Gyrelon jenpani Hu, Fikacek & Matsumoto, 2024

Hu, Fang-Shuo, Arriaga-Varela, Emmanuel, Biffi, Gabriel, Bocak, Ladislav, Bulirsch, Petr, Damaska, Albert Frantisek, Frisch, Johannes, Hajek, Jiri, Hlavac, Peter, Ho, Bin-Hong, Ho, Yu-Hsiang, Hsiao, Yun, Jelinek, Josef, Klimaszewski, Jan, Kundrata, Robin, Loebl, Ivan, Makranczy, Gyoergy, Matsumoto, Keita, Phang, Guan-Jie, Ruzzier, Enrico, Schuelke, Michael, Svec, Zdenek, Telnov, Dmitry, Tseng, Wei-Zhe, Yeh, Lan-Wei, Le, My-Hanh & Fikacek, Martin, 2024, Forest leaf litter beetles of Taiwan: first DNA barcodes and first insight into the fauna, Deutsche Entomologische Zeitschrift 71 (1), pp. 17-47 : 17

Gyrelon jenpani Hu, Fikacek & Matsumoto sp. nov.

Fig. 2

Type material.

Holotype: male (NMNS): 'Taiwan: Nantou County, Huisun Forest Reserve, track to Xiaochushan Mt., 24.0826139°N, 121.03115869°E, 1050 m, 4.v.2019, Damaška, Fikáček, Hu & Liu lgt., 2019-TW15' (DNA voucher: HS2004). Paratypes: 1 male (NMNS): Taiwan: Nantou County, Huisun Forest Reserve, track to Xiaochushan Mt., 24.0744602°N, 121.0366337°E, 1150 m, 11.x.2020, FS Hu & YJ Chen lgt., old overgrown secondary forest on the slope, sparse understory vegetation: sifting of leaf litter accumulations (DNA voucher: 20-10HS506); 1 male (BMNH): same locality, date and collectors; 1 male (NMNS): same locality data, but 1.iii.2021, Hu, Chen, Fikáček & Peng lgt. (DNA voucher: 21-03HS508); 1 female (NMPC): same locality data, but 24.ii.2020, FS Hu lgt. (DNA voucher: 20-02HS509).

Differential diagnosis.

The new species can be easily recognized from G. rugosus ( Ślipiński, 1982) from southern China by the presence of both coarse and smaller punctures on the pronotum (only small and strongly elongated punctures are present in G. rugosus ). The new species can be recognized from G. mila Hinton, 1942 (Sarawak) and G. sumatrensis Dajoz, 1974 (Sumatra) by the following characters: (1) metaventrite with punctures on the anterolateral portion much larger than posterolaterally (in contrast to small widely separated punctures in both latter species), (2) abdominal ventrite 1 with coarse punctures anteriorly and minute ones in the transverse row (with all punctures moderately large and widely spaced in both latter species), (3) transverse rows on abdominal ventrites 2-5 consisting of minute punctures (moderately large punctures in both latter species). The new species differs from G. compactus Dajoz, 1979 from Singapore by (1) the presence of 8 elytral series (7 in G. compactus ), (2) the serially arranged minute seriferous punctures on elytral intervals 1-2 (with irregularly arranged setiferous punctures in G. compactus ), and (3) the parallel-sided posterolateral margins of the pronotum (posteriorly converging in G. compactus ). The comparison is based on the examination of the holotype of G. mila and two non-type specimens of G. sumatrensis in coll. BMNH (from Perak and Fort de Kock). The types of G. sumatrensis and G. compactus could not be examined as they are lost (A. Mantilleri, pers. comm., March 2023). The types of G. rugosus were not examined as the difference is clear from the original description. We also examined unidentified specimens of Gyrelon from Sumatra, Borneo, Sulawesi and Thailand in coll. BMNH. All of them are similar to G. mila and G. sumatrensis in the characters listed above.

Description.

Body widely oval, body length 2.8-3.2 mm (holotype: 3.2 mm), body width 1.7-1.8 mm (holotype: 1.8 mm) (n=5 including holotype). Dorsal and ventral coloration dark reddish brown to black, legs and antennae brown to reddish brown, all body parts bearing yellowish erect setae.

Head relatively small, eyes moderately large, globular; frons with several moderately large punctures, each bearing erect seta, interstices smooth; clypeus weakly concave on anterior margin, dorsal surface bearing many erect setae. Antenna robust, with 11 antennomeres including the 2-segmented club; antennomeres gradually widening from base to apex; antennomere with microsculptures surface, bearing moderately dense erect setae; antennal club covered by dense short setation and moderately dense set of long erect setae; apex of antennal club bluntly pointed. Mentum small, subtriangular, strongly narrowing anteriad. Apical maxillary and labial palpomeres much narrower than the subapical ones.

Thorax. Pronotum subquadratic, nearly parallel-sided in posterior half, strongly narrowing in anterior half; median part of pronotum with elevated longitudinal ridge. Posterior corners nearly rectangular. Pronotal surface with large irregularly circular or oval punctures, each puncture bearing an erect seta; punctures getting smaller in posterolateral direction. Median part of pronotum lacking punctures, surface between punctures micropunctate. Prosternum widely rectangular, smooth, with coarse deep punctures; prosternal process wide, variable in shape, concave to weakly or strongly trifid posteriorly. Procoxal cavities widely separated, antennal grooves moderately wide, hypomeron with coarse punctures similar to those on prosternum. Mesoventrite anteriorly with a series of longitudinal ridges; surface microsculptured. Mesocoxal cavities widely separated by metaventral process. Each elytron with eight slightly irregular longitudinal series of punctures; serial punctures rounded, lacking setae; additional short series of coarse shallow punctures present anteriorly along elytral side; intervals flat dorsally, slightly convex laterally, smooth, each with a series of widely spaced minute punctures, each bearing erect seta; epipleuron present throughout elytral length, wide anteriorly, gradually narrowing posteriad. Scutellar shield widely triangular. Metaventral process with a narrow median projection of variable shape. Metaventrite flat mesally, lateral portions with large closely adjacent punctures along posterior margin of mesocoxal cavities, otherwise with relatively small and widely separated punctures, each bearing a decumbent seta; interstices with mesh-like microsculpture. Metathoracic wings absent.

Abdomen with 5 visible ventrites, ventrite 1 with a row of large closely adjacent punctures along anterolateral margin, posterior part with a transverse series of minute punctures, each with a decumbent seta. Ventrites 2-5 each with a transverse series of minute punctures, each bearing a decumbent seta. Interstices of all ventrites with fine mesh-like microsculpture. Ventrite 5 sexually dimorphic, with posterior margin nearly smooth in ventral view in male (finely crenulate in posteroventral view), and strongly crenulate in female (with a longitudinally ridged bar situated below apical part of elytral epipleuron).

