Rory A. Dow

Chlorocyphidae (Odonata: Zygoptera) represent a family of odonates found in tropical Asia and Africa, composed of 162 species. The family is characterized by a pronounced snout-like clypeus, wings that usually extend beyond the abdomen, and striking male coloration correlated with elaborate courtship behavior. Morphological differences between species and among genera are limited. To date, there is no extensive phylogeny of the group. As a result, the current classification is primarily based on coloration and morphological features that have not been tested in a phylogenetic framework. Our taxon sampling includes 90 species (> 55% of diversity). We targeted 92 AHE genes to test previous hypotheses of chlorocyphid classification. We present an updated classification. We recover Chlorocyphidae as monophyletic and Cyrano as sister to the remaining genera. We describe two new subfamilies, increasing the number to six: Cyranoinae Fomekong-Lontchi, Kalkman and Bybee subfam. nov . and Indocyphinae Fomekong-Lontchi, Kalkman and Bybee subfam. nov . A total of 26 species are reclassified to another genus (highlighting the difficulty with Chlorocyphidae morphology and classification), of which 13 are placed in two newly described genera: Ebenocypha Fomekong-Lontchi, Kalkman and Bybee gen. nov . and Scapanocypha Fomekong-Lontchi, Kalkman and Bybee gen. nov . Also, the genera Melanocypha , Pachycypha , Sclerocypha , and Watuwila are synonymized with Libellago . This new classification provides a framework for future studies of chlorocyphid evolution, including their courtship and biogeography.

We present an updated classification for extant dragonflies and damselflies (Odonata) and summarize new insights gained over the past two decades. Our focus is on taxa of family-level and higher and we indicate subfamilies only when their monophyly is currently undisputed and well-supported by phylogenetic analyses. The superfamily Calopterygoidea was known to be polyphyletic and based on recent phylogenomic data is divided into nine superfamilies, of which eight are recognized for the first time (Amphipterygoidea stat. nov., Euphaeoidea stat. nov., Megapodagrionoidea stat. nov., Mesopod­agrionoidea stat. nov., Philogangoidea stat. nov., Polythoroidea stat. nov., Priscagrionoidea stat. nov., Tatocnemidoidea stat. nov.). At present, midway through 2025, odonates are divided into three suborders, 17 superfamilies, 55 families and 687 genera containing 6447 species (May 1, 2025). We give an overview of the distribution of the families across major biogeographical realms. Except for Amanipodagrionidae, nymphs of at least some of the species of all families are known. Our understanding of the classification of dragonflies and damselflies has greatly improved in the past two decades largely due to phylogenetic inferences based on molecular studies. We expect that in the next few years the last remaining issues regarding the higher-level phylogeny and classification, including the position of the South American genus Sciotropis and the division into subfamilies of families such as Coenagrionidae, Aeshnidae, Gomphidae and Libellulidae, will be better refined by the acquisition of additional morphological and genomic data.

