Exploring the Caste-Specific Multi-Layer Defense Mechanism of Formosan Subterranean Termites, Coptotermes formosanus Shiraki.

The survival and foraging of Coptotermesformosanus Shiraki in a microbe-rich environment reflect the adaptation of an extraordinary, sophisticated defense mechanism by the nest-mates. We aimed to explore the host pathogen interaction by studying caste-specific volatile chemistry and genes encoding the antioxidant defense of winged imagoes, nymphs, soldiers and workers of Formosan subterranean termites. Qualitative analyses of C.formosanus Shiraki performed by HS-SPME/GC-MS showed considerable variations in the chemical composition of volatile organic compounds (VOCs) and their proportions among all the castes. Winged imagoes produced the most important compounds such as naphthalene and n-hexanoic acid. The antifungal activity of these compounds along with nonanal, n-pentadecane, n-tetradecane, n-heptadecane and methyl octanoate against the conidial suspensions of Metarhiziumanisopliae and Beauveriabassiana isolates enable us to suggest that the failure of natural fungal infection in the nest is due to the antiseptic environment of the nest, which is mainly controlled by the VOCs of nest-mates. In addition, conidial germination of M.anisopliae and B.bassiana isolates evaluated on the cuticle of each caste showed significant variations among isolates and different castes. Our results showed that the conidia of M.anisopliae 02049 exhibited the highest germination on the cuticle of all the inoculated castes. Moreover, we recorded the lowest germination of the conidia of B.bassiana 200436. Caste-specific germination variations enabled us to report for the first time that the cuticle of winged imagoes was found to be the most resistant cuticle. The analysis of the transcriptome of C.formosanus Shiraki revealed the identification of 17 genes directly involved in antioxidant defense. Expression patterns of the identified antioxidant genes by quantitative real-time PCR (qPCR) revealed the significantly highest upregulation of CAT, GST, PRXSL, Cu/Zn-SOD2, TXN1, TXN2, TXNL1, TXNL2, TXNL4A and TPx genes among winged imagoes upon infection with the most virulent isolate, M.anisopliae 02049. Furthermore, soldiers showed the least expression of genes encoding antioxidant defense. Our findings indicated that the volatile chemistry of nest-mates and genes encoding antioxidant defense greatly contribute to the survival and foraging of Formosan subterranean termites in a microbe-rich habitat.

Reproductive traits of Glenea cantor (Coleoptera: Cerambycidae: Lamiinae).

Glenea cantor (F.) is a cerambycid species that attacks living trees of at least seven plant families in Asia, and has the potential to become an invasive pest. Here we investigated its reproductive biology in the laboratory to provide vital information for the development of pest control measures and preparation of pest risk analysis. Both sexes required a period of maturation feeding before mating, with the mean premating period of males (5.87 +/- 1.68 d) being significantly longer than that of females (4.59 +/- 2.34 d). This was a synovigenic species with the mean preoviposition and oviposition period being 13.50 +/- 3.15 d and 49.46 +/- 16.16 d, respectively. If no food was available after emergence, adult males died before reaching sexual maturation and females vanished before oviposition. Paired couples mated an average of 15.12 +/- 7.02 times; 28- to 53-d-old beetles mated significantly more frequently than younger and older ones. The mean fecundity was 133.24 +/- 9.67 eggs. The oviposition rate significantly increased in the first 5 wk of the oviposition period, peaking when females were 40- to 54-d-old, and then significantly decreased in the following weeks. Reproductive activities occurred almost exclusively during the day, with most ovipositions taking place in the morning and matings in the afternoon with some overlaps.

Four new cavernicolous species of the millipede genus Eutrichodesmus Silvestri, 1910 from southern China (Diplopoda: Polydesmida: Haplodesmidae).

Four new species of the large oriental genus Eutrichodesmus Silvestri, 1910 are described from caves in southern China: E. digitatus sp. n. (Guangdong), E. simplex sp. n. (Jiangxi), E. spinatus sp. n. (Hunan) and E. planatus sp. n. (Guangxi).

Three new species of the genus Brachinus Weber, 1801 from the Philippines, with a modified key to species (Coleoptera: Carabidae: Brachininae).

