanensis Riedel, sp. n., Trigonopterus klatakanensis Riedel, sp. n., Trigonopterus lima Riedel, sp. n., Trigonopterus mesehensis Riedel, sp. n., Trigonopterus pararugosus Riedel, sp. n., Trigonopterus punctatoseriatus Riedel, sp. n., Trigonopterus ranakensis Riedel, sp. n., Trigonopterus rinjaniensis Riedel, sp. n., Trigonopterus rugosus Riedel, sp. n., Trigonopterus rutengensis Riedel, sp. n., Trigonopterus saltator Riedel, sp. n., Trigonopterus sembilan Riedel, sp. n., Trigonopterus telagensis Riedel, sp. n., Trigonopterus tepalensis Riedel, sp. n., Trigonopterus tiga Riedel, sp. n., Trigonopterus tujuh Riedel, sp. n.

Trigonopterus trigonopterus-group:Trigonopterus trigonopterus Riedel, sp. n.

Trigonopterus wallacei-group:Trigonopterus palawanensis Riedel, sp. n., Trigonopterus schulzi Riedel, sp. n., Trigonopterus wallacei Riedel, sp. n.

Plates

Discussion

Herein, we apply our fast track taxonomy approach (Riedel et al. 2013a) to revise the entire Trigonopterus fauna of a defined region. The proportion of closely related species to be separated was higher than in our earlier work covering representatives of different species groups (Riedel et al. 2013b), but this did not cause any complications. Diagnosis of species using cox1 sequences was straightforward and the question where to draw the line between allopatric populations and putative species entities usually could be answered by the evaluation of morphological, especially genital characters. In a few cases, especially in the Trigonopterus relictus-group, sibling species could be revealed that occur syntopically. After recognizing a subdivision into well separated clades of relatively high divergence we reassessed genital characters. We found that the length of the flagellum was correlated with these clusters. It can only be speculated that their discovery would not have been possible with morphological taxonomy alone as the taxonomist in charge had initially relegated these differences to intraspecific variation by lumping a number of them.

At the start of this project and based on literature the high diversity of Trigonopterus in the Papuan region (Riedel et al. 2010; Tänzler et al. 2012) seemed to contrast with a relatively poor representation in the Philippines (three described species: Heller 1915; Hustache 1925) and Sulawesi (one described species: Riedel 2011), while its presence in Sundaland appeared questionable – the single species described by Marshall (1925) from East Sumatra looked out of place and more like the result of a mistake than representing a reliable record. Step by step, extensive field work revealed this picture to be a simple artifact. In fact, the islands of the Sunda Arc and Borneo each harbour a rich and so far overlooked fauna, at least in places with wet primary forest. However, unlike in New Guinea and Wallacea where many species can be collected from foliage, the fauna of the Sunda Islands is largely restricted to the litter layer more difficult to sample and thus largely neglected by collectors. Moreover, Trigonopterus weevils usually fall into the category of “small beetles” which have a higher chance to be undescribed than beetles of larger body size (Stork et al. 2008).

The field work for this study focused on eastern parts of Sumatra, Java and the Lesser Sunda Islands. Materials stored in museum collections neither contained Trigonopterus from the western parts of Sumatra, nor from the Malay Peninsula. A dedicated search in the field could possibly lead to interesting discoveries, but equally well the genus´ real area of distribution excludes this region or parts of it. More promising for future discoveries of species are Borneo and Palawan where only material from a few localities was available. Borneo obviously possesses a rich fauna of Trigonopterus and its species number will be magnitudes higher than the ten species recorded so far. Species discovery on Java (35 species recorded), Bali (8 species recorded), and Lombok (7 species recorded) may approach saturation, while additional species are likely to be found on Sumatra (7 species recorded), Sumbawa (12 species recorded), and Flores (18 species recorded).