Legs long and robust. Coxae and trochanters of all three pairs relatively small, coxa subglobular, trochanter subconical. Femora conical, with sparse erect setation, surface with mesh-like microsculpture. Tibiae flat, widening from base to apex, slightly more expanded in apical third, apical part with moderately dense erect setation; apical part of protibiae with an area of dense yellowish hair-like setae mesally. Tarsi with 4 tarsomeres, tarsomere 1 long and thick with dense long setae, tarsomeres 2-3 short, tarsomere 4 the longest.

Male genitalia. Aedeagus 1 mm long, simple, rod-like, without parameres, slightly widened at mid-length, rounded at apex.

Etymology.

The species is dedicated to Dr. Jen-Pan Huang (Biodiversity Center, Academia Sinica, Taipei) as thanks for all his support of this project, including the possibility to work in his lab and for numerous inspiring discussions about evolution, diversity, and beetles.

Distribution.

The species is so far only known from the type locality in central Taiwan.

Notes on diagnostic characters.

Most previous studies use the form of the dorsal punctation and the shape of the prosternal process as the main diagnostic characters. Despite examining a few specimens only, we found both characters, especially the shape of the prosternal process, individually variable and/or dependent on the precise position of observation. The prosternal processes illustrated in Fig. 2D-F belong to the examined specimens whose conspecific identity was confirmed by the cox1 barcode (Fig. 2I ). A slight variation was also observed in the shape of the median projection of the metaventral process. In contrast, the character of the punctation of the metaventrite and abdominal ventrites seems to be much more distinct among species, and does not vary among the examined specimens of the new species.

Archeocrypticidae

Sivacrypticus taiwanicus Kaszab, 1964

Fig. 3

Type material examined. Holotype: female (HNMB): 'Formosa, Sauter’, 'Pilam, 908. II’. We have compared our specimens with the photos of the holotype provided to us by Gy. Makranczy in May 2023 (photos are available in the Zenodo Archive under https://doi.org/10.5281/zenodo.10069183).

New material examined. 1 male (NMNS): Taiwan: Nantou County, Huisun Forest Reserve, beginning of Wading trail, 24.0892139°N, 121.0297836°E, 700 wm, 17.viii.2021, M. Fikáček & WR. Liang, stony disturbed forest on the slope, small leaf accumulations (DNA voucher: 21-08HS169); 1 unsexed specimen (NMPC): same locality data, but 5.v.2019, Damaška, Fikáček, Hu & Liu lgt.; 1 female (NMNS): same locality data, but 28.ii.2021, Hu & Chen lgt. (DNA voucher: 21-03HS119); 1 male, 2 females, 3 unsexed specimens (NMNS, NMPC, BHHC): Taiwan: Taichung, Wufeng, Beikeng Creek trail, 24.0451°N, 120.7827°E, 410 m, 24.v. 2023, lgt. F.S. Hu & Y.J. Chen, lowland tropical forest with large accumulation of leaf litter and sparse understory (TW2023-018, DNA voucher WF1-023 and additional non-sequenced specimens).

Description of male genitalia. Male genitalia 1 mm long. Median lobe thin, strongly bent in the lateral view, with a plate-like expansion on the apex. Tegmen small, freely movable along the median lobe; parameres in lateral view narrowly elongated, pointed at apex and moderately pubescent, in dorsal view plate-like, with a small indentation on lateral margin. Sperm pump present, large, bottle-like, with slightly coiled distal ductus.

Comparison with the holotype. Our specimens correspond to the holotype by all diagnostic characters, including body proportions, the coarse and complete series of punctures on elytra, the double-sized punctation on the pronotum, and the shape of the lateral pronotal margin. On the first view, the specimen in Fig. 3A has much weaker elytral series than the holotype. The additional specimens examined, in which most of the dorsal setation was abraded in the same way as in the holotype, prove that the character of the elytra is in fact identical. In the original description ( Kaszab 1964), as well as in subsequent revisions of the genus ( Kaszab 1979, 1981), the lateral pronotal ridge of S. taiwanicus was mentioned as narrow and not widening anteriad. This stands in contrast to the character state in the holotype as well as in our specimens: the lateral pronotal ridge is, in fact, gradually widening from the base to anterior margin of the pronotum and bends inwards and merges with the anterior margin of the pronotum anteriorly. This fact also corresponds to the illustration of S. taiwanicus by Kaszab (1964) in which the anteriorly widening lateral ridge of the pronotum is clearly seen.

Comparison with other species. The previously unknown male of S. taiwanicus allows us to compare the male genitalia of the species (type species of Sivacrypticus ) with those illustrated for other species of the genus. The male genitalia of S. taiwanicus are very distinct from the genitalia of most described species by (1) small tegmen, (2) strongly elongated median lobe, and (3) strongly expanded parameres in dorsal view. Its genitalia are, however, very similar to those of S. philippinus Merkl, 1988 from Luzon (Manila), but clearly differ from them by the apical expansion of the median lobe in lateral view, and in a less lobate shape of the parameres in dorsal view.

Distribution. The species was described from ‘Pilam’ (= Beinan township, Taitung County, southern Taiwan). The sequenced specimens examined by us are from lowland to lower montane forest in cenrtal Taiwan (Taichung and Nantou Counties), indicating that the species is likely widespread in lowland and lower montane forests at least in central and southern Taiwan.

New records for Taiwan

Since the leaf litter fauna of Taiwan has never been studied in detail, even our small starting dataset from the single area in central Taiwan results in many new records for Taiwan at species, genus or even family levels. Below we concisely report these new records, despite the species-level taxonomic treatment of most of them requiring additional study. The material examined is only listed for taxa identified down to species, for genus-level records, it can be found in the Excel sheet with the complete data (Suppl. material 2). List of all species recorded in this project and identified down to genus or species is available in the Appendix 1.

Carabidae

Oodes ( Lachnocrepis ) Oodes japonicus (Bates, 1873) ( Licininae : Oodini )

Material examined. 1 female (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0847025°N, 121.0274161°E, 1000 m, 20.vi.2020, F.S. Hu lgt., mixed Cryptomeria + sparse broadleaf forest on the slope (voucher 20-06HS304); 1 spec. (IDL): Taiwan: Nantou County, Huisun Forest res., Wading trail, 24.0892139°N, 121.0297836°E, 700 m, 17.viii.2021, M. Fikáček & W.R. Liang lgt., stony forest on the slope, small leaf accumulations (voucher 21-08HS115). 9 spec. (IDL): Taiwan: Kaohsiung City, Zuoying district (左營區), Banpingshan (半屏山), SW slope, 22.694262 120.305072, 100 m, 22.vii.2021, M. Fikáček lgt. (TW2021-06d), sifting of large to shallow leaf accumulations with some wood and fungi in the forest with Ficus in karst area (incl. voucher BP1-001); 17 spec. (IDL): same area and date but 90 m, 22.693469, 120.304979 (TW2021-06f) (incl. DNA voucher BP3-001); 8 spec. (IDL): same area, 90 m, 22.693469, 120.304979, 11.viii. 2022, M. Fikáček lgt. (TW2022-006A) (incl. DNA voucher BP4-010); 2 spec. (IDL): same area and date, 100 m, 22.693469, 120.304979 (TW2022-006B) (incl. DNA voucher BP7-010); 1 spec. (IDL): same area, 22.693469, 120.304979, 90 m, 30.v. 2023, M. Fikáček lgt. (TW2023-015, DNA voucher BP10-002); 1 spec. (IDL): same area, 22.694262, 120.30507 2, 90 m, 30.v. 2023, M. Fikáček lgt. (TW2023-016, DNA voucher BP9-003).