Three new Brachinus species namely B. grootaerti n. sp. (Palawan Island), B. palawanensis n. sp. (Palawan Island), B. mindanaoensis n. sp. (Mindanao Island) from Philippines were described. In addition, a modified key to the known Brachinus Philippine species is provided.

A contribution to cavernicolous beetle diversity of South China Karst: eight new genera and fourteen new species (Coleoptera: Carabidae: Trechini).

The specific diversity of Chinese cave fauna of the ground beetle tribe Trechini is further enriched. Eight new genera and species of trechine beetles are reported from limestone caves in southern China karsts: Youtrechus zijunae n. gen., n. sp. (Xianning, southeastern Hubei Province), Wumengius fengi n. gen., n. sp. (Liupanshui, western Guizhou Province), Chenotrechus parvulus n. gen., n. sp. (Laibin, central Guangxi Zhuang Autonomous Region), Sinaphaenopoides zhaoyiae n. gen., n. sp. (Chenzhou, southern Hunan Province), Ecotrechus ruruiae n. gen., n. sp. Hurites yunxianicus n. gen., n. sp. (Shaoyang, central Hunan Province), Pangaphaenops seraphicus n. gen., n. sp. (Bama, northwestern Guangxi Zhuang Autonomous Region) and Zunotrechus zhaojieyui n. gen., n. sp. (Zunyi, northern Guizhou Province). All the above genera are monospecific. Generic diagnosis of each new genus, with discussions on its possible phylogenetic position within the tribe Trechini, and description of the type species are provided. The following new species of cavernicolous trechines are described: Dongodytes (Dongodytodes) pingjingae sp. n. (Hechi, northern Guangxi Zhuang Autonomous Region), Dongodytes (s. str.) mingruoae n. sp. (Du'an, Hechi, northern Guangxi Zhuang Autonomous Region), Deuveaphaenops (Furongius) huangheae n. sp. (Daozhen, Zunyi, northern Guizhou Province), Shilinotrechus zhangfani n. sp. (Mile, Honghe Hani & Yi Autonomous Prefecture, southeastern Yunnan Province), Guizhaphaenops (s. str.) wanghaoi n. sp. (Qinglong, Qianxinan Buyi & Miao Autonomous Prefecture, southwestern Guizhou Province), and Graciliblemus gui n. sp. (Liping, Qiandongnan Miao & Dong Autonomous Prefecture, southeastern Guizhou Province). Further generic characteristics of the genus Graciliblemus Deuve & Tian, 2016 are added. A distribution map for all above new taxa is given.

Discovery of a new cavernicolous trechine genus and species from a deep pit in Anlong, southwestern Guizhou Province (Coleoptera: Carabidae: Trechinae).

Ckacus youcha n. gen. n. sp. is described and illustrated based on a single male discovered in the deep area of Luoshui Dong, a vertical cave in Anlong Xian, southwestern most of Guizhou Province, Southwest China. This new genus seems related to Bathytrechus Uéno, 2005 from Leye Xian, northwestern most Guangxi by its elongate body which is wholly pubescent, the tridentate right mandible and the well-striated elytra with the 1st interval extremely narrowed at apical 1/3, but differs in several important characters. In addition, three individuals of the species Satotrechus rieae Uéno, 2006 were also collected from -340 m to -450 m in the same cave. This species was formerly only known from four caves in Anlong and Ceheng counties.

Two new genera and three new species of cavernicolous trechines from the western Wuling Mountains, China (Coleoptera, Carabidae, Trechinae).

Two new genera and three new species of cave-adapted ground beetles belonging to the tribe Trechini are established and described: Wulongiusqilinger gen. nov. and sp. nov. from limestone cave Qiankou Dong (Chongqing: Wulong), Qianotrechuscongcongae sp. nov. from cave Shigao Dong (Chongqing: Nanchuan), and Qianlongiuszhoui gen. nov. and sp. nov. from cave Qianlong Dong (Guizhou: Songtao). Wulongiusqilinger sp. nov. is a small aphaenopsian beetle with a thin and elongated body, while Qianlongiuszhoui sp. nov. is a semi-aphaenopsian with a stout body. Both new genera are not closely related to any genus of Trechini occurring in the South China Karst, and so their systematic positions remain unclear.