Most localities sampled by us harbour at least one species of Trigonopterus; in some places many more were found, such as seven species each at Mount Sawal of West Java, and Mount Ranaka of Flores Island. Although these figures are dwarfed by comparable localities of New Guinea (Riedel et al. 2010), the cumulative number of highly endemic species found on the Sunda Islands is substantial. A few of the species found in the montane forests of West Java are of relatively wide distribution, (e.g. Trigonopterus allopatricus Riedel, sp. n., Trigonopterus javensis Riedel, sp. n., Trigonopterus vulcanicus Riedel, sp. n.), but none cover the entire island. Species with a narrower distribution range can be found on somewhat isolated mountain blocks with suitable vegetation, e.g. Trigonopterus dimorphus Riedel, sp. n. and Trigonopterus halimunensis Riedel, sp. n. on Mt. Halimun-Salak, Trigonopterus angulicollis Riedel, sp. n. and Trigonopterus gedensis Riedel, sp. n. on Mt. Gede. For lowland habitats the original degree of endemism is harder to judge since the areas of lowland primary forest remaining today are very restricted themselves. The two species recorded for Ujung Kulon National Park are found nowhere else but in this last major area of primary lowland forest of western Java. Similarly, the species found in Alas Purwo National Park and the two species found in Meru Betiri National Park are also restricted to their respective localities, although the two are separated by less than 60 km distance. Such a distance could easily straddle geological, respectively ecological boundaries effectively separating populations of flightless beetles. At the same time, it raises the question how many species once inhabiting the other areas of Java´s lowland forest have become extinct when the land was converted to agricultural use.

Wide areas of East Java and the Lesser Sunda Islands are markedly influenced by seasonal climate resulting in extensive monsoon forests, while wet evergreen forests persist on the slopes of higher mountains (Whitten et al. 1996). Trigonopterus species are restricted to the latter areas, and the monsoon forests are a habitat unsuitable to them, just like the vegetation altered by human influence. Even before the arrival of humans on the Sunda Islands the habitats of Trigonopterus were probably somewhat patchy for climatic reasons. With agriculture changing the land and human population increasing, areas of wet evergreen forests became increasingly fragmented and at times limited to a few square kilometers or less. Besides the usual risks that come with small areas of habitat for animals, additional factors pose serious hazards to the species inhabiting these forests: 1) Many of the mountains supporting these forests are active or dormant volcanoes. Eruptions may destroy major portions of the habitat. During our sampling campaign violent eruptions of Mount Merapi destroyed the last remaining area of primary forest on this mountain. 2) With increasing accessibility spots of natural beauty such as pockets of wet evergreen forests become more vulnerable to illegal logging and to disasters such as wildland fires. During times of drought a cigarette or a campfire may be enough to wipe out the remaining habitat. A wide and sealed road built for the purpose of recreation gave us easy access to a small fragment of wet montane forest on slope of Mount Wilis. It can only be hoped that the road´s impact will not be fatal to Trigonopterus acuminatus Riedel, sp. n., a species exclusive to this area. 3) Many of the remaining wet evergreen forests stand on fertile grounds highly suitable to cultivate vegetables. With some of the highest population densities worldwide, it is not surprising that in Java, Bali and Lombok gardens tend to expand on the mountain slopes and encroach even on protected forests. Many Trigonopterus species have constraints in terms of elevation: For example, Trigonopterus argopurensis Riedel, sp. n. was found to a maximum elevation of 1785 m. Although a larger forest area exists on the upper slopes of Mt. Argopuro, it seems to be unsuitable to this particular species which is cornered by coffee gardens in its lower range.

Looking at the distribution of Trigonopterus species in the Sunda Arc and the areas of remaining primary forests, it appears likely that some of these newly discovered species will become extinct in the next few decades. The blame will be largely on human activities which have reduced the once extensive areas of forest to small fragments. It is our responsibility to help preserving these remaining areas of habitat. We should intensify our efforts especially with those forests harbouring endemic species but nevertheless holding a relatively weak conservation status, as exemplified by the reserves of Mount Sawal and Mount Ranaka. In these cases, a limited investment in better protection could achieve a significant improvement for species conservation.

Herein, we provide faces to 98 hitherto unknown species by publishing their descriptions and images, in parallel also on the wiki-site Species-ID. The new names will be useful to deposit scientific data reliably and sustainably. Separate studies on the phylogeny and distribution patterns of Trigonopterus are in progress and will build on this taxonomic foundation. Most of all, it is hoped that knowledge of this previously unknown fauna will create an awareness for the conservation value of the remaining fragments of wet evergreen primary forest in the region. There may be a certain redundancy of conservation areas when we look at the distribution of many vertebrates – however, the larger portion of biodiversity as represented by arthropods exhibits a different picture. The message that a study of flightless weevils of the Sunda Arc conveys is clear: with almost every area of remaining primary forest being destroyed we will lose some more species – with or without names.

