Rory A. Dow

Evolution of endemism on a young tropical mountain

Tropical mountains are hot spots of biodiversity and endemism, but the evolutionary origins of their unique biotas are poorly understood. In varying degrees, local and regional extinction, long-distance colonization, and local recruitment may all contribute to the exceptional character of these communities. Also, it is debated whether mountain endemics mostly originate from local lowland taxa, or from lineages that reach the mountain by long-range dispersal from cool localities elsewhere. Here we investigate the evolutionary routes to endemism by sampling an entire tropical mountain biota on the 4,095-metre-high Mount Kinabalu in Sabah, East Malaysia. We discover that most of its unique biodiversity is younger than the mountain itself (6 million years), and comprises a mix of immigrant pre-adapted lineages and descendants from local lowland ancestors, although substantial shifts from lower to higher vegetation zones in this latter group were rare. These insights could improve forecasts of the likelihood of extinction and ‘evolutionary rescue’ in montane biodiversity hot spots under climate change scenarios.

Redefining the damselfly families: a comprehensive molecular phylogeny of Zygoptera (Odonata)

An extensive molecular phylogenetic reconstruction of the suborder Zygoptera of the Odonata is presented, based on mitochondrial (16S, COI) and nuclear (28S) data of 59% of the 310 genera recognized and all (suspected) families except the monotypic Hemiphlebiidae. A partial reclassification is proposed, incorporating morphological characters. Many traditional families are recovered as monophyletic, but reorganization of the superfamily Coenagrionoidea into three families is proposed: Isostictidae, Platycnemididae and Coenagrionidae. Archboldargia Lieftinck, Hylaeargia Lieftinck, Palaiargia Förster, Papuargia Lieftinck and Onychargia Selys are transferred from Coenagrionidae to Platycnemididae, and Leptocnemis Selys, Oreocnemis Pinhey and Thaumatagrion Lieftinck from Platycnemididae to Coenagrionidae. Each geographically well‐defined clade of Platycnemididae is recognized as a subfamily, and thus Disparoneurinae (i.e. Old World ‘Protoneuridae’) is incorporated, Calicnemiinae is restricted, and Allocnemidinae (type genus: Allocnemis Selys) subfam.n., Idiocnemidinae (type genus: Idiocnemis Selys) subfam.n. and Onychargiinae (type genus: Onychargia Selys) subfam.n. and Coperini trib.n. (type genus: Copera Kirby) are described. Half of Coenagrionidae belongs to a well‐supported clade incorporating Coenagrion Kirby and the potential subfamilies Agriocnemidinae, Ischnurinae and Pseudagrioninae. The remainder is less well defined, but includes the Pseudostigmatidae and New World Protoneuridae that, with Argiinae and Teinobasinae, may prove valid subfamilies with further evidence. Ninety‐two per cent of the genera formerly included in the polyphyletic Amphipterygidae and Megapodagrionidae were studied. Pentaphlebiidae, Rimanellidae and Devadattidae fam.n. (type genus: Devadatta Kirby) are separated from Amphipterygidae, and Argiolestidae, Heteragrionidae, Hypolestidae, Philogeniidae, Philosinidae and Thaumatoneuridae from Megapodagrionidae. Eight further groups formerly placed in the latter are identified, but are retained as incertae sedis; the validity of Lestoideidae, Philogangidae and Pseudolestidae is confirmed. For some families (e.g. Calopterygidae, Chlorocyphidae) a further subdivision is possible; Protostictinae subfam.n. (type genus: Protosticta Selys) is introduced in Platystictidae. Numerous new combinations are proposed in the Supporting Information. Many long‐established families lack strong morphological apomorphies. In particular, venation is incongruent with molecular results, stressing the need to review fossil Odonata taxonomy: once defined by the reduction of the anal vein, Protoneuridae dissolves completely into six clades from five families.

A review of the Teinobasis of Sundaland, with the description of Teinobasis cryptica sp. nov. from Malaysia (Odonata: Coenagrionidae)

Abstract Teinobasis cryptica sp. nov. (holotype ♂: Borneo, Sarawak, Bahagian Samarahan, Kota Samarahan, old UNIMAS campus, disturbed peatswamp forest, 25 ii 2008, RMNH) from Malaysia is described from both sexes and compared with other Teinobasis species known to occur in Malaysia. The members of the genus known from Sundaland are reviewed. Records of T. ruficollis from Borneo are clarified. New records of other Bornean species are listed. The females of T. laidlawi, T. rajah and T. ruficollis are described for the first time. Keys are given to both sexes of all named species from the genus known from peninsular Malaysia, Singapore and the Greater Sunda Islands and the species are placed provisionally into two groups: the laidlawi-group and ruficollis-group. The former group also includes T. rubricauda from the Palawan region of the Philipplnes, which may be a junior synonym of T. laidlawi.