Comments. Multiple species and genera of the Oodini are reported from Japan or southern China ( Guéorguiev 2014; Löbl and Löbl 2017; Guéorguiev and Liang 2020), with only Oodes desertus Motschulsky, 1858 reported from Taiwan so far ( Guéorguiev and Liang 2020). The species barcoded here belongs to Oodes (Lachnotrepis) based on the width of elytral interval 7 and 8 and setation of tarsomeres, and corresponds to O. japonicus based on all characters in the identification key by Guéorguiev and Liang (2020). The species is widespread from the Russian Far East through China and Japan to Laos and Vietnam ( Guéorguiev and Liang 2020). It is recorded from Taiwan for the first time; based on our data it may be widespread in lowland to lower montane forests of central and southern Taiwan. For larval morphology, see below.

Histeridae

Anapleus Horn, 1873 ( Dendrophilinae : Anapleini )

Comments. The genus was first recorded from Taiwan by Bickhardt (1913) based on A. stigmaticus (Schmidt, 1892). Mazur (2007) mentioned that this record might be based on a misidentification and removed the genus and species from his updated list of the Histeridae of Taiwan. The specimen sequenced here is morphologically different from A. stigmaticus ; its identification will be done in the future.

Leiodidae

Dermatohomoeus sp.

Material examined. 4 females (ZSPC): Taiwan: Nantou County Huisun Forest reserve, track to Xiaochushan Mt., 24.0744602°N, 121.0366337°E, 1150 m, 24.ii.2020, F.S. Hu lgt., primary forest on the slope with sparse understory: sifting of small leaf accumulations (incl. DNA voucher 20-02HS511); 17 females (ZSPC): same locality, 20.vi.2020, F.S. Hu lgt. (incl. DNA voucher 20-06HS519); 31 females (ZSPC): same locality, 11.x.2020, F.S. Hu & Y.J. Chen lgt. (incl. DNA voucher 20-10HS521); 19 females (ZSPC): same locality, 16.viii.2021, M. Fikáček & W.R. Liang lgt. (incl. voucher 21-08HS526); 1 female (ZSPC): same locality, 1.iii.2021, M. Fikáček, F.S. Hu & G.J. Peng lgt. (voucher 21-03HS507); 10 females (ZSPC): same locality, 4.v.2019, M. Fikáček, F.S. Hu, A. Damaska & H.C. Liu lgt. (incl. DNA voucher HS1020); 4 females (ZSPC): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0847025°N, 121.0274161°E, 1000 m, 16.viii.2021, mixed Cryptomeria + sparse broadleaf forest on the slope, 16.viii.2021, M. Fikáček & W.R. Liang lgt. (incl. DNA voucher 21-08HS339); 1 female (ZSPC): same locality, 20.vi.2020, F.S. Hu lgt. (voucher 20-06HS317); 1 female (ZSPC): same locality, 11.x.2020, F.S. Hu & Y.J.Chen lgt. (voucher 20-10HS310); 8 females (ZSPC): Taiwan: Nantou County, Huisun Forest Reserve, Xiaochushan Mt. track, 0.5 km above hotels 24.0887444°N, 121.0355063°E, 850 m, 4.v.2019; Damaška, Fikáček, Hu & Liu lgt.; large accumulations of leaf litter in a small gorge with lower montane/lowland broad-leaf forest (incl. voucher HS4031); 2 females (ZSPC): Taiwan: Nantou County, Huisun Forest res., Wading trail, 24.0892139°N, 121.0297836°E, 700 m, 28.ii.2020, F.S. Hu & Y.J. Chen lgt., stony forest on the slope, small leaf accumulations (incl. DNA voucher 21-03HS120); 1 female (ZSPC): same locality, 5.v.2019, M. Fikáček, F.S. Hu, A. Damaska & H.C. Liu lgt. (DNA voucher HS5013).

Comments. The genus is newly recorded from Taiwan in the present paper. The previous records of the genus from Taiwan are based on the transfer of Colenisia miyatakei (Hisamatsu, 1985) to the Dermatohomoeus by Hoshina (1999) that is however not supported by diagnostic characters of Dermatohomoeus ( Švec 2022). Consequently, Dermatohomoeus has not been reported from Taiwan before. All DNA-barcoded specimens from the Huisun Reserve are conspecific, and the examination of additional non-sequenced specimens confirms that all collected specimens are conspecific. Yet, they cannot be identified to species, as all of them are females (in total 99 specimens from 12 collecting events at four different collecting sites). The species of the genus are morphologically uniform, with species-specific characters being the shape of the aedeagus, including the endophallus. Female genitalia and the spermatheca of Dermatohomoeus species are of the unique shape within the tribe Pseudoliodini but lack species-specific morphological features. External morphological characters detectable in Dermatohomoeus females are hardly sufficient for species identifications. The population of Dermatohomoeus consisting exclusively of females found in this study is not the first case of the absence of males. No males have been found so far for Dermatohomoeus terrenus (Hisamatsu, 1985), despite altogether several dozen specimens attributed to this species having been examined ( Hisamatsu 1985; Hoshina 1999; Park and Ahn 2007; Švec 2022). The species is known from the Japanese islands of Honshu, Shikoku, Kyushu, Izu, Goto, from four Ryukyus islands ( Hoshina 1999) and the Awaji Island ( Švec 2022). Besides them, the species was recorded also from southern Korea ( Park and Ahn 2007). Hoshina (1999) published a hypothesis that D. terrenus may be a parthenogenetic species. Perhaps, this type of reproduction is more widespread in Dermatohomoeus species or their populations, including those occurring in Taiwan.

Staphylinidae

Drusilla obliqua (Bernhauer, 1916) ( Aleocharinae : Lomechusini )

Material examined. 12 spec. (FSHC, IDL): same locality, 20.vi.2020, lgt. F.S. Hu (voucher 20-06HS129, and non-extracted specimens); 1 spec. (IDL): same locality, 17.viii.2021, lgt. M. Fikáček & W.R. Liang (voucher 21-08HS133).