Supplementary Material

XML Treatment for Trigonopterus

zookeys.467.8206-treatment1.xml^{(2.8KB, xml)}

XML Treatment for Trigonopterus acuminatus

zookeys.467.8206-treatment2.xml^{(7.8KB, xml)}

XML Treatment for Trigonopterus aeneomicans

zookeys.467.8206-treatment3.xml^{(15.1KB, xml)}

XML Treatment for Trigonopterus alaspurwensis

zookeys.467.8206-treatment4.xml^{(8.4KB, xml)}

XML Treatment for Trigonopterus allopatricus

zookeys.467.8206-treatment5.xml^{(9KB, xml)}

XML Treatment for Trigonopterus allotopus

zookeys.467.8206-treatment6.xml^{(7.2KB, xml)}

XML Treatment for Trigonopterus angulicollis

zookeys.467.8206-treatment7.xml^{(7.1KB, xml)}

XML Treatment for Trigonopterus amphoralis

zookeys.467.8206-treatment8.xml^{(10.3KB, xml)}

XML Treatment for Trigonopterus argopurensis

zookeys.467.8206-treatment9.xml^{(7.7KB, xml)}

XML Treatment for Trigonopterus arjunensis

zookeys.467.8206-treatment10.xml^{(7.1KB, xml)}

XML Treatment for Trigonopterus asper

zookeys.467.8206-treatment11.xml^{(6.3KB, xml)}

XML Treatment for Trigonopterus attenboroughi

zookeys.467.8206-treatment12.xml^{(6.8KB, xml)}

XML Treatment for Trigonopterus baliensis

zookeys.467.8206-treatment13.xml^{(12.3KB, xml)}

XML Treatment for Trigonopterus batukarensis

zookeys.467.8206-treatment14.xml^{(7.4KB, xml)}

XML Treatment for Trigonopterus bawangensis

zookeys.467.8206-treatment15.xml^{(6.7KB, xml)}

XML Treatment for Trigonopterus binodulus

zookeys.467.8206-treatment16.xml^{(6.5KB, xml)}

XML Treatment for Trigonopterus bornensis

zookeys.467.8206-treatment17.xml^{(6.2KB, xml)}

XML Treatment for Trigonopterus cahyoi

zookeys.467.8206-treatment18.xml^{(8.6KB, xml)}

XML Treatment for Trigonopterus costipennis

zookeys.467.8206-treatment19.xml^{(7.5KB, xml)}

XML Treatment for Trigonopterus cuprescens

zookeys.467.8206-treatment20.xml^{(6.6KB, xml)}

XML Treatment for Trigonopterus cupreus

zookeys.467.8206-treatment21.xml^{(6.6KB, xml)}

XML Treatment for Trigonopterus dacrycarpi

zookeys.467.8206-treatment22.xml^{(7.6KB, xml)}

XML Treatment for Trigonopterus delapan

zookeys.467.8206-treatment23.xml^{(6.9KB, xml)}

XML Treatment for Trigonopterus dentipes

zookeys.467.8206-treatment24.xml^{(9KB, xml)}

XML Treatment for Trigonopterus diengensis

zookeys.467.8206-treatment25.xml^{(5.5KB, xml)}

XML Treatment for Trigonopterus dimorphus

zookeys.467.8206-treatment26.xml^{(6.9KB, xml)}

XML Treatment for Trigonopterus disruptus

zookeys.467.8206-treatment27.xml^{(12.6KB, xml)}

XML Treatment for Trigonopterus dua

zookeys.467.8206-treatment28.xml^{(6.4KB, xml)}

XML Treatment for Trigonopterus duabelas

zookeys.467.8206-treatment29.xml^{(6.8KB, xml)}

XML Treatment for Trigonopterus echinatus

zookeys.467.8206-treatment30.xml^{(6KB, xml)}

XML Treatment for Trigonopterus empat

zookeys.467.8206-treatment31.xml^{(6.8KB, xml)}

XML Treatment for Trigonopterus enam

zookeys.467.8206-treatment32.xml^{(7KB, xml)}

XML Treatment for Trigonopterus fissitarsis

zookeys.467.8206-treatment33.xml^{(6.8KB, xml)}

XML Treatment for Trigonopterus florensis

zookeys.467.8206-treatment34.xml^{(7.3KB, xml)}

XML Treatment for Trigonopterus foveatus

zookeys.467.8206-treatment35.xml^{(6.1KB, xml)}

XML Treatment for Trigonopterus fulgidus