A contribution to the knowledge of cave-adapted ground beetles from Guiyang, central Guizhou Province, southwestern China (Coleoptera, Carabidae, Trechini).

A new genus and two new species of cavernicolous trechines are reported from central Guizhou Province, southwestern China. Haixiaphaenops gen. nov. is established to place a new species discovered in two limestone caves in northern Qingzhen Shi: H.jinxiaohongae sp. nov. (Dawan Dong cave and Changtu Dong and Dawan Dong caves). This new genus is allied to Zhijinaphaenops Uéno & Ran, 2002. Zhijinaphaenopszhaofeii sp. nov. is described from Zhangkou Dong cave in northern Jiuzhuang Zhen of Xifeng County. In addition, two new localities of the species Zhijinaphaenopsjingliae Deuve & Tian, 2015, and two new localities of SinaphaenopschengguangyuaniMa et al. 2020 are provided. A distribution map for all cavernicolous trechine beetles known in Guiyang is provided.

Contribution to the knowledge of subterranean ground beetles from eastern Wuling Mountains, China (Coleoptera: Carabidae: Trechinae).

Two new genera and three new species of troglobitic trechine beetles are described and illustrated from the eastern part of Wuling Mountains: Xiangxius jiangi n. gen., n. sp. and Guizhaphaenopsodes solidior n. gen., n. sp. from the limestone cave Tangle Dong (Jishou, western Hunan Province); Sinotroglodytes hefengensis n. sp. from the limestone cave Qizimei Dong (Hefeng, southwestern Hubei Province). Xiangxius jiangi is a highly modified cave-adapted beetle with elongate body and appendages, recognized by its peculiar elytral shape and remarkable dorsal and marginal umbilicate pores of elytra. Guizhaphaenopsodes solidior is peculiar for its stout body, robust antennae and pigmented forebody, antennae and legs though it shares several generic important characteristics with the genus Guizhaphaenops Vigna Taglianti, 1997. Sinotroglodytes hefengensis has an elongated and parallel-sided head.

Morphological Adaptation of Cave-Dwelling Ground Beetles in China Revealed by Geometric Morphometry (Coleoptera, Carabidae, Trechini).

Cave-dwelling ground beetles in China represent the most impressive specific diversity and morphological adaptations of the cavernicolous ground beetles in the world, but they have not been systematically examined in quantitative terms. The present study focuses on the application of geometric morphological methods to address the morphological adaptations of the tribe Trechini, the most representative group in China. We have employed a geometric morphometry analysis of the head, pronotum, and elytra of 53 genera of Trechini, including 132 hypogean and 8 epigean species. Our results showed that the overall morphological variation of cave carabids has gradually specialized from an anophthalmic to semi-aphaenopsian to aphaenopsian type. There were extremely significant differences (p < 0.01) among four different adaptive types including aphaenopsian, semi-aphaenopsian, anophthalmic, and surface-dwelling Trechini when their adaptability to a cave environment was used as the basis for grouping. Furthermore, there were differences in the phenotypic tree of the head, pronotum, and elytra, and an integrated morphology. To the best of our knowledge, this is the first report on the analysis of the head, pronotum, and elytra of four different adaptive types of ground beetles in order to clarify the morphological adaptations of cavernicolous carabids to the cave environment.

Differential fluctuation in virulence and VOC profiles among different cultures of entomopathogenic fungi.

