Adrian R. Plant

Aquatic insects in the Dinarides: identifying hotspots of endemism and species richness shaped by geological and hydrological history using Empididae (Diptera)

We identify local hotspots of elevated species richness and endemicity of aquatic insects in the Dinarides (Balkans) using aquatic Empididae (Diptera) as models and compared observed patterns with historical geological and hydrological changes in the region. We analysed species richness and reciprocal weighted endemicity. Parsimony analysis of endemism was used to generate an historical hypothesis of the relationships between the aquatic Empididae biotas of different regions of local endemism within the Dinarides and the results reconciled with the geological and hydrological history of the region. Hotspots of high endemicity were coincident with areas of elevated species richness and were located primarily in the northwest Dinarides, central Bosnia & Herzegovina, and southeast Montenegro. Parsimony analysis of endemism retrieved three weakly supported area clades comprising areas of southeast Dinarides together with disjunct areas at the western Dinarides, and central Bosnia & Herzegovina together with the disjunct northwest Dinarides. The aquatic Empididae fauna of the northwest Dinarides has been strongly influenced by dispersal from the European Alps, whereas that at the southern end of the range may have been shaped by immigration from ranges further south. In the central Dinarides, progressive fragmentation of the hydrological network through orogenesis, karstification, redistribution of flysch deposits, and saline ingressions may underlie current diversity patterns with populations becoming increasingly isolated and focused within refugia. Hotspots of endemism and diversity are not necessarily coincident with protected areas and further work will be needed if they are to be reconciled with future planning of conservation priorities.

Extreme male leg polymorphic asymmetry in a new empidine dance fly (Diptera: Empididae).

A new dance fly (Empididae: Empidinae) with hugely modified male fore tarsus, either on the right, left, both or neither sides, is described from Japan. Such massive polymorphic asymmetry occurring with so high an incidence in a population is previously unreported. In view of the courtship behaviour of other Empidinae, we hypothesize that the oversized tarsus is a secondary sexual character employed by males for attracting females. Alternative hypotheses are also discussed. We suggest that this extraordinary new species is a potential model for the study of mating biology in Empidinae and the evolution of mating systems in general.

Hemerodromiinae (Diptera: Empididae): a tentative phylogeny and biogeographical discussion

A cladistic analysis of adult morphological characters was used to hypothesize phylogenetic and zoogeographical relationships in Hemerodromiinae (Diptera: Empididae). Afrodromia Smith, Drymodromia Becker and an undescribed Chilean genus subtended a sister‐group relationship between the tribes Chelipodini and Hemerodromiini. Chelipodozus Smith and an undescribed Australian genus were supported only weakly in Hemerodromiini, and are regarded as incertae sedis within Hemerodromiinae. In Chelipodini, Anaclastoctedon Plant and an undescribed Australian genus subtended all others that form two sister‐group clades. (i) A widespread Chelipoda‐like group comprising Achelipoda Yang, Zhang & Zhang, Ptilophyllodromia Bezzi, Chelipoda Macquarts.s. and Phyllodromia Zetterstedt s.s. has a worldwide distribution, excepting the Afrotropical, considered to date from before or during the early phase of Gondwanan fragmentation. Phyllodromia s.s. is an exclusively Palaearctic genus of doubtful validity, to which Southern Hemisphere forms have been assigned incorrectly. Ptilophyllodromia Bezzi syn.n. is regarded as a junior synonym of Chelipoda, with the included species relegated to a species group. (ii) An austral Chelipoda‐like group confined to New Zealand, New Caledonia, Lord Howe Island and Vanuatu, includes Monodromia Collin and species currently incorrectly assigned to Chelipoda and Phyllodromia. The group is hypothesized as a relictual Gondwanan element that has survived Oligocene drowning as metapopulations persisting in situ on ephemeral islands along arcs, ridges and buoyant crustal blocks overlying hot spots in New Zealand and the Southwest Pacific. In the tribe Hemerodromiini, many Southern Hemisphere species assigned currently to Hemerodromia Meigen, Cladodromia Bezzi and Neoplasta Coquillett require reassignment. The sister‐group relationship between the southern African endemic Afrodromia and other Hemerodromiinae is viewed as evidence of early divergence of Hemerodromiinae and Empidinae by the early Cretaceous, pre‐dating major Gondwanan fragmentation. An assessment of fossil forms indicated that Chelifera detestata (Meunier) from Eocene/Oligocene Baltic amber is the only genuine described fossil representative of Hemerodromiinae.

Phylogenetic systematics of the Gondwanan Empis macrorrhyncha group (Diptera, Empididae, Empidinae)

The Empis macrorrhyncha group (Diptera: Empididae) from cool to warm temperate areas of South America and Australia is diagnosed and cladistically analysed, and five new species, Empis animosasp.n., E. austerasp.n., E. maculosasp.n., E. occidentalissp.n. and E. pedivillosulasp.n., are described. Cladistic analysis of 23 adult morphological characters for 14 species of the group generated a single tree of 28 steps (CI = 0.82; RI = 0.93). Monophyly was established on the basis of a single apomorphy, possession of a bilobed cercus of the male hypopygium. Three main clades were inferred: clade 1 included three Patagonian and a single southwestern Australian species; clade 2 included two species from southeastern Australia; clade 3 included a large Patagonian group of five species and a single southeastern Australian species. The E. fulvicollis complex (clade 1) is a sister‐group of the E. macrorrhyncha complex (clades 2 + 3). A provisional historical biogeographic hypothesis is advanced correlating the appearance of the South American and Australian sister lineages with the timing of the break‐up of Gondwana.