zookeys.467.8206-treatment36.xml^{(10.1KB, xml)}

XML Treatment for Trigonopterus gedensis

zookeys.467.8206-treatment37.xml^{(7.3KB, xml)}

XML Treatment for Trigonopterus halimunensis

zookeys.467.8206-treatment38.xml^{(8.4KB, xml)}

XML Treatment for Trigonopterus honjensis

zookeys.467.8206-treatment39.xml^{(6KB, xml)}

XML Treatment for Trigonopterus ijensis

zookeys.467.8206-treatment40.xml^{(6.4KB, xml)}

XML Treatment for Trigonopterus javensis

zookeys.467.8206-treatment41.xml^{(8.6KB, xml)}

XML Treatment for Trigonopterus kalimantanensis

zookeys.467.8206-treatment42.xml^{(7KB, xml)}

XML Treatment for Trigonopterus kintamanensis

zookeys.467.8206-treatment43.xml^{(6.6KB, xml)}

XML Treatment for Trigonopterus klatakanensis

zookeys.467.8206-treatment44.xml^{(7.1KB, xml)}

XML Treatment for Trigonopterus lampungensis

zookeys.467.8206-treatment45.xml^{(7.8KB, xml)}

XML Treatment for Trigonopterus latipes

zookeys.467.8206-treatment46.xml^{(6.1KB, xml)}

XML Treatment for Trigonopterus lima

zookeys.467.8206-treatment47.xml^{(6.2KB, xml)}

XML Treatment for Trigonopterus lombokensis

zookeys.467.8206-treatment48.xml^{(11.3KB, xml)}

XML Treatment for Trigonopterus merubetirensis

zookeys.467.8206-treatment49.xml^{(6KB, xml)}

XML Treatment for Trigonopterus mesehensis

zookeys.467.8206-treatment50.xml^{(8.3KB, xml)}

XML Treatment for Trigonopterus micans

zookeys.467.8206-treatment51.xml^{(7.8KB, xml)}

XML Treatment for Trigonopterus misellus

zookeys.467.8206-treatment52.xml^{(5.7KB, xml)}

XML Treatment for Trigonopterus palawanensis

zookeys.467.8206-treatment53.xml^{(5.5KB, xml)}

XML Treatment for Trigonopterus pangandaranensis

zookeys.467.8206-treatment54.xml^{(6.1KB, xml)}

XML Treatment for Trigonopterus paraflorensis

zookeys.467.8206-treatment55.xml^{(6.4KB, xml)}

XML Treatment for Trigonopterus pararugosus

zookeys.467.8206-treatment56.xml^{(9.1KB, xml)}

XML Treatment for Trigonopterus parasumbawensis

zookeys.467.8206-treatment57.xml^{(6.3KB, xml)}

XML Treatment for Trigonopterus pauxillus

zookeys.467.8206-treatment58.xml^{(7.9KB, xml)}

XML Treatment for Trigonopterus payungensis

zookeys.467.8206-treatment59.xml^{(5.9KB, xml)}

XML Treatment for Trigonopterus porcatus

zookeys.467.8206-treatment60.xml^{(6.3KB, xml)}

XML Treatment for Trigonopterus pseudoflorensis

zookeys.467.8206-treatment61.xml^{(7.7KB, xml)}

XML Treatment for Trigonopterus pseudosumbawensis

zookeys.467.8206-treatment62.xml^{(6KB, xml)}

XML Treatment for Trigonopterus punctatoseriatus

zookeys.467.8206-treatment63.xml^{(7.7KB, xml)}

XML Treatment for Trigonopterus ranakensis

zookeys.467.8206-treatment64.xml^{(7.3KB, xml)}

XML Treatment for Trigonopterus relictus

zookeys.467.8206-treatment65.xml^{(9KB, xml)}

XML Treatment for Trigonopterus rinjaniensis

zookeys.467.8206-treatment66.xml^{(8.9KB, xml)}

XML Treatment for Trigonopterus roensis

zookeys.467.8206-treatment67.xml^{(7.1KB, xml)}

XML Treatment for Trigonopterus rugosostriatus

zookeys.467.8206-treatment68.xml^{(7.5KB, xml)}

XML Treatment for Trigonopterus rugosus

zookeys.467.8206-treatment69.xml^{(10.2KB, xml)}

XML Treatment for Trigonopterus rutengensis

zookeys.467.8206-treatment70.xml^{(6.8KB, xml)}

XML Treatment for Trigonopterus saltator

zookeys.467.8206-treatment71.xml^{(7KB, xml)}