Insect-passaged cultures of entomopathogenic fungi grown on potato dextrose agar media have been shown to have altered virulence and profiles of volatile compounds. The present study demonstrated the pathogenic status of FS(0) (in vitro) and FS(1) and FS(2) (insect-passaged cultures grown on PDA) cultures of Metarhizium anisopliae (strains 406 and 02049) and Beauveria bassiana by a non-choice assay, in which filter paper was inoculated with fungal spores at a concentration of 1 x 10(7) spores/ml. The FS(1) and FS(2) cultures of M. anisopliae strain 02049 and B. bassiana produced conidia with high virulence, and the volatile profiles of these conidia comprised relatively lower percentages of branched-alkanes than conidia from the FS(0) cultures. In contrast, the conidia from an FS(0) culture of M. anisopliae strain 406 had somewhat elevated virulence levels, but their volatile profile had <2% branched-alkanes. The FS(1) and FS(2) cultures of M. anisopliae strain 406 did not gain virulence, and these cultures showed a decline in virulence along with major alteration of their volatile profiles. Their volatile profiles mainly comprised branched-alkanes. The volatile profiles of the FS(1) and FS(2) cultures lacked n-tetradecane, which was an important component of all the virulent cultures. Four compounds, 2-phenylpropenal, 2,5,5-trimethyl-1-hexene, n-tetradecane and 2,6-dimethylheptadecane, were detected only from the virulent cultures, suggesting that low LT(50) values were probably due to the production of these compounds. This is the first report to characterize volatiles from FS(0), FS(1) and FS(2) cultures of entomopathogenic fungi; its utility in different aspects opens an interesting area for further investigations.

A contribution to the knowledge of cavernicolous ground beetles from Sichuan Province, southwestern China (Coleoptera, Carabidae, Trechini, Platynini).

Two new genera and four new species of cave-adapted ground beetles are described from Sichuan Province, southwestern China. Uenoaphaenops gen. nov. is established to place the trechine species Qianotrechus fani Uéno, 2003 occurring in the limestone cave Hua'er Dong, southeastern Sichuan (Luzhou: Gulin). Chu pheggomisetoides gen. nov. & sp. nov., from the limestone cave Hanwang Dong, northeastern Sichuan (Guangyuan: Chaotian), is somewhat like the European cavernicolous trechine genus Pheggomisetes Knirsch, 1923, from Bulgaria and Serbia, in particular in the configurations of head and pronotum. Boreaphaenops liyuani sp. nov., also from Hanwang Dong, is the second representative of the genus and the first record in Sichuan. Agonotrechus sinotroglophilus Deuve, 1999, a troglophile, is reported from Sichuan for the first time. The other two new species belong to the platynine genus Jujiroa Uéno, 1952: J. uenoi sp. nov. from the cave Banche Dong on the northern side of the Dadu River (Leshan: Shawan) and J. wangzheni sp. nov. from the cave Hua'er Dong, which is sympatric with Uenoaphaenops fani (Uéno, 2003) comb. nov. A distribution map for the localities of all abovementioned caves and a key to Jujiroa species known in Sichuan are provided.

New species and new record of cavernicolous ground beetles from Sichuan Province, China (Coleoptera: Carabidae: Pterostichini and Platynini).

Two new species of cave-adapted ground beetles are described and illustrated from Sichuan Province, southwestern China: Jujiroa zhouchaoi sp. nov. from the calcareous conglomerate cave Shenxian Dong in Mt. Zhaogong (Dujiangyan, Chengdu) and Pterostichus (Huaius) hanwang sp. nov. from the limestone cave Hanwang Dong (Chaotian, Guangyuan). A new locality for Jujiroa deliciola Uéno and Kishimoto, 2001 is given. Keys to species of the genus Jujiroa Uéno, 1952 and the subgenus Huaius Tian and Huang, 2019 (in Tian et al. 2019) are also provided.

New cavernicolous ground beetles from Anhui Province, China (Coleoptera, Carabidae, Trechini, Platynini).

A new genus and three new species of cave-adapted ground beetles are reported from the limestone cave Shenxian Dong in Huangshan Shi, southeastern Anhui Province, China. Shenoblemus gen. nov. is proposed to place the anophthalmic trechine species S. minusculus sp. nov. This genus is characterized by the tiny but stout body, sub-moniliform antennae, serrated elytral margins near the base, and a wide distance between the fifth and sixth pores of the marginal umbilicate series on the elytra. In addition, two new species, Wanoblemus huangshanicus sp. nov. (an anophthalmic trechine) and Jujiroa inexpectata sp. nov. (a microphthalmic platynine), are also described and illustrated from the same cave.

A new anophthalmic trechine genus and two new species from southern Guizhou, China (Coleoptera: Carabidae: Trechini).