Comments. Drusilla obliqua is a widespread species; it has been recorded from India, Nepal, Myanmar, China (Yunnan), Vietnam and Malaysia ( Assing 2017, 2019). The species is newly recorded from Taiwan in the present paper.

Paraploderus cf. thailandicus Makranczy, 2016 ( Oxytelinae : Thinobiini )

Fig. 4

Material examined. 16 spec. (HNHM, IDL): TAIWAN: Nantou County Huisun Forest reserve, track to Xiaochushan Mt., 24.0744602°N, 121.0366337°E; 1150 m 11.x.2020; Hu & Chen lgt., primary forest on the slope with sparse understory: sifting of small accumulations of leaves (DNA voucher 20-10HS531 and non-extracted specimens). 13 spec. (MHNG): TAIWAN: Taoyuan Co. Twnsh.Fushing S-BaLing km 54, road 7, 22.ii.2010 1140m, decaying wood + forest litter, leg. S. Vit #2; 3 spec. (MHNG): TAIW: Chiayi County Alishan Natural Scenic Area, 11.iv.2009 2350m, leg. S. Vit #18//Road 18, km 02 Old Lulin Tree Track, decaying Wood litter #18.

Comments. The genus is newly recorded from Taiwan in the present paper. György Makranczy examined the specimens of this Paraploderus species from Taiwan already earlier, based on the material collected by S. Vít deposited in MHNG (see under Material examined). The male genitalia of these specimens (Fig. 4 ) show rather slight differences from those of Paraploderus thailandicus Makranczy, 2016. Therefore, it requires confirmation whether the Taiwanese populations represent a distinct species or not. This is best done by a comparison of DNA sequences from Taiwan and the mainland, including Thailand from where the species was described.

Thinocharis Kraatz, 1859 ( Paederinae : Lathrobiini )

Comments. The genus is newly recorded from Taiwan in the present paper. The species identification will need to be done in the future.

Tribe Trichonychini ( Pselaphinae )

Comments. The tribe is newly recorded from Taiwan in the present paper, as well as the supertribe Euplectitae . There are at least two species in our samples. A generic revision of the Trichonychini needs to be done before the confirmation of the generic identifications.

Tribe Ctenistini ( Pselaphinae )

Comments. The tribe is here newly recorded from Taiwan. The generic revision of the Ctenistini needs to be done before the confirmation of the generic identifications.

Tribe Bythinoplectini ( Pselaphinae )

Comments. The tribe, as well as the supertribe Euplectitae , are newly recorded from Taiwan here. There are at least two species in our samples.

Batraxis Reitter, 1882 ( Pselaphinae : Brachyglutini )

Comments. The genus was listed for Taiwan in the Catalogue of Life, based on the occurrence of B. obesa Raffray, 1894 ( Chung and Shao 2022). However, the source of the record was online only and the link is not available anymore. We formally record the genus from Taiwan for the first time.

Cephennodes Reitter, 1884 ( Scydmaeninae : Cephenniini )

Comments. The genus is newly recorded from Taiwan in the present paper. The species identification will be done in the future.

Napoconnus Franz, 1957 ( Scydmaeninae : Stenichnini )

Comments. The genus has been newly recorded from Taiwan in the present paper. The species identification will need to be done in the future.

Cerylonidae

Thyroderus porcatus Sharp, 1885 ( Ceryloninae )

Material examined. 2 spec. (FSHC, IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0744602°N, 121.0366337°E; 1150 m, 1.iii.2021, M. Fikáček, F.S. Hu & G.J. Peng lgt. (voucher 21-03HS511 and an additional non-sequenced specimen).

Comments. The species was only known from Japan previously ( Löbl and Smetana 2007), representing the only species of the genus that occurred in the Palearctic region. The genus and the species are newly recorded from Taiwan.

Cautomus Sharp, 1885 ( Ceryloninae )

Comments. The genus is newly recorded from Taiwan in this study based on two species from the Huisun Forest Reserve. Both species differ both by the DNA barcode sequences and morphologically. The species identification will be done in the future.

Sphindidae

Aspidiphorus Ziegler, 1821

Comments. The family and genus are newly recorded from Taiwan. There are two species in our Huisun samples identified by the DNA barcode sequences; their species identification needs to be done in the future.

Latridiidae

Bicava Belon, 1884

Comments. The genus is newly recorded from Taiwan in the present paper. The species identification will be done in the future.

Cartodere sp.

Comments. The genus was first recorded from Taiwan by Yao et al. (2011) based on C. (s. str.) constricta (Gyllenhal, 1827). The specimens sequenced in this study differ from C. (s. str.) constricta by having three antennomeres clubbed (in contrast to two clubbed antennomeres in C. constricta ). The species identification will be done in the future.

Curculionidae

Otibazo Morimoto, 1961

Comments. The genus is newly recorded from Taiwan in the present paper. An extensive taxonomic study on this genus in Taiwan is in preparation and will be published in the near future (Wei-Zhe Tseng, in prep.).

Seleuca Pascoe, 1871

Comments. The genus is newly recorded from Taiwan in the present paper. The species identification needs to be completed in the future.

Acallinus Morimoto, 1962

Comments. The genus is newly recorded from Taiwan in the present paper. Based on the DNA barcodes, the samples reported here (Taiwan: Nantou County, Huisun Forest Reserve) contain two or three species. The species identification needs to be done in the future.

Coccotrypes advena Blandford, 1894

Material examined. 1 female (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0826139°N, 121.0315869°E; 1050 m, 4.v.2019, Damaška, Fikáček, Hu & Liu lgt., sparse secondary forest with dense understory incl. tree ferns on the margin of a tree plantation (voucher HS2015); 1 female (IDL): Taiwan: Nantou County, Huisun Forest res., Xiaochushan Mt. track, 0.5 km above hotels 24.0887444°N, 121.0355063°E, 850 m, 4.v.2019; Damaška, Fikáček, Hu & Liu lgt., large accumulations of leaf litter in a small gorge with lower montane/lowland broad-leaf forest (voucher HS4007); 4 females (IDL): Taiwan: Nantou County, Huisun Forest reserve, Wading trail, 24.0892139°N, 121.0297836°E, 700 m, 11.x.2020, F.S.Hu & Y.J.Chen lgt. (incl. voucher 20-10HS114); 2 females (IDL): same locality, 17.viii.2021, M. Fikáček & W.R. Liang lgt. (incl. voucher 21-08HS170); 1 female (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0847025°N, 121.0274161°E, 1000 m, 11.x.2020, F.S.Hu & Y.J.Chen lgt. (voucher 20-10HS308); 1 female (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0744602°N, 121.0366337°E, 1150 m, 16.viii.2021, M. Fikáček & W.R.Liang lgt. (voucher 21-08HS559).