XML Treatment for Trigonopterus santubongensis

zookeys.467.8206-treatment72.xml^{(7.4KB, xml)}

XML Treatment for Trigonopterus sasak

zookeys.467.8206-treatment73.xml^{(8.4KB, xml)}

XML Treatment for Trigonopterus satu

zookeys.467.8206-treatment74.xml^{(5.8KB, xml)}

XML Treatment for Trigonopterus schulzi

zookeys.467.8206-treatment75.xml^{(6.7KB, xml)}

XML Treatment for Trigonopterus sebelas

zookeys.467.8206-treatment76.xml^{(8.2KB, xml)}

XML Treatment for Trigonopterus sembilan

zookeys.467.8206-treatment77.xml^{(6.4KB, xml)}

XML Treatment for Trigonopterus sepuluh

zookeys.467.8206-treatment78.xml^{(7KB, xml)}

XML Treatment for Trigonopterus seriatus

zookeys.467.8206-treatment79.xml^{(8.4KB, xml)}

XML Treatment for Trigonopterus serratifemur

zookeys.467.8206-treatment80.xml^{(6.3KB, xml)}

XML Treatment for Trigonopterus setifer

zookeys.467.8206-treatment81.xml^{(6KB, xml)}

XML Treatment for Trigonopterus silvestris

zookeys.467.8206-treatment82.xml^{(6.5KB, xml)}

XML Treatment for Trigonopterus singkawangensis

zookeys.467.8206-treatment83.xml^{(5.7KB, xml)}

XML Treatment for Trigonopterus singularis

zookeys.467.8206-treatment84.xml^{(6.5KB, xml)}

XML Treatment for Trigonopterus sinuatus

zookeys.467.8206-treatment85.xml^{(9KB, xml)}

XML Treatment for Trigonopterus squalidus

zookeys.467.8206-treatment86.xml^{(6.4KB, xml)}

XML Treatment for Trigonopterus sumatrensis

zookeys.467.8206-treatment87.xml^{(6.8KB, xml)}

XML Treatment for Trigonopterus sumbawensis

zookeys.467.8206-treatment88.xml^{(8KB, xml)}

XML Treatment for Trigonopterus sundaicus

zookeys.467.8206-treatment89.xml^{(8.3KB, xml)}

XML Treatment for Trigonopterus suturalis

zookeys.467.8206-treatment90.xml^{(8.3KB, xml)}

XML Treatment for Trigonopterus syarbis

zookeys.467.8206-treatment91.xml^{(5.7KB, xml)}

XML Treatment for Trigonopterus telagensis

zookeys.467.8206-treatment92.xml^{(7.1KB, xml)}

XML Treatment for Trigonopterus tepalensis

zookeys.467.8206-treatment93.xml^{(5.3KB, xml)}

XML Treatment for Trigonopterus tiga

zookeys.467.8206-treatment94.xml^{(7KB, xml)}

XML Treatment for Trigonopterus trigonopterus

zookeys.467.8206-treatment95.xml^{(8KB, xml)}

XML Treatment for Trigonopterus tujuh

zookeys.467.8206-treatment96.xml^{(7.2KB, xml)}

XML Treatment for Trigonopterus ujungkulonensis

zookeys.467.8206-treatment97.xml^{(6.6KB, xml)}

XML Treatment for Trigonopterus variolosus

zookeys.467.8206-treatment98.xml^{(6.2KB, xml)}

XML Treatment for Trigonopterus vulcanicus

zookeys.467.8206-treatment99.xml^{(12KB, xml)}

XML Treatment for Trigonopterus wallacei

zookeys.467.8206-treatment100.xml^{(7.4KB, xml)}

Acknowledgements

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, and we thank all of them very warmly. Michi Schrödl (Munich) gave inspiration to use numbers as species epithets. We are grateful to Lyubomir Penev (Sofia) for his constant support during the publication of this study. We thank the German Research Foundation DFG (RI 1817/1-1, 3-1, 3-3) for financial support.

Citation

Riedel A, Tänzler R, Balke M, Rahmadi C, Suhardjono YR (2014) Ninety-eight new species of Trigonopterus weevils from Sundaland and the Lesser Sunda Islands. ZooKeys 467: 1–162. doi: 10.3897/zookeys.467.8206