A new genus and two new species of cavernicolous trechine beetles are reported from southern Guizhou Province, Southwest China: Miaotrechus mahua n. gen., n. sp. from the cave Miaoting, Getuhe cave system, Ziyun Miao Buyi Zizhi Xian (Autonomous County), Anshun Shi; M. heweii n. sp. from the cave Jingua Dong, danzhai County, Qiandongnan Miao Dong Zizhizhou (Autonomous Prefecture). Miaotrechus might be related to the genus Guizhaphaenops Vigna Taglianti, 1997, but it is an Anophthalmus-like, whereas Guizhaphaenops species are semi-aphaenopsian.

Two new species of cavernicolous trechines from central Guizhou and northwestern Guangxi, China (Coleoptera: Carabidae: Trechinae).

Two highly modified new species of cave-adapted trechine beetles are described from Guizhou Province and Guangxi Zhuang Autonomous Region: Sinaphaenops (Sinaphaenops) chengguangyuani n. sp. (Guizhou: Longli: cave Shuijing Dong), belonging to the banshanicus species group of the genus Sinaphaenops Uéno Wang, 1991; Giraffaphaenops brevicephalus n. sp. (Guangxi: Leye: cave Dalongguan Dong), being the third species of the genus Giraffaphaenops Deuve, 2002, with a shorter but broader head than its congeners. Keys to identify the species of the banshanicus species group and the genus Giraffaphaenops are provided respectively.

Discovery of a cavernicolous trechine species from the Sanwang Dong-Erwang Dong cave system, Wulong, Chongqing, southwestern China (Coleoptera: Carabidae: Trechinae).

Qianaphaenops (Sanwangius) rowselli Tian et Chen, n. subgen., n. sp. is described from the Sanwang Dong-Erwang Dong cave system, northeastern Wulong, southern Chongqing, southwestern China. Sanwangius is markedly distinct by several important features from subgenera Qianaphaenops (s. str.) Uéno, 2000 and Tiankengius Tian et Huang, 2018. Larval characteristics of this new species are also provided. This is the first report of a Qianaphaenops species from Chongqing.

Contributions to the knowledge of the genus Tonkinaphaenops Deuve, 2013 from China and Vietnam (Coleoptera: Carabidae: Trechinae).

Cavernicolous trechine beetles of the genus Tonkinaphaenops Deuve, 2013, known so far only from Vietnam, are reported for the first time from southern China, and new data are also provided regarding this genus from northern Vietnam. Two new Chinese species, T. yinquanicus sp. nov. from cave Yinquan Dong and T. jingxicus sp. nov. from cave Nianluo Dong in Jingxi County, southwesternmost Guangxi Zhuang Autonomous Region, are described and illustrated, together with the descriptions of two new Vietnamese species: T. anthonyi sp. nov. from the cave Hang Dõi and T. impunctatus sp. nov. from the cave Hang Ran in Cao Bang province. A key to all known species of Tonkinaphaenops is also provided.

Contribution to the knowledge of the cavernicolous beetle genus Aspidaphaenops Uéno from Guizhou (Coleoptera: Carabidae: Trechinae).

The hypogean ground beetle genus Aspidaphaenops Uéno, 2006 is reviewed. Some new information of A. reflexus Uéno, 2006 is provided, including the male genital characteristics as well as a new locality. Two new species are described and illustrated from southwestern Guizhou Province, southwest China: A. xiongda n. sp. (Guizhou: Anlong: Fengyan Dong), A. dudou n. sp. (Guizhou: Xingyi: Xiaozi Dong). A key to all Aspidaphaenops species is also provided.

A contribution to Dongodytes (s. str.) Deuve, 1993 (Coleoptera, Carabidae, Trechinae).

The hypogean genus Dongodytes Deuve, 1993, one of the most cave-adapted genera of ground beetles, is distributed in northern Guangxi, ranging from Mashan through Du'an, Bama, and Fengshan to Tian'e. Review of nominate subgenus Dongodytes Deuve, 1993, with new records for D. fowleri Deuve, 1993 and D. grandis Uéno, 1998 are provided. Meanwhile, Dongodytes tonywhittenisp. n. is described from a limestone cave in Bama County. This interesting species is dedicated to the late Dr. Tony Whitten, a well-known cave biodiversity conservationist in Asia. A key to all species of Dongodytes (s. str.) is also provided.