Comments. This is a generalist seed-boring scolytine species widespread in SE Asia, Australia and Oceania, America from Florida through the Caribbean to Suriname ( Wood and Bright 1992; Bright 2021) and also recorded from Africa (Uganda: Jordal et al. 2002). In Asia, it has been recorded from India, Sri Lanka, Thailand, Vietnam, Indonesia, Malaysia, the Philippines, and Japan; here we are recording it from Taiwan for the first time. Jordal et al. (2002) report a high intraspecific variation of cox1 sequences, possibly indicating that it represents a complex of species. The cox1 sequences of our specimens cluster with those of the Japanese specimen sequenced by Jordal et al. (2002) (uncorrected p -distance to the Japanese specimen: 0.7-1.6%).

Examples of larvae associated with adults

Carabidae

Oodes (Lachnocrepis) japonicus (Bates, 1873) ( Licininae : Oodini )

Fig. 5

Material examined. Larvae: 1 larva (IDL): Taiwan: Nantou County, Huisun Forest res., Wading trail, 24.0892139°N, 121.0297836°E, 700 m, 20.vi.2020, F.S. Hu lgt., stony forest on the slope, small leaf accumulations (voucher 20-06HS179); 1 larva (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0847025°N, 121.0274161°E, 1000 m, 16.viii.2021, M. Fikáček & W.R. Liang lgt., mixed Cryptomeria + sparse broadleaf forest on the slope (voucher 21-08HS350). Adults: see above under New records for Taiwan.

Comments. The knowledge on larval morphology of the Oodini is quite limited so far, with larvae of several species of Oodes Bonelli, 1810 described and illustrated ( van Emden 1942; Lindroth 1942; Chu 1945; Thomson 1979); the larva of an unidentified North American Oodes illustrated by Chu (1945) differs from others in very narrow mandibles, transverse head, multidentate nasale and frontale reaching posterior margin of the head, and may actually represent a different taxon than Oodes or Oodini . The larva of Oodes (Lachnocrepis) japonicus corresponds to Oodes s.str. larvae illustrated by van Emden (1942) and Lindroth (1942) by general morphology, but differs from them in the shape of the nasale ( O. japonicus with 4 sharp teeth, compared to 3 or 5 low rounded teeth in O. helopioides and O. gracilis , respectively), more slender mandibles, shorter and more robust antennomeres, and wider and more robust labial palpomere 2.

Perigona cf. nigriceps Dejean, 1831 ( Lebiinae : Perigonini )

Fig. 6

Material examined. Larvae: 2 larvae (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0847025°N, 121.0274161°E, 1000 m, 20.vi.2020, F.S. Hu lgt., mixed Cryptomeria + sparse broadleaf forest on the slope (voucher 20-06HS344 and one additional specimen). Adults: 1 spec. (IDL): same locality, date and collector (voucher 20-06HS305); 1 spec. (IDL): same locality but 16.viii.2021, M. Fikáček & W.R. Liang lgt. (voucher 21-08HS313).

Comments. Perigona Laporte, 1835 is a species-rich world-wide genus (e.g., Baehr 2014) with larva only illustrated for P. (Xenogona) termitis Jeannel, 1941 ( Jeannel 1941, 1942). Sequenced and examined adult specimens from Huisun belong to the subgenus Perigona Trechicus LeConte, 1853 based on the triangular arrangement of the subapical elytral punctures. Genetically it stands close (uncorrected p -distance 6.4-6.6%) but does not cluster with available sequences of the world-wide invasive P. nigriceps Dejean, 1831 for which DNA barcodes are available from Europe, Africa, South America, the Caribbean and New Zealand in the BOLD database (these moreover form two separate clusters). The larva examined and illustrated here corresponds to that of P. termitis in all characters including the multidentate slightly projecting nasale; it slightly differs from the larva of P. termitis by more robust labial palpomere 1.

Ptilodactylidae : Ptilodactylinae

Ptilodactyla sp.

Fig. 7

Material examined. Larvae: 3 larvae (IDL): Taiwan: Nantou County, Huisun Forest reserve, Wading trail, 24.0892139°N, 121.0297836°E, 700m, 24.ii.2020, F.S. Hu lgt., stony disturbed forest on the slope, small leaf accumulations (incl. sequenced voucher 20-02HS155). Adults: 3 adults (NMPC): same locality, 5.v.2019, Damaška, Fikáček, Hu & Liu lgt. (2019-TW18) (incl. sequenced voucher HS5011).

Comments. Larvae of Ptilodactyla Illiger, 1807 have been mentioned and illustrated by numerous authors (e.g., Costa et al. 1988), including that of P. exotica Chapin, 1927 which is introduced with tropical plants in the USA and Europe (e.g., Aberlenc and Allemand 1997; Mann 2006; Viñolas et al. 2020). Here we are concisely illustrating the sequenced larva of Ptilodactyla from subtropical lowland forest in central Taiwan. The examined specimen has clearly visible proventriculus armored with numerous spines (Fig. 7A ), a structure not yet documented for larval Ptilodactylidae ; we suppose this may be an adaptation for processing the food, indicating that Ptilodactyla larvae likely feed also on decaying wood and detritus, not only on plant roots as stated by some authors (e.g., Lawrence 2005).

Cantharidae : Malthininae

Maltypus ryukyuanus Wittmer, 1970 ( Malthodini )

Fig. 8

Material examined. Larva: 1 larva (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0826139°N, 121.0315869°E; 1050 m, 4.v.2019; Damaška, Fikáček, Hu & Liu lgt., sparse secondary forest with dense understory incl. tree ferns on the margin of a tree plantation: sifting (2019-TW15) (sequenced voucher HS4055L). Adult: 1 specimen (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0847025°N, 121.0274161°E, 1000 m, 20.vi.2020, F.S. Hu lgt., mixed conifer/broadleaf forest + sparse broadleaf forest on the slope (sequenced voucher 20-06HS319).

Comments. In Malthininae , larvae are only known for two genera, Malthinus Latreille, 1806 ( Malthinini ) and Malthodes Kiesenwetter, 1852 ( Malthodini ), with the data about their morphology are scattered. Klausnitzer (1997) assembled all the data and proposed a key to species. Fitton (1976) presented the similarities and differences between both genera. The examined larva of Maltypus Motschulsky, 1860 is similar to that of Malthodes sp. illustrated by Fitton (1976) in the shape of the median tooth of nasale and the absence of setae on the median tooth, but resembles the larva of Malthinus in the inner tooth of the mandible situated more basally. The larva of Maltypus is illustrated for the first time here.

