Illuminating the darkness: an exceptionally diverse fauna of subterranean Pselaphinae (Coleoptera: Staphylinidae) in Asian karsts

Abstract Subterranean ecosystems, particularly the karst environments of Asia, represent a significant but underexplored frontier for biodiversity exploration. This paper focuses on the rove beetle subfamily Pselaphinae (Coleoptera: Staphylinidae) from caves and related subterranean habitats throughout the continent. Drawing upon extensive material accumulated primarily from China and Thailand, supplemented by records from Vietnam, Laos, and the Philippines, an exceptionally diverse subterranean fauna is herein delineated, including the description of 43 new species: Araneibatrus antennatus sp. nov., A. huangsunbini sp. nov., A. leclerci sp. nov., A. thamluang sp. nov., A. thamphamon sp. nov., A. thamphathong sp. nov., A. thamprakaiphet sp. nov.; Batrisiella longlina sp. nov.; Batrisocenus affinis sp. nov., Batrisocenus dushizi sp. nov., Batrisocenus filum sp. nov., Batrisocenus mojiachengi sp. nov., Batrisocenus shilin sp. nov., Batrisocenus trungtrang sp. nov., Tribasodites circinatus sp. nov., T. disuensis sp. nov., T. duanus sp. nov., T. guilinensis sp. nov., T. huchanghaoi sp. nov., T. miaoting sp. nov., T. pakchong sp. nov., T. shui sp. nov., T. uncinus sp. nov., T. zhouhanpingi sp. nov.; Zopherobatrus excavatus sp. nov., Z. kaiyangus sp. nov., Z. sinanus sp. nov., Z. wulongensis sp. nov.; Batraxis soparki sp. nov.; Nipponobythus haozhu sp. nov., N. jinji sp. nov., N. liyunchuni sp. nov., N. long sp. nov., N. minor sp. nov. (soil-dwelling), N. niubi sp. nov., N. platycephalus sp. nov., N. proiectus sp. nov., N. punctatus sp. nov., N. qian sp. nov., N. sanxian sp. nov., N. suxian sp. nov., N. wufengensis sp. nov., and N. zengtingkaii sp. nov. The present work nearly doubles the documented diversity of cavernicolous Pselaphinae in Asia, increasing the number of known species from 48 to 90, highlighting the profound geographical biases in previous research and the immense, cryptic diversity harbored within these habitats. Identification keys to supertribes, tribes, genera, and species are provided, along with a checklist of cavernicolous Pselaphinae recorded from Asia. Moreover, this study reports novel instances of pronounced male antennal polymorphism and identifies potential morphological correlates of sexual conflict, specifically, putative female resistance traits, within these cave-adapted lineages, thereby illuminating evolutionary dynamics operative in aphotic ecosystems. Collectively, this work establishes a crucial taxonomic framework for the Asian cavernicolous fauna and stresses the urgent need for continued biospeleological exploration to effectively catalogue and conserve the unique biodiversity endemic to these karst systems. 地下生态系统, 特别是亚洲喀斯特环境, 构成了生物多样性探索中一片重要但尚未充分研究的领域。本研究聚焦亚洲洞穴及相关地下生境的蚁甲亚科（鞘翅目：隐翅虫科）, 以中国和泰国的大量标本为主要材料, 辅以越南、老挝和菲律宾的记录, 系统描述了高度多样化的地下动物区系, 包含43新种：弯角蛛蚁甲Araneibatrus antennatus sp. nov., 黄氏蛛蚁甲A. huangsunbini sp. nov., 勒氏蛛蚁甲A. leclerci sp. nov., 銮蛛蚁甲A. thamluang sp. nov., 帕蒙蛛蚁甲A. thamphamon sp. nov., 帕通蛛蚁甲A. thamphathong sp. nov., 佩卡菲特蛛蚁甲A. thamprakaiphet sp. nov., 隆林小毛唇蚁甲Batrisiella longlina sp. nov., 拟穴居隆脊蚁甲Batrisocenus affinis sp. nov., 独石子隆脊蚁甲Batrisocenus dushizi sp. nov., 腹毛隆脊蚁甲Batrisocenus filum sp. nov., 莫氏隆脊蚁甲Batrisocenus mojiachengi sp. nov., 石林隆脊蚁甲Batrisocenus shilin sp. nov., 中庄隆脊蚁甲Batrisocenus trungtrang sp. nov., 圆茎脊胸蚁甲Tribasodites circinatus sp. nov., 地苏脊胸蚁甲T. disuensis sp. nov., 都安脊胸蚁甲T. duanus sp. nov., 桂林脊胸蚁甲T. guilinensis sp. nov., 胡氏脊胸蚁甲T. huchanghaoi sp. nov., 苗厅脊胸蚁甲T. miaoting sp. nov., 巴冲脊胸蚁甲T. pakchong sp. nov., 水族脊胸蚁甲T. shui sp. nov., 刺钩脊胸蚁甲T. uncinus sp. nov., 周氏脊胸蚁甲T. zhouhanpingi sp. nov., 洼腹幽蚁甲Zopherobatrus excavatus sp. nov., 开阳幽蚁甲Z. kaiyangus sp. nov., 思南幽蚁甲Z. sinanus sp. nov., 武隆幽蚁甲Z. wulongensis sp. nov., 索氏珠蚁甲Batraxis soparki sp. nov., 豪猪奇首蚁甲Nipponobythus haozhu sp. nov., 金鸡奇首蚁甲N. jinji sp. nov., 李氏奇首蚁甲N. liyunchuni sp. nov., 龙洞奇首蚁甲N. long sp. nov., 迷你奇首蚁甲N. minor sp. nov.（土栖）、牛鼻奇首蚁甲N. niubi sp. nov., 宽头奇首蚁甲N. platycephalus sp. nov., 隆额奇首蚁甲N. proiectus sp. nov., 粗点奇首蚁甲N. punctatus sp. nov., 黔奇首蚁甲N. qian sp. nov., 三仙奇首蚁甲N. sanxian sp. nov., 苏仙奇首蚁甲N. suxian sp. nov., 五峰奇首蚁甲N. wufengensis sp. nov.以及曾氏奇首蚁甲N. zengtingkaii sp. nov.。本研究将亚洲已知穴居蚁甲亚科物种数从48种增至90种, 近乎翻倍, 揭示了以往研究中存在的显著地域偏见, 并凸显了地下生境蕴含的丰富而隐秘的生物多样性。本研究详细编制了超族、族、属及种检索表并整理了亚洲穴居蚁甲亚科物种名录；此外, 还报道了雄性触角显著多态的新例, 揭示了性冲突的潜在形态学关联（疑似雌性抗拒性状）, 为洞悉无光环境下的进化动态提供了新视角。本研究奠定了亚洲穴居蚁甲分类的基础框架, 并强调持续开展洞穴生物调查的迫切需要, 以系统编目并有效保护亚洲喀斯特生态系统独特的生物多样性。