Empis (Coptophlebia) hyalea, Melander, 1946
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publication ID |
https://doi.org/10.1111/j.1096-3642.2005.00189.x |
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https://treatment.plazi.org/id/039D7838-FFDE-FFC8-3C6E-DB17FACDFD5E |
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treatment provided by |
Diego (2021-08-31 18:22:02, last updated by Plazi 2023-11-06 11:23:43) |
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scientific name |
Empis (Coptophlebia) hyalea |
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E. (C.) HYALEA View in CoL -GROUP AND THEIR DISTRIBUTION
As a result of the phylogenetic analysis, six main clades are recognized within the E. (C.) hyalea -group ( Fig. 1 View Figure 1 ), namely E. (C.) acris , E. (C.) aspina , E. (C.) atratata , E. (C.) hyalea , E. (C.) jacobsoni and E. (C.) nahaeoensis . Two main lineages can be distinguished: the first one is represented by the E. (C.) nahaeoensis and E. (C.) aspina clades, which are sister-groups on the basis of the reduction of the hypandrium; the second is represented by the E. (C.) acris , E. (C.) atratata , E. (C.) hyalea and E. (C.) jacobsoni clades on the basis of the presence of minute spines at the tip of the cercus.
The first lineage is presently known from the Oriental Region ( Figs 15 View Figure 15 , 40 View Figure 40 ), especially in Southern China, Laos and Thailand, with an extension of the E. (C.) aspina clade in Japan [ E. (C.) urumae is recorded from Ryukyu Islands]. Preliminary studies of material collected in Japan, Taiwan and South Korea show a large presence of this lineage in these countries with many new species to describe.
Within the second lineage, the E. (C.) atratata clade, distributed in South-east Asia ( Fig. 16 View Figure 16 ), is represented only by the two species included in the analysis as well as E. (C.) velutinella Frey. It is the most basal clade, sister-group of the E. (C.) acris + E. (C.) hyalea + E. (C.) jacobsoni clade.
The E. (C.) acris clade is the sister-group of the E. (C.) hyalea + E. (C.) jacobsoni clade on the basis of the reduction of the two long dorsal epandrial projections to two small protuberances. This clade is distributed only in New Guinea ( Fig. 11 View Figure 11 ), mainly in mountainous areas. Finally the E. (C.) hyalea and E. (C.) jacobsoni clades are sister-groups on the basis of the more or less deformed male fore tibia. They seem to have radiated together: they represent half of the species included in the analysis and twice the number of the species of their sister-group. The distribution of the E. (C.) jacobsoni clade is very wide, ranging from tropical areas of Africa to New Caledonia ( Fig. 35 View Figure 35 ). However, this clade has not been recorded in New Guinea. The distribution of the E. (C.) hyalea clade is confined to South-east Asia ( Fig. 23 View Figure 23 ).
THE E. (C.) HYALEA -GROUP: A POTENTIAL MODEL TO STUDY THE IMPACT OF QUATERNARY CLIMATE CHANGE ON ASIAN BIODIVERSITY
The E. (C.) hyalea -group shows a remarkable diversification in tropical and subtropical areas of the Oriental and Australasian regions. Species are found mainly in middle and high altitude primary or secondary rain forests, as well as in coastal areas where they occur in mangroves and swamp forests ( Daugeron & Grootaert, 2004a). The phylogeny obtained shows sister-group relationships between altitude and coastal species (or species groups), i.e. species (or species groups) occurring in disjunct areas, are displayed in a repeated way. For example, within the E. (C.) jacobsoni clade, E. (C.) mengluensis + E. (C.) mengyangensis (mountainous region, Yunnan Province, China) is the sister-group of E. (C.) jacobsoni + E. (C.) producta (mangroves, Java and Singapore, respectively); within the E. (C.) hyalea clade, E. (C.) thiasotes (mountainous region, Java) is the sister-species of E. (C.) vitisalutatoris (low-altitude rain forests, Singapore); within the E. (C.) nahaeoensis clade, E. (C.) nahaeoensis (middle-altitude mountainous region, Loei Province, Thailand) is the sisterspecies of E. (C.) kuaensis (mangroves, Songkhla Province, Thailand); within the E. (C.) atratata clade, E. (C.) velutinella (mountainous region, Kambaiti, Myanmar), not included in the phylogeny, seems to be the sister-group of E. (C.) atratata + E. (C.) kosametensis (both found in rain forests of Koh Samet Island, Thailand).
During the Quaternary, Asian primary rain forests were considerably fragmented, especially during ice ages, because of the generalized reduction in rainfall. In South-east Asia, forests persisted in still rainy, refuge areas such as mountain and coastal regions (e.g. see Brandon-Jones, 1998).
