Keywords: Coranus; 16S rRNA; Cyt b; COI and 28S rRNA; Harpactorinae; Biocontrol Agents; Intrageneric Molecular Biosystematics; Speciation; Ecotypes; Geographical Isolation;
Hence, classifications of Reduviidae based on morphological characters may at times become insufficient, and there is an urgent need for a cohesive meaningful classification of Reduviidae based on ecological, morphological, behavioural, cytological, and biochemical data [1,2,3,4,5]. Moreover, a multidisciplinary biosystematics is imperative to accurately identify reduviids and employ them against a particular insect pest [4,5,6,7]. Though the literature available on multi-disciplinary biosystematics of Reduviidae including molecular tools is available at family or species level it is very meager [5,8,9,10,11,12,13,14].
Curtis established the genus Coranus with Cimex subapterus De Geer as the type species. Coranus is one of the largest genera of subfamily Harpactorinae in the family Reduviidae with 100 known species worldwide [2,3,4,15]. The members of Coranus are widely distributed and occur throughout the Eastern Hemisphere with 30 Palaearctic, 21 Oriental, 41 Ethiopian and 17 Australian species [11]. Malipatil revised the Australian Coranus with redescription of seven species, description of eight new species and formulated key to identify the fifteen species [16]. Liu . inferred the phylogenetic relationship of the harpactorine genus Velinoides Matsmura with Coranus Curtis based on three mitochondrial genes (cyt b, CoI and 16S rRNA) and one nuclear (28S rRNA) gene [11]. Since they found molecular affinity between these two genera supported with morphological and cytogenetic characteristics they validated the status of genus Velinoides and its phylogenetic affinity with the genus Coranus. They suggested that these two genera could be two subgenera of the genus Coranus. They further reported that the 28S rDNA gene alone might not be an optimal marker for the phylogeny of the genus Coranus. Though twelve species of Coranus have been recorded from India none of its gene sequence is available. Except the work of Liu et al. no work on the molecular phylogenetics of the genus Coranus is available [4,11].
Hence, this study was undertaken based on the sequences of three mitochondrial genes, 16S rRNA, Cyt b, COI and one nuclear gene, 28S rRNA of thirteen species of Coranus Curtis and probably two ecotypes of Coranus callosus Stål from four countries viz., Australia, Brunei, China and Nigeria of three continents viz., Africa, Asia and Australia and probably two ecotypes of C. callosus Stål from western Australia downloaded from the GenBank (Table 1). The inclusion of Coranus species from four countries of three continents further enhances the scope of the work at the intraspecific level and the understanding on the role of geographical isolation in biosystematics.
Species |
Locality |
Coranus sp.1 |
Brunei |
Coranus sp.2 |
Nigeria: Ondo |
Coranus sp.3 |
Australia : South Australia |
Coranus lativentris Jakovlev |
China: Xiaowutai Mt., Hebei |
Coranus hammarstroemi Reuter |
China: Lvliang Mt., Shanxi |
Coranus dilatatus (Matsumura) |
China: Lvliang Mt., Shanxi |
Coranus marginatus Hsiao |
China: Yingjiang, Dehong, Yunnan |
Coranus emodicus Kiritschenko |
China: Yingjiang, Dehong, Yunnan |
Coranus fuscipennis Reuter |
China: Yunji Mt., Xinfeng, Guangdong |
Coranus sichuensis Hsiao & Ren |
China: Tengchong, Baoshan, Yunnan |
Coranus spiniscutis Reuter |
China: Jinghong, Xishuangbanna,Yunnan |
Coranus subapterus (De Geer) |
China: Tianchi, Urumchi, Xinjiang |
Coranus callosus Stål* |
Australia: Western |
Coranus callosus Stål** |
Australia: Western |
Mitochondrial and nuclear genes |
Species |
Genbank accession number |
|
Coranus sp.1 |
JQ888411.1 |
Coranus sp.2 |
JQ888412.1 |
|
Coranus sp.3 |
JQ888413.1 |
|
Coranus lativentris Jakovlev |
EU128688.1 |
|
Coranus hammarstroemi Reuter |
EU128689.1 |
|
Coranus dilatatus (Matsumura) |
EU128690.1 |
|
Coranus marginatus Hsiao |
EU128691.1 |
|
Coranus emodicus Kiritschenko |
EU128692.1 |
|
Coranus fuscipennis Reuter |
EU128693.1 |
|
Coranus sichuensis Hsiao & Ren |
EU128694.1 |
|
Coranus spiniscutis Reuter |
EU128695.1 |
|
Coranus subapterus (De Geer) |
EU128696.1 |
|
Coranus callosus Stål* |
FJ230433.1 |
|
|
Coranus lativentris Jakovlev |