Staphylinidae

Aleocharinae

Drusilla obliqua (Bernhauer, 1916) ( Lomechusini )

Fig. 9

Material examined. Larvae: 2 larvae (IDL): Taiwan: Nantou County, Huisun Forest res., Wading trail, 24.0892139°N, 121.0297836°E, 700 m, 30.vi.2020, F.S. Hu lgt., stony forest on the slope, small leaf accumulations (vouchers 20-06HS176 and 20-06HS187); 2 larvae (IDL): same locality but 17.viii.2021, M. Fikáček & W.R. Liang lgt. (vouchers 21-08HS152 and 21-08HS153). Adults: 1 spec. (IDL): same locality, 20.vi.2020, lgt. F.S. Hu (voucher 20-06HS129); 1 spec. (IDL): same locality, 17.viii.2021, lgt. M. Fikáček & W.R. Liang (voucher 21-08HS133).

Comments. Larvae of two species of Drusilla Leach, 1819 have been described: Drusilla canaliculata (Fabricius, 1787) ( Paulian 1941; Topp 1978; Schminke 1982) and D. italica (Bernhauer, 1903) ( De Marzo 2007). Larvae of all species of Drusilla are very similar and further comparisons are needed to distinguish them.

Myrmecocephalus brevisulcus (Pace, 2008) ( Falagriini )

Fig. 10

Material examined. Larvae: 1 larva (IDL): Taiwan: Nantou County, Huisun Forest res., Wading trail, 24.0892139°N, 121.0297836°E, 700 m, 20.ii.2020, F.S. Hu lgt., stony forest on the slope, small leaf accumulations (voucher 20-02HS154); 2 larvae (IDL): same locality but 11.x.2020, F.S. Hu & Y.J. Chen lgt. (vouchers 20-10HS163-164); 1 larva (IDL): same locality but 28.ii.2021, F.S. Hu & Y.J. Chen lgt. (voucher 21-03HS157); 1 larva (IDL): same locality but 5.v.2019, Fikáček, Hu, Damaška & Liu lgt. (voucher HS5071L); 1 larva (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0847025°N, 121.0274161°E, 1000 m, 4.v.2019; Damaška, Fikáček, Hu & Liu lgt., mixed conifer/broadleaf forest + sparse broadleaf forest on the slope (2019-TW16) (voucher HS3067L); 1 larva (IDL): same locality but 20.vi.2020, F.S. Hu lgt. (voucher 20-06HS348); 1 larva (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0744602°N, 121.0366337°E, 1150 m, 20.vi.2020, F.S. Hu lgt., oldgrown secondary forest on the slope with sparse understory (voucher 20-06HS573); 1 larva (IDL): same locality but 11.x.2020, F.S. Hu & Y.J. Chen lgt. (voucher 20-10HS563). Adults: 1 adult (FSHC): Taiwan: Nantou County, Huisun Forest res., Wading trail, 24.0892139°N, 121.0297836°E, 700 m, 20.ii.2020, F.S. Hu lgt., stony forest on the slope, small leaf accumulations (voucher 20-02HS132); 1 adult (FSHC): same locality but 20.vi.2020 (voucher 20-06HS130); 1 adult (FSHC): same locality but 11.x.2020, F.S. Hu & Y.J. Chen lgt. (voucher 20-10HS135); 1 adult (IDL): same locality but 28.ii.2021, F.S. Hu & Y.J. Chen lgt. (voucher 21-03HS139); 1 adult (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0744602°N, 121.0366337° E, 1150 m, 20.vi.2020, F.S. Hu lgt., oldgrown secondary forest on the slope with sparse understory (voucher 20-06HS533); 1 adult (IDL): same locality but 11.x.2020, F.S. Hu & Y.J. Chen lgt. (voucher 20-10HS529); 1 adult (IDL): same locality but 16.viii.2021, M. Fikáček & W.R. Liang lgt.

Comments. Larvae of several genera of Falagriini have been described or illustrated, including Cordalia Jacobs, 1925, Falagria Leach, 1819 and Myrmecopora Saulcy, 1864 ( Topp 1978; De Marzo 2000, 2002, 2008, 2009). The larva of Myrmecocephalus brevisulcus is similar to that of Myrmecopora by the posterior part of the head becoming remarkably narrower. Myrmecocephalus can be distinguished from the Myrmecopora by the longer and stouter first antennal segment. The larva of Myrmecocephalus is illustrated for the first time here.

Staphylininae

Diochus sp. ( Diochini )

Fig. 11

Material examined. Larva: 1 larva (IDL): Taiwan: Nantou County, Huisun Forest res., Wading trail, 24.0892139°N, 121.0297836°E, 700 m, 20.vi.2020, F.S. Hu lgt., stony forest on the slope, small leaf accumulations (voucher 20-06HS182). Adults: 1 adult (coll. J. Janák, Prague): same locality but 11.x.2020, F.S. Hu & Y.J. Chen lgt. (voucher 20-10HS136); 1 adult (coll. J. Janák, Prague): same locality but 17.viii.2021, M. Fikáček & W.R. Liang lgt. (voucher 21-08HS124); 1 adult (IDL): Taiwan: Kaohsiung City, Zuoying district (左營區), Banpingshan (半屏山), SW slope, 22.694296°N, 120.305797°E, 100 m, 22.vii. 2021, M. Fikáček lgt., sifting of shallow leaf accumulations with some wood and fungi and fallen figs in the forest with Ficus in karst area (TW2021-06e) (voucher BP2-012).

Comments. The tribe Diochini contains two genera: Antarctothius Coiffait & Sáiz, 1969 and Diochus Erichson, 1839; the larva of Antarctothius is unknown. The larva of the American Diochus schaumii Kraatz, 1860 is currently the only known larva in the tribe; it has been mentioned in the phylogenetic study by Solodovnikov and Newton (2005) and listed in the material examined by Irmler (2017), but neither of these works provides a detailed description of the larva. Newton (1990) illustrated an unidentified larva of Diochus from Mexico, which is very similar to the Diochus sp. from Taiwan. Here we document the larva of Diochus sp. which seems to be widespread in lowland forests of Taiwan because this species was found in central (Huisun) and southern Taiwan (Banpingshan) in this study. The adult of this species is similar to one of D. japonicus Cameron, 1930 based on the shorter second antennal segment, but the morphology of aedeagus is completely different. The species identification needs to be done by further comparisons.

Paederinae

Mimopinophilus sp. ( Pinophilini )

Fig. 12

Material examined. Larvae: 1 larva (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0744602°N, 121.0366337°E, 1150 m, 16.viii.2021, M. Fikáček & W.R. Liang lgt., old-grown secondary forest on the slope with sparse understory (voucher 21-08HS568); 1 larva (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0847025°N, 121.0274161°E, 1000 m, 16.viii.2021, M. Fikáček & W.R. Liang lgt., mixed conifer/broadleaf forest + sparse broadleaf forest on the slope (voucher 21-08HS346). Adults: 1 spec. (IDL): same locality, but 24.ii.2020, F.S. Hu lgt. (voucher 20-02HS316); 1 spec. (IDL): same locality, 20.vi.2020, lgt. F.S. Hu (voucher 20-06HS321); 1 spec. (IDL): same locality, 11.x.2020, lgt. F.S. Hu & Y.J. Chen (voucher 20-10HS313).