References

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

XML Treatment for Trigonopterus

zookeys.467.8206-treatment1.xml^{(2.8KB, xml)}

XML Treatment for Trigonopterus acuminatus

zookeys.467.8206-treatment2.xml^{(7.8KB, xml)}

XML Treatment for Trigonopterus aeneomicans

zookeys.467.8206-treatment3.xml^{(15.1KB, xml)}

XML Treatment for Trigonopterus alaspurwensis

zookeys.467.8206-treatment4.xml^{(8.4KB, xml)}

XML Treatment for Trigonopterus allopatricus

zookeys.467.8206-treatment5.xml^{(9KB, xml)}

XML Treatment for Trigonopterus allotopus

zookeys.467.8206-treatment6.xml^{(7.2KB, xml)}

XML Treatment for Trigonopterus angulicollis

zookeys.467.8206-treatment7.xml^{(7.1KB, xml)}

XML Treatment for Trigonopterus amphoralis

zookeys.467.8206-treatment8.xml^{(10.3KB, xml)}

XML Treatment for Trigonopterus argopurensis

zookeys.467.8206-treatment9.xml^{(7.7KB, xml)}

XML Treatment for Trigonopterus arjunensis

zookeys.467.8206-treatment10.xml^{(7.1KB, xml)}

XML Treatment for Trigonopterus asper

zookeys.467.8206-treatment11.xml^{(6.3KB, xml)}

XML Treatment for Trigonopterus attenboroughi

zookeys.467.8206-treatment12.xml^{(6.8KB, xml)}

XML Treatment for Trigonopterus baliensis

zookeys.467.8206-treatment13.xml^{(12.3KB, xml)}

XML Treatment for Trigonopterus batukarensis

zookeys.467.8206-treatment14.xml^{(7.4KB, xml)}

XML Treatment for Trigonopterus bawangensis

zookeys.467.8206-treatment15.xml^{(6.7KB, xml)}

XML Treatment for Trigonopterus binodulus

zookeys.467.8206-treatment16.xml^{(6.5KB, xml)}

XML Treatment for Trigonopterus bornensis

zookeys.467.8206-treatment17.xml^{(6.2KB, xml)}

XML Treatment for Trigonopterus cahyoi

zookeys.467.8206-treatment18.xml^{(8.6KB, xml)}

XML Treatment for Trigonopterus costipennis

zookeys.467.8206-treatment19.xml^{(7.5KB, xml)}

XML Treatment for Trigonopterus cuprescens

zookeys.467.8206-treatment20.xml^{(6.6KB, xml)}

XML Treatment for Trigonopterus cupreus

zookeys.467.8206-treatment21.xml^{(6.6KB, xml)}

XML Treatment for Trigonopterus dacrycarpi

zookeys.467.8206-treatment22.xml^{(7.6KB, xml)}

XML Treatment for Trigonopterus delapan

zookeys.467.8206-treatment23.xml^{(6.9KB, xml)}

XML Treatment for Trigonopterus dentipes

zookeys.467.8206-treatment24.xml^{(9KB, xml)}

XML Treatment for Trigonopterus diengensis

zookeys.467.8206-treatment25.xml^{(5.5KB, xml)}

XML Treatment for Trigonopterus dimorphus

zookeys.467.8206-treatment26.xml^{(6.9KB, xml)}

XML Treatment for Trigonopterus disruptus

zookeys.467.8206-treatment27.xml^{(12.6KB, xml)}

XML Treatment for Trigonopterus dua

zookeys.467.8206-treatment28.xml^{(6.4KB, xml)}

XML Treatment for Trigonopterus duabelas

zookeys.467.8206-treatment29.xml^{(6.8KB, xml)}

XML Treatment for Trigonopterus echinatus

zookeys.467.8206-treatment30.xml^{(6KB, xml)}

XML Treatment for Trigonopterus empat

zookeys.467.8206-treatment31.xml^{(6.8KB, xml)}

XML Treatment for Trigonopterus enam

zookeys.467.8206-treatment32.xml^{(7KB, xml)}

XML Treatment for Trigonopterus fissitarsis

zookeys.467.8206-treatment33.xml^{(6.8KB, xml)}

XML Treatment for Trigonopterus florensis

zookeys.467.8206-treatment34.xml^{(7.3KB, xml)}

XML Treatment for Trigonopterus foveatus

zookeys.467.8206-treatment35.xml^{(6.1KB, xml)}

XML Treatment for Trigonopterus fulgidus

zookeys.467.8206-treatment36.xml^{(10.1KB, xml)}

XML Treatment for Trigonopterus gedensis

zookeys.467.8206-treatment37.xml^{(7.3KB, xml)}

XML Treatment for Trigonopterus halimunensis

zookeys.467.8206-treatment38.xml^{(8.4KB, xml)}

XML Treatment for Trigonopterus honjensis