Sumatra, the second largest island in Sundaland, has been poorly studied for Odonata. Sumatra has ca. 72,300 km2 of peatlands which, prior to human alteration, were almost entirely covered by Peat Swamp Forest (PSF); Riau Province has the largest peatlands on the island. Less than 20% of Sumatra’s peatlands remain as PSF (pristine or degraded). PSF is known to provide habitat for specialised and/or threatened plant and animal species, including Odonata, but has received little sampling effort. No study on Odonata of any PSF area in Sumatra has been reported until now. The Kampar Peninsula in Riau Province has the largest remaining PSF (3,445.73 km2) in Sumatra, with 1,300.95 km2 specifically managed by the Restorasi Ekosistem Riau (RER) programme with the aim of protecting and restoring the degraded PSF at the core of the peninsula. In early 2020 the first author, together with Universitas Muhammadiyah Riau, commenced surveys of Odonata in the Kampar Peninsula, funded and facilitated by RER. After a long pause due to the global Covid-19 pandemic, surveys resumed in 2023. All surveys were conducted under research permits issued by the Indonesian National Research and Innovation Agency. The results of the Odonata surveys in the Kampar Peninsula are presented here. Surveys were conducted in January 2020, February 2023, May/June 2023 and September/October 2023, the spacing of and weather conditions during the surveys are discussed. Ninety-seven individual sites where specimens were collected are listed, grouped into 19 general areas. One hundred species and subspecies recorded during the surveys are listed with notes on habitat and in some cases other remarks. Taxonomic notes and remarks are given for some of the taxa recorded (Podolestes buwaldai Lieftinck, 1940, Copera vittata (Selys, 1863), Mortonagrion sp. cf. aborense (Laidlaw, 1914), M. megabinluyog Dow & Choong, 2015, M. sp. cf. forficulatum Lieftinck, 1953, Teinobasis ruficollis (Selys, 1877), T. sp. cf. suavis Lieftinck, 1953, Gynacantha dohrni Krüger, 1899 and G. maclachlani Krüger, 1899 (under the first name), Heliaeschna crassa Krüger, 1899 and H. idae (Brauer, 1865) (under the first name). In particular three forms of penile organ found in the M. aborense complex in Sumatra are discussed and illustrated with sketches; the structure of the female prothorax in T. ruficollis is discussed; the taxonomic history, status of type material, and diagnostic characters of G. dohrni and G. maclachlani are discussed in detail and illustrated with stacking images of the male anal appendages of type material of both species. Ninety-four of the taxa recorded are identified to a named species or subspecies, five are only identified with certainty to genus level, pending further taxonomic work and collection of more samples, and one taxon is identified as an unnamed but already known species. Of the distinct taxa recorded, 42 are from the Zygoptera and 58 from the Anisoptera. Most sampled habitats can be classified as either lowland swamp forest or open and disturbed; typically for both classes of habitat, the majority of species recorded are from the Coenagrionidae and Libellulidae. Eighty-nine of the 94 taxa identified to a named species/subspecies have also been recorded in Peninsular Malaysia and/or Singapore. At least 38 of the taxa recorded represent first published records for either Indonesia (five species: Amphicnemis bebar Dow, Choong & Ng, 2010, Amphicnemis hoisen Dow, Choong & Ng, 2010, Amphicnemis reri Dow, Gesriantuti & Lupiyaningdyah, 2025, M. megabinluyog and Chalybeothemis chini Dow, Choong & Orr, 2007), Sumatra including satellite islands (two taxa), mainland Sumatra (six taxa) or Riau Province (25 taxa); these first records are discussed. With these results 271 species of Odonata are known from Sumatra and 126 from Riau Province. Of the 100 distinct taxa recorded in the surveys in the Kampar Peninsula we regard 49 as forest-dependent (requiring the presence of forest for their survival) and 14 as specialists of low pH waters. A discussion of the species found in natural forest, planted forest and open habitats is included. Fifteen of the species recorded currently have IUCN Red List assessments other than Least Concern (four Data Deficient, one Endangered, three Vulnerable and seven Near Threatened. One species, A. reri, has not yet been assessed but would likely fall in one of the threatened categories with the data available. Fifteen percent is a high proportion of non-Least Concern species for a sample of Odonata in a relatively small area but fairly typical for low pH water habitats in Sundaland; all but one of the species regarded as low pH specialists have non-Least Concern Red List assessments. A comparison of the odonate fauna of the Kampar Peninsula with that of PSF areas in Borneo and Peninsular Malaysia is made and odonate diversity at low pH waters in Sundaland is discussed.

Amphicnemis reri sp. nov. is described from tropical Peat Swamp Forest (PSF) in the Kampar Peninsula, Riau Province, Sumatra, Indonesia (holotype ♂, pools in PSF, Meranti Estate Block B Transect, Kampar Peninsula, Pelalawan, Riau, Sumatra, Indonesia, 0.3363N, 102.6440E, 7 ii 2023, deposited with the Indonesian National Research and Innovation Agency). The female (supposition) is also described. Amphicnemis bebar Dow, Choong & Ng, 2010 and A. hoisen Dow, Choong & Ng, 2010, both originally described from Peninsular Malaysia, are recorded from Sumatra for the first time; variation in these species is discussed and, in the case of A. bebar is illustrated. Species groups within Amphicnemis Selys, 1863 are defined and briefly discussed.