The genus Hybos Meigen (Diptera: Empidoidea: Hybotidae) in Thailand.

The genus Hybos Meigen in Thailand is revised and full descriptions and keys provided for all 41 species. Thirty-four new species are recognized: H. aceriformis sp. nov., H. ancyclochiles sp. nov., H. anisoserratus sp. nov., H. bispinatus sp. nov., H. chaweewani sp. nov., H. daugeroni sp. nov., H. divisus sp. nov., H. grootaerti sp. nov., H. hylobates sp. nov., H. inthanonensis sp. nov., H. kaluang sp. nov., H. khamfui sp. nov., H. konkaogwang sp. nov., H. lannaensis sp. nov., H. mangraii sp. nov., H. meeamnat sp. nov., H. men sp. nov., H. merzi sp. nov., H. ngachang sp. nov., H. paknok sp. nov., H. phahompokensis sp. nov., H. pisadaanus sp. nov., H. saenmueangmai sp. nov., H. shamshevi sp. nov., H. sinclairi sp. nov., H. songbai sp. nov., H. steatopygus sp. nov., H. stigmaticus sp. nov., H. subapicalis sp. nov., H. tetricus sp. nov., H. thaosaeo sp. nov., H. thepkaisoni sp. nov., H. tilokarati sp. nov., H. yungyak sp. nov. Seven species known previously from China are recognized: H. ancistroides Yang & Yang, H. apicihamatus Yang & Yang, H. longus Yang & Yang, H. particularis Yang, Yang & Hu, H. serratus Yang & Yang, H. xishuangbannaensis Yang & Yang, and H. zhejiangensis Yang & Yang. Eight informal species-groups are tentatively proposed based on characters of male and female terminalia and attention is drawn to the many previously overlooked taxonomically useful characters of the female terminalia. Distribution maps of all species are presented and distributions categorised as ‘widespread’, ‘eastern’, ‘southern’, north-eastern’ or ‘northern and western’. Species richness and endemicity are greatest in mid to high elevation evergreen forest biotopes of the northern mountains and areas of endemism are identified on the Luang Prabang, Daen Lao, Thanon Thong-chai ranges and on the Isaan Plateau at least. Adult phenology is correlated with the rainy season in many species and preliminary analyses reveal that many high-elevation species have short emergence periods and restricted distributions, whereas some lowland species have longer emergence periods and wider distributions.

The genus Icasma Collin (Diptera: Empididae: Ceratomerinae), and the description of a new species.

A new species, Icasma aequbilis, is described from specimens collected in the Waikato region, North Island, New Zealand. This species is compared with Icasma singularis Collin, a widespread species in the North Island.

The female of Austropeza Plant, 1989 (Diptera: Empidoidea: Hybotidae; Ocydromiinae)

The female of the genus Austropeza Plant, 1989 (Diptera, Hybotidae) is described and illustrated for the first time from a specimen of Austropeza insolita (Collin, 1928) captured in New Zealands South Island.

The Hemerodromiinae (Diptera: Empididae) of New Zealand III. Antipodromia new genus

Antipodromia new genus (Diptera: Empididae: Hemerodromiinae) is described. The genus is monotypic with Hemerodromia radialis Collin, 1928 its type species and is endemic to New Zealand. Systematic relationships with Hemerodromia Meigen and other Hemerodromiinae are discussed.

Phylogenetic reappraisal and taxonomic review of the Empis (Coptophlebia) hyalipennis-group (Diptera : Empididae : Empidinae)

The Empis (Coptophlebia) hyalipennis-group is redefined, keyed, cladistically analysed and three new species, Empis doi, sp. nov., Empis pseudohystrichopyga, sp. nov. and Empis pseudovillosipes, sp. nov., are described (from Thailand, China and Taiwan respectively). A cladistic analysis of 27 adult morphological characters was performed for Empis hyalipennis (the type-species of the subgenus Coptophlebia), and 13 Nearctic, Oriental and Palaearctic species hypothesised as being closely related, which resulted in a single tree of 31 steps (CI = 0.96; RI = 0.98). Monophyly of the group was established on the basis of two apomorphic characters of the male hypopygium. Based on these results and previous studies of the Empis-Coptophlebia complex of subgenera within the genus Empis, the E. (C.) hyalipennis-group appears to be much richer in species, exhibit greater morphological heterogeneity and be more widely distributed than expected. It is strongly recommended that any future revision of the subfamily Empidinae should apply the name Coptophlebia only to the monophyletic Empis (Coptophlebia) hyalipennis-group defined in this study.

Neotropical Hemerodromia Meigen (Diptera: Empididae), a world of discovery II: New species from Atlantic forest, Brazil.

Eight new species of Hemerodromia Meigen are described from the Brazilian Atlantic forest, H. brevicercata sp. nov., H. carioca sp. nov., H. cornuhypandrialis sp. nov., H. megalamellata sp. nov., H. membranosa sp. nov., H. mourai sp. nov., H. cummingi sp. nov., and H. ubajaraensis sp. nov. are described and illustrated. This is the first record of the genus from the Brazilian southeast and northeast.