Even if the main six clades of the E. (C.) hyalea - group probably originated after major geological (e.g. tectonic) changes, the distribution of the present species, on the one hand, and the sister-groups relationships between coastal and altitude species (or species groups), on the other, suggest a potential connection between the present specific diversity of the E. (C.) hyalea -group and forest fragmentations during successive ice ages ( Daugeron & Grootaert, 2004b). The isolation of the ancestral populations in the refuge areas resulted in their genetic divergence and the development of new species. According to these preliminary results, the E. (C.) hyalea -group could be used as a model to analyse the impact of Quaternary climate changes on tropical biodiversity.
Brandon-Jones D. 1998. Pre-glacial Bornean primate impoverishment and Wallace's line. In: Hall R, Holloway JD, eds. Biogeography and geological evolution of SE Asia. Leiden: Backhuys, 393 - 403.
Daugeron C, Grootaert P. 2004 a. The Empis hyalea - group (Diptera: Empididae: Empidinae): A potential model for studying the impact of quaternary climate changes on the Asian biodiversity. In: Biology in Asia, International Conference, Singapore, 7 - 10 / 12 / 2004, abstracts book, 44 - 45.
Daugeron C, Grootaert P. 2004 b. The Empis hyalea - group (Diptera: Empididae: Empidinae) in Asia: specific richness, distribution and habitats. In: Biology in Asia, International Conference, Singapore, 7 - 10 / 12 / 2004, abstracts book, 109.
Figure 1. Cladogram (consensus tree) of the Empis (Coptophlebia) hyalea-group (outgroup omitted): length = 52 steps; CI = 0.76; RI = 0.92, showing the distribution of the characters and the main monophyletic lineages: A, E. (C.) nahaeoensis clade; B, E. (C.) aspina clade; C, E. (C.) atratata clade; D, E. (C.) acris clade; E, E. (C.) jacobsoni clade; F, E. (C.) hyalea clade.
Figure 11. Distribution of the Empis (Coptophlebia) acris clade: E. (C.) acris sp. nov. = 7; E. (C.) duplex sp. nov. = 3; E. (C.) lamellalta sp. nov. = 5; E. (C.) lata sp. nov. = 3, 4, 5; E. (C.) longiseta sp. nov. = 6; E. (C.) missai sp. nov. = 2; E. (C.) parvula sp. nov. = 1; E. (C.) submetallica sp. nov. = 8; E. (C.) woitapensis sp. nov. = 7.
Figure 15. Distribution of the Empis (Coptophlebia) aspina clade: E. (C.) aspina sp. nov. = 1; E. (C.) miranda Daugeron & Grootaert = 4; E. (C.) nganga Daugeron & Grootaert = 6; E. (C.) projecta sp. nov. = 2; E. (C.) pulchra Daugeron & Grootaert = 5; E. (C.) ratburiensis Daugeron & Grootaert = 3; E. (C.) urumae sp. nov. = 7.
Figure 16. Distribution of the Empis (Coptophlebia) atratata clade: E. (C.) atratata Daugeron & Grootaert = 2; E. (C.) kosametensis Daugeron & Grootaert = 2; E. (C.) velutinella Frey = 1.
Figure 23. Distribution of the Empis (Coptophlebia) hyalea clade: E. (C.) hyalea Melander = 2; E. (C.) inferiseta sp. nov. = 4, 5; E. (C.) lamruensis Daugeron & Grootaert = 6; E. (C.) leptargyra Frey = 1; E. (C.) ostentator Melander = 2; E. (C.) pakensis Daugeron & Grootaert = 8; E. (C.) pseudospinotibialis Daugeron & Grootaert = 7; E. (C.) sinensis Melander = 2; E. (C.) spinotibialis Daugeron & Grootaert = 3; E. (C.) thiasotes Melander = 10; E. (C.) vitisalutatoris sp. nov. = 9.
Figure 35. Distribution of the Empis (Coptophlebia) jacobsoni clade: E. (C.) jacobsoni De Meijere = 6 (Java); E. (C.) lachaisei sp. nov. = 1 (Ivory Coast); E. (C.) loici sp. nov. = 2 (Central African Republic); E. (C.) menglunensis sp. nov. = 3 (China); E. (C.) mengyangensis sp. nov. = 3 (China); E. (C.) cameronsensis sp. nov. = 4 (Malaysia); E. (C.) nimbaensis sp. nov. = 1 (Guinea); E. (C.) nondouensis Daugeron = 7 (New Caledonia); E. (C.) padangensis sp. nov. = 5 (Sumatra).
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