Revision of the Sundaland species of the genus Dysphaea Selys, 1853 using molecular and morphological methods, with notes on allied species (Odonata: Euphaeidae)

The Sundaland species of the genus Dysphaea were studied using molecular and morphological methods. Four species are recognized: D. dimidiata Selys, D. lugens Selys, D. ulu spec. nov. (holotype ♂, from Borneo, Sarawak, Miri division, Upper Baram, Sungai Pejelai, Ulu Moh, 24 viii 2014; deposited in RMNH) and D. vanida spec. nov. (holotype ♂, from Thailand, Ranong province, Khlong Nakha, Khlong Bang Man, 12-13 v 1999; deposited in RMNH). The four species are described and illustrated for both sexes, with keys provided. The type specimens of the four Dysphaea taxa named by E. de Selys Longchamps, i.e. dimidiata, limbata, semilimbata and lugens, were studied and their taxonomic status is discussed. Lectotypes are designated for D. dimidiata and D. limbata. D. dimidiata is recorded from Palawan (the Philippines) for the first time. A molecular analysis using three markers (COI, 16S and 28S) is presented. This includes specimens of three Sundaland species of the genus (D. lugens missing) and two congeners from other regions (D. basi-tincta and D. gloriosa). Notes and photographs of the male holotype of D. walli Fraser (from Maymyo, Burma) are provided.

Taxonomy and molecular phylogeny of the Platystictidae of Sri Lanka ( Insecta: Odonata)

The 22 Sri Lankan representatives of the family Platystictidae, all endemic to the island and belonging to the distinct endemic subfamily Platystictinae, are revised, and a new reconstruction of the phylogeny based on molecular characters is provided. Five new species are described: Ceylonosticta venusta sp. nov. (holotype ♂: Rambodde Falls, at the tunnel; Nuwara Eliya District, Central Province; N7.0489, E80.6961; 12-vii-2012; to be deposited at National Museum of Natural History, Colombo, Sri Lanka), C. inferioreducta sp. nov. (holotype ♂: Norton Bridge, stream on the B43 road 1.5km WNW of Norton Bridge; Nuwara Eliya District, Central Province; N6.9171, E80.5075; 28-vii-2009; to be deposited at National Museum of Natural History, Colombo, Sri Lanka), C. mirifica sp. nov. (holotype ♂: Uwella, primary forest on the road Uwella-Ratnapura, 11.5km NW of Balangoda; Ratnapura District, Sabaragamuwa Province; N6.6968, E80.6059; 16-vii-2012; to be deposited at National Museum of Natural History, Colombo, Sri Lanka), Platysticta secreta sp. nov. (holotype ♂: Hasalaka; Kandy District, Central Province; N7.3535, E80.9509; 31-v-1975; deposited at National Museum of Natural History, Smithsonian Institution, Washington, USA) and P. serendibica sp. nov. (holotype ♂: Kanneliya; Galle District, Southern Province; N6.2291, E80.3834; 8 & 9-vi-1975; deposited at National Museum of Natural History, Smithsonian Institution, Washington, USA). Additionally, a determination key, figures showing morphological details and coloration in life, as well as distribution maps for all species are presented. Based on molecular analysis of 21 taxa, the phylogeny of Platystictinae is presented and discussed from the zoogeographical and paleogeographical point of view. Sri Lankan species, traditionally placed in the genera Platysticta Selys and Drepanosticta Laidlaw / Ceylonosticta Fraser, separated into distinct clades within the subfamily as presently defined, but the monophyletic nature of the Platystictinae and its Sri Lankan endemicity is confirmed. For the South Indian species, formerly known as Platysticta deccanensis, morphological and molecular analyses demonstrated that it does not belong to the Sri Lankan clade and a new genus Indosticta gen. nov. is erected to accommodate it.