zookeys.467.8206-treatment39.xml^{(6KB, xml)}

XML Treatment for Trigonopterus ijensis

zookeys.467.8206-treatment40.xml^{(6.4KB, xml)}

XML Treatment for Trigonopterus javensis

zookeys.467.8206-treatment41.xml^{(8.6KB, xml)}

XML Treatment for Trigonopterus kalimantanensis

zookeys.467.8206-treatment42.xml^{(7KB, xml)}

XML Treatment for Trigonopterus kintamanensis

zookeys.467.8206-treatment43.xml^{(6.6KB, xml)}

XML Treatment for Trigonopterus klatakanensis

zookeys.467.8206-treatment44.xml^{(7.1KB, xml)}

XML Treatment for Trigonopterus lampungensis

zookeys.467.8206-treatment45.xml^{(7.8KB, xml)}

XML Treatment for Trigonopterus latipes

zookeys.467.8206-treatment46.xml^{(6.1KB, xml)}

XML Treatment for Trigonopterus lima

zookeys.467.8206-treatment47.xml^{(6.2KB, xml)}

XML Treatment for Trigonopterus lombokensis

zookeys.467.8206-treatment48.xml^{(11.3KB, xml)}

XML Treatment for Trigonopterus merubetirensis

zookeys.467.8206-treatment49.xml^{(6KB, xml)}

XML Treatment for Trigonopterus mesehensis

zookeys.467.8206-treatment50.xml^{(8.3KB, xml)}

XML Treatment for Trigonopterus micans

zookeys.467.8206-treatment51.xml^{(7.8KB, xml)}

XML Treatment for Trigonopterus misellus

zookeys.467.8206-treatment52.xml^{(5.7KB, xml)}

XML Treatment for Trigonopterus palawanensis

zookeys.467.8206-treatment53.xml^{(5.5KB, xml)}

XML Treatment for Trigonopterus pangandaranensis

zookeys.467.8206-treatment54.xml^{(6.1KB, xml)}

XML Treatment for Trigonopterus paraflorensis

zookeys.467.8206-treatment55.xml^{(6.4KB, xml)}

XML Treatment for Trigonopterus pararugosus

zookeys.467.8206-treatment56.xml^{(9.1KB, xml)}

XML Treatment for Trigonopterus parasumbawensis

zookeys.467.8206-treatment57.xml^{(6.3KB, xml)}

XML Treatment for Trigonopterus pauxillus

zookeys.467.8206-treatment58.xml^{(7.9KB, xml)}

XML Treatment for Trigonopterus payungensis

zookeys.467.8206-treatment59.xml^{(5.9KB, xml)}

XML Treatment for Trigonopterus porcatus

zookeys.467.8206-treatment60.xml^{(6.3KB, xml)}

XML Treatment for Trigonopterus pseudoflorensis

zookeys.467.8206-treatment61.xml^{(7.7KB, xml)}

XML Treatment for Trigonopterus pseudosumbawensis

zookeys.467.8206-treatment62.xml^{(6KB, xml)}

XML Treatment for Trigonopterus punctatoseriatus

zookeys.467.8206-treatment63.xml^{(7.7KB, xml)}

XML Treatment for Trigonopterus ranakensis

zookeys.467.8206-treatment64.xml^{(7.3KB, xml)}

XML Treatment for Trigonopterus relictus

zookeys.467.8206-treatment65.xml^{(9KB, xml)}

XML Treatment for Trigonopterus rinjaniensis

zookeys.467.8206-treatment66.xml^{(8.9KB, xml)}

XML Treatment for Trigonopterus roensis

zookeys.467.8206-treatment67.xml^{(7.1KB, xml)}

XML Treatment for Trigonopterus rugosostriatus

zookeys.467.8206-treatment68.xml^{(7.5KB, xml)}

XML Treatment for Trigonopterus rugosus

zookeys.467.8206-treatment69.xml^{(10.2KB, xml)}

XML Treatment for Trigonopterus rutengensis

zookeys.467.8206-treatment70.xml^{(6.8KB, xml)}

XML Treatment for Trigonopterus saltator

zookeys.467.8206-treatment71.xml^{(7KB, xml)}

XML Treatment for Trigonopterus santubongensis

zookeys.467.8206-treatment72.xml^{(7.4KB, xml)}

XML Treatment for Trigonopterus sasak

zookeys.467.8206-treatment73.xml^{(8.4KB, xml)}

XML Treatment for Trigonopterus satu

zookeys.467.8206-treatment74.xml^{(5.8KB, xml)}

XML Treatment for Trigonopterus schulzi

zookeys.467.8206-treatment75.xml^{(6.7KB, xml)}

XML Treatment for Trigonopterus sebelas

zookeys.467.8206-treatment76.xml^{(8.2KB, xml)}

XML Treatment for Trigonopterus sembilan

zookeys.467.8206-treatment77.xml^{(6.4KB, xml)}

XML Treatment for Trigonopterus sepuluh