The final stadium larvae of the following four species of south-east Asian Chlorocyphidae are described and compared: Aristocypha fenestrella (Rambur), Heliocypha biseriata (Selys), Libellago hyalina (Selys) and Sundacypha petiolata (Selys), including both sexes for the latter two species. Excepting one L. hyalina specimen from Brunei, identified by supposition based on habitat, all specimens were identified by comparing and matching the mitochondrial marker COI with that of known adult specimens from Sarawak, Brunei and several localities throughout tropical Asia. The specimens presented close matches with all adults in this gene. An assessment of the efficacy of this method of identification is provided, noting that in some cases close species cannot be separated by bar-code matching and ultimate determination is partially based on known distributions of adults. Some aspects of the relationships among genera revealed by the genetic analyses are also discussed. In addition, an exuvia of Libellago lineata (Burmeister) from northern Thailand, identified by supposition, is partially described for the purpose of comparison with L. hyalina. For the morphological analysis the unique features of chlorocyphid anatomy are discussed, and some new terminology is introduced. Overall, the morphological analysis revealed numerous clear differences between the four species studied, and comparisons with available literature suggest that some of these may be characteristic of their genera. It is also evident that in some cases clear interspecific differences occur within genera. It is however concluded that a generic level larval key for the Oriental region Chlorocyphidae based on morphology may never be attainable, although local generic or even species level keys addressing the fauna of limited geographic areas may be possible in many places, especially as the larvae of more species come to be known and described in detail.

A checklist, based on a database containing published data, of the Odonata (dragonflies and damselflies) occurring in Sundaland and Wallacea is presented. The presence of (sub)species is indicated for eight main regions (Singapore & Peninsular Malaysia, South China Sea (islands in the South China Sea that are not sensibly treated as satellites of larger landmasses), Borneo, Sumatra, Java & Bali, Lesser Sunda, Sulawesi, Moluccas), 22 subregions and 80 smaller islands and island groups. In total 743 full species are recorded from the entire area with 549 species known from Sundaland and 270 from Wallacea. Of these 482 are not found outside Sundaland and Wallacea, 385 (ca. 52% of the fauna) of which are single region endemics; the majority of these are actually single island endemics. Notes are provided on taxonomic problems or indicating problematic distribution records. Prodasineura lansbergei is considered to be a nomen nudum (stat nov.). For each of the eight main regions the history of the study of odonates is briefly discussed, information is provided on the coverage of the available data and the faunal composition is described. An overview is given of genera for which no larvae have been described. A brief comparison is made between the faunas of Sundaland and Wallacea showing that they only share 10% of the species between them (76 of 743).

A modified and expanded definition of the Drepanosticta carmichaeli-group is given. This includes the species: D. annandalei Fraser, 1924, D. brownelli Tinkham, 1938, D. carmichaeli (Laidlaw, 1915), D. emtrai Dow, Kompier & Phan, 2018, D. hongkongensis Wilson, 1997, D. jurzitzai Hämäläinen, 1999, D. sumatrana Sasamoto & Karube, 2007, D. tenella Lieftinck, 1935, D. vietnamica Asahina, 1997 and D. wildermuthi sp. nov.). The species of the group are reviewed and in most cases illustrated, diagnostic notes are given wherever possible. Drepanosticta polychromatica Fraser, 1931 is considered to be a junior synonym of D. carmichaeli and variation in D. carmichaeli is discussed. Drepanosticta wildermuthi sp. nov. from the Central Highlands of Vietnam (holotype male from Bao Loc district, Lam Dong Province) is described. The female of D. jurzitzai Hämäläinen, 1999 is described for the first time. A key to the males of the Drepanosticta carmichaeli-group is provided.

Coeliccia furcata Hämäläinen, 1986, an atypical species of Coeliccia Kirby, 1890 with bifurcated male cerci, had been known solely from the holotype male collected in Myanmar in 1938. In recent years the species has been rediscovered in Yunnan, China. The first description of the female is given, along with descriptive notes on the male and illustrations of both sexes, including fresh images of the holotype. Another of the unusual features of C. furcata is the extremely simple form of its penile organ, this is discussed and compared with other species from mainland Asia and Japan currently placed in Coeliccia and with a simple penile organ.

Revision of the genus Coeliccia Kirby in Borneo part III: Two new species from the C. membranipes –group from Sarawak and Brunei (Odonata: Zygoptera: Platycnemididae)