Revision of the genus Devadatta Kirby, 1890 in Borneo based on molecular and morphological methods, with descriptions of four new species (Odonata: Zygoptera: Devadattidae)

Species of Devadatta from Borneo are studied using both morphological and molecular methods. As well as D. podolestoides Laidlaw, four new species are recognised from the island: D. aran spec. nov. (holotype ♂, from Pulong Tau National Park, Miri division, Sarawak, Malaysia, deposited in RMNH), D. clavicauda spec. nov. (holotype ♂, from Bukit Mina, Bukit Mina Wildlife Corridor, Sarawak Planted Forest Project, Bintulu division, Sarawak, Malaysia, deposited in RMNH), D. somoh spec. nov. (holotype ♂, from the Sungai Kahei area, Ulu Balui, Kapit division, Sarawak, Malaysia, deposited in RMNH) and D. tanduk spec. nov. (holotype ♂, from Poring Hot Springs, Kinabalu National Park, West Coast division, Sabah, Malaysia, deposited in RMNH). The Philippine taxon D. basilanensis Laidlaw is considered a good species rather than a subspecies of D. podolestoides. The Bornean species plus D. basilanensis are provisionally considered to form a species group, the podolestoides-group, within Devadatta. The species of the podolestoides-group are so similar in morphology and colouration that they are close to truly cryptic species. Two species appear to exhibit character displacement where their ranges overlap with other Devadatta species. A molecular analysis using four markers (COI, 16S, ITS and 28S) is presented. This analysis includes specimens of all species from the podolestoides-group and two Devadatta species from mainland Asia.

Underwater oviposition behavior in two species of Euphaea in Borneo and Hong Kong (Odonata: Euphaeidae)

Abstract Submerged oviposition behaviour by female Euphaea decorata and E. subcostalis is reported. E. decorata descended to within 10 cm of the stream bottom and oviposited endophytically for a total of 59 min. An E. subcostalis female descended 3 cm to dead leaves at the stream bottom and stayed submerged for 20-25 s. Non-contact guarding by the male was observed in both cases. Submerged oviposition into substrates near to the stream bottom may be common behaviour for members of the Euphaeidae.

Out of Australia: the Argiolestidae reveal the Melanesian Arc System and East Papua Composite Terrane as possible ancient dispersal routes to the Indo-Australian Archipelago (Odonata: Argiolestidae)

Information on the origin of distribution patterns shown by freshwater invertebrates in the Indo-Australian Archipelago is poor. Here we present a molecular based hypothesis of the phylogenetic relationships of Argiolestidae, a family of damselflies found throughout the tropical parts of the Eastern Hemisphere. We use this to address the following questions: (1) did Argiolestidae colonize Wallacea and the Philippines from the Eurasian or from the Australian continent? (2) Is the presence of Argiolestidae in New Guinea the result of a single colonization event, i.e. are the Argiolestidae found in New Guinea monophyletic? The results show that clades occurring in the Philippines, Wallacea and New Guinea all originate from Australian ancestors. Representatives in Sundaland are most closely related to African genera and failed to reach the Philippines and Wallacea. The presence of Argiolestidae north of Australia is the result of at least three colonization events from Australia to areas that presently compose New Guinea and probably a fourth from Australia to Sulawesi. The two most diverse lineages found north of Australia show different distribution patterns. One reaching north as far as Luzon, presumably facilitated by Late Oligocene to Miocene islands arcs (Melanesian Arc System). The other clade shows a diversification of two genera and numerous species in the eastern tail of New Guinea, an area largely corresponding with the East Papuan Composite Terrane (EPCT) followed by the expansion of one genus into the rest of New Guinea. The EPCTs importance as source area for the New Guinean fauna has been suggested on the basis of distribution patterns, but we present the first evidence based on phylogeny reconstruction of strong diversification on this formerly isolated landmass.

Telosticta fugispinosa sp. nov. from Sabah (Odonata: Zygoptera: Platystictidae)

Telosticta fugispinosa sp. nov. (holotype male, from Borneo, Sabah, West Coast division, Crocker Range National Park, Inobong, Kimamabang waterfall stream system, 21 ix 2012, deposited in RMNH) is described from Kinabalu National Park and Crocker Range National Park in Sabah, Malaysian Borneo. It is distinguished from all other species of Telosticta by the form of the male anal appendages.

Revision of the genus Coeliccia Kirby in Borneo part II: Two new species from the membranipes -group, with a redescription of C. macrostigma Laidlaw (Odonata: Zygoptera: Platycnemididae)

Coeliccia matok sp. nov. (holotype male from Borneo, Sarawak, Samarahan Division, peat swamp forest at old UNIMAS campus, 25 ii 2008, to be deposited in BMNH) and Coeliccia paludensis sp. nov. (holotype male from Borneo, Kalimantan Tengah, peat swamp forest in ex Mega Rice Project Block E, 18 vi 2012, in RMNH) are described from Borneo. The two new species are apparently confined to peat swamp forest (C. paludensis) or largely confined to peat swamp forest and related forest formations (C. matok). Coeliccia macrostigma Laidlaw is redescribed and all available information on it is summarised. Additional terminology for characters of the prothorax in Coeliccia species is introduced. Distribution maps are given for all three species considered.