zookeys.467.8206-treatment78.xml^{(7KB, xml)}

XML Treatment for Trigonopterus seriatus

zookeys.467.8206-treatment79.xml^{(8.4KB, xml)}

XML Treatment for Trigonopterus serratifemur

zookeys.467.8206-treatment80.xml^{(6.3KB, xml)}

XML Treatment for Trigonopterus setifer

zookeys.467.8206-treatment81.xml^{(6KB, xml)}

XML Treatment for Trigonopterus silvestris

zookeys.467.8206-treatment82.xml^{(6.5KB, xml)}

XML Treatment for Trigonopterus singkawangensis

zookeys.467.8206-treatment83.xml^{(5.7KB, xml)}

XML Treatment for Trigonopterus singularis

zookeys.467.8206-treatment84.xml^{(6.5KB, xml)}

XML Treatment for Trigonopterus sinuatus

zookeys.467.8206-treatment85.xml^{(9KB, xml)}

XML Treatment for Trigonopterus squalidus

zookeys.467.8206-treatment86.xml^{(6.4KB, xml)}

XML Treatment for Trigonopterus sumatrensis

zookeys.467.8206-treatment87.xml^{(6.8KB, xml)}

XML Treatment for Trigonopterus sumbawensis

zookeys.467.8206-treatment88.xml^{(8KB, xml)}

XML Treatment for Trigonopterus sundaicus

zookeys.467.8206-treatment89.xml^{(8.3KB, xml)}

XML Treatment for Trigonopterus suturalis

zookeys.467.8206-treatment90.xml^{(8.3KB, xml)}

XML Treatment for Trigonopterus syarbis

zookeys.467.8206-treatment91.xml^{(5.7KB, xml)}

XML Treatment for Trigonopterus telagensis

zookeys.467.8206-treatment92.xml^{(7.1KB, xml)}

XML Treatment for Trigonopterus tepalensis

zookeys.467.8206-treatment93.xml^{(5.3KB, xml)}

XML Treatment for Trigonopterus tiga

zookeys.467.8206-treatment94.xml^{(7KB, xml)}

XML Treatment for Trigonopterus trigonopterus

zookeys.467.8206-treatment95.xml^{(8KB, xml)}

XML Treatment for Trigonopterus tujuh

zookeys.467.8206-treatment96.xml^{(7.2KB, xml)}

XML Treatment for Trigonopterus ujungkulonensis

zookeys.467.8206-treatment97.xml^{(6.6KB, xml)}

XML Treatment for Trigonopterus variolosus

zookeys.467.8206-treatment98.xml^{(6.2KB, xml)}

XML Treatment for Trigonopterus vulcanicus

zookeys.467.8206-treatment99.xml^{(12KB, xml)}

XML Treatment for Trigonopterus wallacei

zookeys.467.8206-treatment100.xml^{(7.4KB, xml)}

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PERMALINK

Ninety-eight new species of Trigonopterus weevils from Sundaland and the Lesser Sunda Islands

Alexander Riedel

Rene Tänzler

Michael Balke

Cahyo Rahmadi

Yayuk R Suhardjono

Abstract Abstract

Introduction

Materials and methods

Systematics

Trigonopterus

Type species.

Diagnosis.

Descriptions of the species

1. Trigonopterus acuminatus

Diagnostic description.

Figure 1.

Material examined.

Distribution.

Etymology.

Notes.

2. Trigonopterus aeneomicans

Diagnostic description.

Figure 2.

Material examined.

Distribution.

Etymology.

Notes.

3. Trigonopterus alaspurwensis

Diagnostic description.

Figure 3.

Material examined.

Distribution.

Etymology.

Notes.

4. Trigonopterus allopatricus

Diagnostic description.

Figure 4.

Material examined.

Distribution.

Etymology.

Notes.

5. Trigonopterus allotopus

Diagnostic description.

Figure 5.

Material examined.

Distribution.

Etymology.

Notes.

6. Trigonopterus angulicollis

Diagnostic description.

Figure 6.

Material examined.

Distribution.

Etymology.

Notes.

7. Trigonopterus amphoralis

Diagnostic description.

Figure 7.

Type material.

Material examined

Distribution.

Notes.

8. Trigonopterus argopurensis

Diagnostic description.

Figure 8.

Material examined.

Distribution.

Etymology.

Notes.

9. Trigonopterus arjunensis

Diagnostic description.

Figure 9.

Material examined.

Distribution.

Etymology.

Notes.

10. Trigonopterus asper

Diagnostic description.