The Six Dwarfs of the Middle East: revision of the enigmatic praying mantis genus Holaptilon (Mantodea: Gonypetidae: Gonypetinae) with the description of four new species under integrative taxonomy

Abstract

The dwarf-mantid genus Holaptilon Beier, 1964 is composed of small-sized ground-runner species distributed in the Middle East. Due to their elusive lifestyle, little is known about their behaviour, distribution, and phylogeny. The genus Holaptilon was once established for a single species, H. pusillulum Beier, 1964, based on material collected in Jerusalem, Israel. Later, H. brevipugilis Kolnegari, 2018, and H. yagmur Yılmaz and Sevgili, 2023 were described from Iran and Turkey, respectively. In this study, integrated morphology, molecular analyses, and ecology were used to revise the genus Holaptilon and define the boundaries of its species. New data on this genus are presented, based on Holaptilon specimens collected from various provinces of Iran, Israel, Jordan, and Turkey. Extensive analyses, including examinations of male and female genitalia, morphometrical analysis, and morphological hypervolumes were conducted to distinguish its species morphologically. In addition, four molecular markers (mitochondrial and nuclear) were studied to gain a better understanding of species delimitation and phylogenetic relationships. As a result, impressive inter- and intraspecific variability was recovered. In addition to the three already known species, four new species with their distributions restricted to Iran (H. abdullahii sp. nov., H. khozestani sp. nov., H. iranicum sp. nov., and H. tadovaniensis sp. nov.) are here described, and H. yagmur Yılmaz and Sevgili, 2023 is synonymized with H. brevipugilis Kolnegari, 2018. The integrative approach was essential for an adequate classification in Holaptilon taxonomy and also helpful in the clarification of problematic and cryptic Mantodea species. Additional information concerning the life cycle, ecological aspects, spermatophore feeding, as well as geographic range and historical biogeography of Holaptilon species is also provided.