Comments. The larvae of Pinophilini are poorly understood ( Staniec et al. 2022). Paulian (1941) described and illustrated a larva of Pinophilini from Brazil; however, the genus to which the larva belongs was not determined. Grebennikov and Newton (2009) coded the larval character states of Paederinae from Australia for the phylogenetic work, which is putative as a larva of Pinophilus Gravenhorst 1802. Assing (2022) subdivided the former Pinophilus into several separate genera; the species examined here (as well as all other Taiwanese species) correspond to the recently established Mimopinophilus Assing, 2022.

Nitidulidae

Stelidota multiguttata Reitter, 1877

Fig. 13A-H

Material examined. Larvae: 1 larva (IDL): Taiwan: Nantou County, Huisun Forest res., Wading trail, 24.0892139°N, 121.0297836°E, 700 m, 20.vi.2020, F.S. Hu lgt., stony forest on the slope, small leaf accumulations (voucher 20-06HS169); 1 larva (IDL): same locality, 17.viii.2021, M. Fikáček & W.R. Liang lgt. (voucher 21-08HS158). Adults: 1 spec. (IDL): same locality, 24.ii.2020, lgt. F.S. Hu (voucher 20-02HS116); 1 spec. (IDL): same locality, 11.x.2020, lgt. F.S. Hu & Y.J. Chen (voucher 20-10HS111); 1 spec. (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0847025°N, 121.0274161°E, 1000 m, 24.ii.2020, F.S. Hu lgt., mixed Cryptomeria + sparse broadleaf forest on the slope (voucher 20-02HS302); 1 spec. (IDL): same locality, 11.x.2020, F.S. Hu & Y.J. Chen lgt. (voucher 20-10HS302); 1 spec. (IDL): same locality, 1.iii.2021, lgt. M. Fikáček, F.S. Hu & G.J. Peng; 1 spec. (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0744602°N, 121.0366337°E; 1150 m, 11.x.2020, lgt. F.S. Hu & Y.J. Chen (voucher 20-10HS501).

Comments. The larvae of Nearctic species, Stelidota geminata (Say, 1825), S. ferruginea Reitter, 1873 and S. octomaculata (Say, 1825), have been described ( Peng et al. 1990). The larva of S. multiguttata is very similar to S. geminata ; both species possess longer second antennomere. Further comparision between Stelidota multiguttata and other species is needed to distinguish these similar species.

Lasiodites inaequalis (Grouvelle, 1914)

Fig. 13I-N

Material examined. Larva: 1 spec. (IDL): Taiwan: Nantou County, Huisun Forest res., Wading trail, 24.0892139°N, 121.0297836°E, 700 m, 20.vi.2020, F.S. Hu lgt., stony forest on the slope, small leaf accumulations (voucher 20-06HS172). Adults: 1 spec. (IDL): same locality, 17.viii.2021, lgt. M. Fikáček & W.R. Liang (voucher 21-08HS107); 1 spec. (IDL): same locality, 11.x.2020, lgt. F.S. Hu & Y.J. Chen (voucher 20-10HS110); 1 spec. (NMPC): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0847025°N, 121.0274161°E, 1000 m, 4.v.2019; Damaška, Fikáček, Hu & Liu lgt., mixed conifer/broadleaf forest + sparse broadleaf forest on the slope: sifting (2019-TW16).

Comments. Although the larvae of the invasive Lasiodites picta are sometimes reported in literature (e.g., Serri et al. 2023), the larva of the genus has never been illustrated. Here, we are illustrating an early instar larva of L. inaequalis . It differs from the examined larvae of Stelidota Erichson, 1843 by the form of the urogomphi and by the multidentate mandibles. The species is sometimes placed in Phenolia Erichson, 1943 which comprises similar-looking yet unrelated American species (see Jelínek 1999; Lawrence 2019).

Tenebrionidae : Lagriinae

Lagria scutellaris Pic, 1910 ( Lagriini )

Fig. 14

Material examined. Larvae: 1 larva (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0744602°N, 121.0366337°E; 1150 m, 24.ii.2020, lgt. F.S. Hu, old-grown forest on the slope with sparse understory (voucher 20-02HS537); 1 larva (IDL): same locality, 11.x.2020, lgt. F.S. Hu & Y.J. Chen (voucher 20-10HS556); 1 larva (IDL): Taiwan: Nantou County,

Huisun Forest reserve, Wading trail, 24.0892139°N, 121.0297836°E, 700m, 5.v.2019, Damaška, Fikáček, Hu & Liu lgt., stony forest on the slope, small leaf accumulations (2019-TW18) (voucher HS5060L); 1 larva (IDL): same locality, 24.ii.2020, lgt. F.S. Hu (voucher 20-02HS159); 1 larva (IDL): same locality, 20.vi.2020, lgt. F.S. Hu (voucher 20-06HS167); 1 larva (IDL): same locality, 11.x.2020, lgt. F.S. Hu & Y.J. Chen (voucher 20-10HS159); 1 larva (IDL): same locality, 28.ii.2021, lgt. F.S. Hu & Y.J. Chen (voucher 21-03HS102); 1 larva (IDL): same locality, 17.viii.2021, M. Fikáček & W.R. Liang lgt. (voucher 21-08HS163). Adults: 1 spec. (IDL): same locality, 28.ii.2021, leg. F.S. Hu & Y.J. Chen (voucher 21-03HS102).

Comments. Although some species of Lagria Fabricius, 1775 are recognized as pests and are also used as model organisms and their life cycle is hence well known and studied (e.g., Zhou 1996, 2001; Janke et al. 2022), the larval morphology is rarely illustrated in detail, and are mostly available for European species L. hirta (Linnaeus, 1758) and the invasive African L. villosa Fabricius, 1781 (see Spilman 1978 and online resources). We illustrate the larva of the Taiwan-endemic L. scuttelaris as it is often a dominant larval morphotype in forest leaf litter samples in Taiwan. It resembles the larva of L. hirta by dorsal color patterns (in contrast to uniformly black larva of L. villosa ), but differs from it by larger and more widely separated urogomphi (very small and closely situated in L. hirta ).

Anaedus spinicornis Kaszab, 1973 ( Goniaderini )

Fig. 15

Material examined. Larvae: 1 larva (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0847025°N, 121.0274161°E, 1000 m, 20.vi.2020, F.S. Hu lgt., mixed conifer/broadleaf forest + sparse broadleaf forest on the slope (voucher 20-06HS334); 1 larva (IDL): same locality, 4.v.2019, Damaška, Fikáček, Hu & Liu lgt. (2019-TW16) (voucher HS3055L); 1 larva (IDL): Taiwan: Nantou County, Huisun Forest reserve, track to Xiaochushan Mt., 24.0744602°N, 121.0366337°E; 1150 m, 20.vi.2020, lgt. F.S. Hu, old-grown forest on the slope with sparse understory (voucher 20-06HS557); 1 larva (IDL): same locality, 4.v.2019, Fikáček, Hu, Damaška, Liu lgt. (voucher HS1062L). Adults: 1 spec. (IDL): same locality, 24.ii.2020, F.S. Hu lgt. (voucher 20-02HS502); 1 spec. (IDL): same locality, 20.vi.2020, lgt. F.S. Hu (voucher 20-06HS501).