EU128710.1 |
Coranus hammarstroemi Reuter |
EU128711.1 |
|
Coranus dilatatus (Matsumura) |
EU128712.1 |
|
Coranus marginatus Hsiao |
EU128713.1 |
|
Coranus fuscipennis Reuter |
EU128714.1 |
|
Coranus sichuensis Hsiao & Ren |
EU128715.1 |
|
Coranus spiniscutis Reuter |
EU128716.1 |
|
Coranus subapterus (De Geer) |
EU128717.1 |
|
|
Coranus sp.1 |
JQ888572.1 |
Coranus sp.2 |
JQ888573.1 |
|
Coranus sp.3 |
JQ888574.1 |
|
Coranus callosus Stål* |
JQ888571.1 |
|
Coranus callosus Stål** |
JQ942321.1 |
|
28S ribosomal RNA |
Coranus sp.1 |
JQ888911.1 |
Coranus sp.2 |
JQ888756.1 |
|
Coranus lativentris Jakovlev |
EU128677.1 |
|
Coranus hammarstroemi Reuter |
EU128678.1 |
|
Coranus dilatatus (Matsumura) |
EU128679.1 |
|
Coranus marginatus Hsiao |
EU128680.1 |
|
Coranus emodicus Kiritschenko |
EU128681.1 |
|
Coranus fuscipennis Reuter |
EU128682.1 |
|
Coranus sichuensis Hsiao & Ren |
EU128683.1 |
|
Coranus spiniscutis Reuter |
EU128684.1 |
|
Coranus subapterus (De Geer) |
EU128685.1 |
|
Coranus callosus Stål* |
FJ230594.1 |
The substitution type based nucleotide sequences and the codon positions included were 1st+2nd+3rd+Noncoding and all the positions containing gaps and missing data were eliminated in all the five methods. Five phylograms were thus constructed based on maximum likelihood (ML), neighbor-joining (N-J), maximum evolution (ME), UPGMA and maximum parsimony (MP) methods for three mitochondrial genes, 16S rRNA, Cyt b and Cyt c oxidase subunit I and one nuclear gene, 28S rRNA. The trees were analyzed based on the arrangement of each species in the tree.
The NJ (Figure 2) and ME trees (Figure 3) replicate the second major cluster as in ML tree. However, in the first major cluster, the positions of C. dilatatus and C. hammarstroemi vary. Though almost a similar kind of phylogeny is observed for UPGMA (Figure 4) and MP methods (Figure 5) slight deviations were found in relation to C. dilatatus in UPGMA and C. hammarstroemi and C. callosus in MP tree.
Coranus sp.2 of Australia instead of clustering with C. callosus of Australia clustered with Coranus sp.3 of Nigeria and Coranus sp.1 of Brunei. These species exhibit affinity despite their geographical isolation as observed by Mahendran et al. in silk producing insects and Ambrose et al. in R. fuscipes (Fabricius) of India with R. segmentarius (Germar) of South Africa [5,28].
Cyt b. The five phylograms observed for eight Coranus species of China except C. emodicus formed into two major clusters (Figure 6,7,8,9 and 10). The first cluster had C. spiniscutis, C. hammarstroemi, C. subapterus and C. sichuensis and the second cluster had C. dilatatus, C. lativentris, C. fuscipennis and C. marginatus with slight modification in different phylograms, revealing monophyly as observed by Liu et al. in Coranus species of China, Baskar et al. and Ambrose et al. in Rhynocoris species of India [5,11,29].
Cyt c. The five phylograms (Figure 11,12,13,14 and 15) of Cyt c gene of three undetermined species of Coranus from Australia, Brunei and Nigeria C. callosus from western Australia, probably from two localities, i.e., two ecotypes revealed affinity between Coranus sp.1 of Brunei with Coranus sp.2 of Australia. Coranus sp.2 of Australia thus instead of clustering with C. callosus of Australia aligns with that of Coranus sp.1 from Brunei. Similarly two ecotypes of Coranus callosus of Western Australia instead of
28S rRNA. Five phylograms (Figure16,17,18,19,20) were constructed for twelve species of Coranus, i.e., except Coranus sp.3. In maximum likelihood method (Figure 16), all the nine Coranus species from China formed a major cluster. Coranus sp.1 of Brunei diversified as a separate lineage. From this common node a sub cluster formed with two Australian species viz., C. callosus and Coranus sp.2. An almost similar kind of phylogency is revealed by NJ, ME, UPGMA and MP methods (Figure 17,18,19,20). Thus, the affinity between the nine species of Coranus from China and that of two species from Australian is well pronounced. Although Liu et al. (2009) reported that 28S rRNA is a highly conserved gene and may not be an optimum molecular marker for Coranus, the present analysis contradicts their view and suggests its usefulness in phylogenetics [11].
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