Comments. The larva of American Anaedus brunneus (Ziegler, 1844) has been illustrated without a detailed description ( Böving and Craighead 1930). Arndt (1993) described the African species Anaedus camerunus Gebien, 1920. The larvae of A. spinicornis can be distinguished from the two known species by a relatively shorter and broader head. It can also be distinguished from A. camerunus by the coloration without a pair of spots on the anterior portion of the pronotum and with longer stripes on the lateral portion of the meso- and metanota.

Figures

Figure 2. Gyrelon jenpani sp. nov. (Cerylonidae). A, B. Habitus (A. Dorsal view, female; B. Ventral view, male); C. Tegmen of the aedeagus; D-F. Variability of the shape of the prosternal process and metaventral process (D, E. Males; F. Female). Last abdominal ventrite in ventral and postero-ventral views (G. Male; H. Female); I. Maximum likelihood tree based on cox 1 barcodes of the sequenced Cerylonidae specimens.

Figure 3. Sivacrypticus taiwanicus Kaszab, 1964 (Archeocrypticidae). A-C. Habitus (A. Dorsal; B. Lateral; C. Ventral); D. Abdominal ventrites, male; E-H. Male genitalia (E. Median lobe and the sperm pump, lateral view; F. Median lobe and parameres, lateral view; G. Detail of median lobe and parameres, lateral view; H. Detail of parameres, dorsal view).

Figure 4. Male genitalia of Paraploderus cf. thailandicus Makranczy, 2016 from Taiwan. A. Frontal view (parameral setation shown on the left); B. Lateral view.

Figure 5. Carabidae: Oodini: larva of Oodes (Lachnocrepis) japonicus (Bates, 1873) (OTU 159, voucher 20 - 08 HS 350) associated with adults by DNA. A, B. Head (A. Dorsal view; B. Ventral view); C. Middle and hind legs; D. Nasale and mandibles, dorsal view; E. Antenna, dorsal view; F. Mouthparts, ventral view; G, H. Abdominal apex (G. Dorsal view; H. Ventral view).

Figure 6. Carabidae: Perigonini: larva of Perigona cf. nigriceps Dejean, 1831 (OTU 158, voucher 20 - 06 HS 344) associated with adults by DNA. A, B. Head (A. Ventral view; B. Dorsal view); C. Antenna; D. Mouthparts, ventral view; E. Nasale; F. Thorax, dorsal view; G. Middle leg; H. Abdominal apex.

Figure 7. Ptilodactylidae: larva of Ptilodactyla sp. (OTU 83, voucher 20 - 02 HS 155) associated with adults by DNA. A. Head and thorax in dorsal view; B. Head, ventral view; C. Head, lateral view; D. Anterior part of the head, dorsal view; E. Detail of labrum; F. Antenna in lateral view; G. Front and middle leg.

Figure 8. Cantharidae: larva of Maltypus ryukyuanus (OTU 66, voucher HS 4055 L) associated with adults by DNA. A. Head and pro- and mesothorax, ventral view; B. Head, dorsal view; C. Detail of anterior part of the head, dorsal view; D. Detail of the head surface, with smooth anterior and sculptured posterior part; E. Antenna; F. Front leg.

Figure 9. Staphylinidae: Aleocharinae: Lomechusini: larva of Drusilla obliqua. (OTU 216, voucher 21 - 08 HS 152) associated with adults by DNA. A. Dorsal habitus; B-F. Head: B. Dorsal view; C. Ventral view; D. Details of anterior part in dorsal view; E. Maxilla; F. Labium; G. Abdominal apex in ventral view; H. Hind legs.

Figure 10. Staphylinidae: Aleocharinae: Falagriini: larva of Myrmecocephalus brevisulcus (OTU 84, voucher 20 - 06 HS 573) associated with adults by DNA. A. Head and prothorax in ventral view; B. Head in dorsal view; C. Clypeus, labrum and mandibles in dorsal view; D. Mouthparts in ventral view; E. Antenna; F. Thorax in dorsal view; G. Hind leg; H. Abdominal apex.

Figure 11. Staphylinidae: Staphylininae: Diochini: larva of Diochus sp. (OTU 206, voucher 20 - 08 HS 182) associated with adults by DNA. A-D. Head: A. Dorsal view; B. Ventral view; C. Details of anterior part in ventral view; D. Antenna; E. Thorax in dorsal view; F. Fore leg; G. Apex of abdomen in lateral view.

Figure 12. Staphylinidae: Paederinae: Pinophilini: larva of Mimopinophilus sp. (OTU 271, vouchers 21 - 08 HS 346 and 21 - 08 HS 568) associated with adults by DNA. A. Head and anterior part of thorax, dorsal view; B. Maxilla; C. Antenna; D. Nasale; E. Eye in lateral view; F. Labium; G. End of the abdomen with urogomphi, dorsal view; H. Front legs.

Figure 13. Nitidulidae. A-G. Late instar larva of Stelidota multiguttata (OTU 119, voucher 21 - 08 H 158); H-M. Early instar larva of Lasiodites inaequalis (OTU 180, voucher 20 - 08 HS 172). A, I. Habitus in dorsal view; B. Abdominal apex in lateral view; C. Head in dorsal view; D, K. Antenna; E. L. Mouthparts in ventral view; F. Head and thorax in ventral view; G, M. Detail of leg; H, N. End of abdomen in dorsal view; K. Detail of antenna and mandible in dorsal view.

Figure 14. Tenebrionidae: Lagriinae: Lagriini: larvae of Lagria scutellaris (OTU 174) associated with adults by DNA. A-C. Habitus of late instar larva (voucher 20 - 10 HS 556; A. Dorsal; B. Lateral; C. Ventral); D, E. Head of the late instar larva (voucher 21 - 08 HS 163; D. Dorsal; E. Ventral); F-H. Early instar larva (voucher HS 5060 L): F. Mouthparts, ventral view; G. Hind leg; H. Head and thorax in dorsal view.

Figure 15. Tenebrionidae: Lagriinae: Goniaderini: larva of Anaedus spinicornis (OTU 49, voucher HS 1062 L) associated with adults by DNA. A-C. Habitus (A. Dorsal; B. Ventral; C. Lateral); D-F. Head (D. Dorsal; E. Ventral; F. Ventral, close-up).