journal article
LitStream Collection
Phylogeny and revision of the Oriental leafhopper genus Amritodus (Hemiptera: Cicadellidae: Idiocerini)
2019 Zoological Journal of the Linnean Society
doi: 10.1093/zoolinnean/zlz129
Abstract The phylogeny of the Oriental leafhopper genus Amritodus is reconstructed, for the first time, based on 47 discrete morphological characters and DNA sequence data from one nuclear and two mitochondrial genes. The phylogenetic results show that Amritodus is not monophyletic, and its concept is narrowed here to include four species: Amritodus atkinsoni, Amritodus brevis, Amritodus brevistylus and Amritodus saeedi. The phylogenetic results support establishment of a new genus, Paramritodus gen. nov., with three new species,Paramritodus triangulus sp. nov. (type species), Paramritodus introflexus sp. nov., Paramritodus spatiosus sp. nov. and three species previously included in Amritodus: Paramritodus pistacious comb. nov., Paramritodus flavocapitatus comb. nov. and Paramritodus podocarpus comb. nov. In addition, Amritodus flavoscutatus is transferred from Amritodus to Hyalinocerus as Hyalinocerus flavoscutatus comb. nov. Keys to species of Amritodus and Paramritodus are provided. Auchenorrhyncha, Eurymelinae, Homoptera, Idiocerinae, morphology, phylogenetic analysis, taxonomy INTRODUCTION Idiocerini Baker, 1915 is one of the largest tribes of arboreal leafhoppers. The relationships among genera of this tribe remain poorly understood. Krishnankutty & Dietrich (2011) were the first to study idiocerine leafhoppers using phylogenetic methods. Their analysis supported the monophyly of Nesocerus Freytag & Knight, 1966 and resolved relationships among species of that genus. Moreover, in her unpublished PhD dissertation, Krishnankutty (2012) presented results of a molecular phylogenetic analysis of Idiocerini and related groups that included representatives from most biogeographical regions of the world. That analysis and a more recent phylogenomic analysis of Membracoidea (Dietrich et al., 2017) indicated that the endemic Australian group Eurymelinae is derived from Idiocerinae. Dietrich & Thomas (2018) treated Idiocerinae as a junior synonym of Eurymelinae, which now includes both former subfamilies, Eurymelinae and Idiocerinae, as its tribes. Anufriev (1970) established the genus Amritodus based on the species Idiocerus atkinsoni Lethierry, 1889, from India and Pakistan. Amritodus is widespread in the Oriental region, and its species often feed on Anacardiaceae (Viraktamath, 1997; Gnaneswaran et al., 2007). Viraktamath (1976) added Amritodus brevistylus Viraktamath and Amritodus mudigerensis Viraktamath from India. Viraktamath & Murphy (1980) transferred A. mudigerensis to Busoniomimus Maldonado-Capriles, 1977. Ahmed et al. (1980) described Amritodus saeediAhmed, Naheed & Ahmed, 1980 from Pakistan. Huang & Maldonado-Capriles (1992) reported Amritodus for the first time in China and described Amritodus pistacious Huang & Maldonado-Capriles, 1992. Viraktamath (1997) described Amritodus brevis from India, and suggested that A. pistacious is sufficiently different from other species of Amritodus that it should be removed from the genus. Cai & Shen (1998) described Amritodus flavoscutatus, Cai et al. (2001) added Amritodus flavocapitatus Cai & He, 2001, and Zhang & Li (2010) described Amritodus podocarpus Zhang & Li, 2010 from China. Until this study, Amritodus included eight species. Amritodus was previously defined based on characters of the external morphology and male genitalia, but in our studies, we found that some species do not conform to the diagnosis of this genus. In the present paper, the phylogeny of Amritodus is reconstructed based on DNA sequence data and adult morphological data, and a new classification scheme for Amritodus is proposed based on these phylogenetic results. Furthermore, a new genus is established to include some species excluded from Amritodus together with some additional new species, and Amritodus flavoscutatus is transferred to Hyalinocerus. MATERIAL AND METHODS Taxon sampling Nineteen taxa were included in the datasets analysed (Table 1). The ingroup comprised all eight previously recognized Amritodus species. The outgroup comprised 11 species belonging to eight related genera based on a previous phylogenetic analysis (Xue Q, Dietrich C & Zhang Y, in prep.), including Anidiocerus brevispinus Xue & Zhang, Busonia albilateralis Maldonado-Capriles, Busoniomimus polydoros (Kirkaldy), Chunra quadrispinosa Zhang, Li & Xu, Hyalinocerus nigrimaculatus Zhang & Li, Idioscopus nagpurensis (Pruthi), Idioscopus nitidulus (Walker), Jogocerus concavus Xue & Zhang and three undescribed species. Five taxa were represented only by morphological data. Specimens of all taxa except A. podocarpus were examined. The character states for A. podocarpus were determined based on the detailed description and illustrations by Zhang & Li (2010). Table 1. GenBank accession numbers for the respective DNA fragments Taxon Location 28S D2 16S COI Chunra quadrispinosa China: Hainan MK064017 MK063942 MK055892 Busonia albilateralis China: Yunnan MK064007 MK063938 MK055888 Anidiocerus brevispinus China: Shaanxi MK064000 MK063932 MK055881 Busoniomimus polydoros Australia: QLD MK064009 MK063939 MK055889 Hyalinocerus flavoscutatus comb. nov. China: Guizhou MK063996 MK063928 MK055878 Hyalinocerus nigrimaculatus China: Shaanxi MK064025 MK063950 MK055899 Idioscopus nagpurensis China: Yunnan MK064032 MK063956 MK055903 Idioscopus nitidulus China: Yunnan MK064033 NC029203 MK055904 Jogocerus concavus China: Yunnan MK064039 MK063963 MK055910 Amritodus atkinsoni India – – HQ268819 Amritodus brevis China: Yunnan MK063994 MK063926 MK055876 Amritodus brevistylus Sri Lanka – – HQ268817 Amritodus saeedi India – – – Paramritodus introflexus sp. nov. China: Yunnan – – – Paramritodus flavocapitatus comb. nov. China: Zhejiang MK063995 MK063927 MK055877 Paramritodus pistacious comb. nov. China: Taiwan – – – Paramritodus podocarpus comb. nov. China: Guizhou – – – Paramritodus spatiosus sp. nov. China: Yunnan – – – Paramritodus triangulus sp. nov. China: Yunnan MK064057 MK063977 MK055926 Taxon Location 28S D2 16S COI Chunra quadrispinosa China: Hainan MK064017 MK063942 MK055892 Busonia albilateralis China: Yunnan MK064007 MK063938 MK055888 Anidiocerus brevispinus China: Shaanxi MK064000 MK063932 MK055881 Busoniomimus polydoros Australia: QLD MK064009 MK063939 MK055889 Hyalinocerus flavoscutatus comb. nov. China: Guizhou MK063996 MK063928 MK055878 Hyalinocerus nigrimaculatus China: Shaanxi MK064025 MK063950 MK055899 Idioscopus nagpurensis China: Yunnan MK064032 MK063956 MK055903 Idioscopus nitidulus China: Yunnan MK064033 NC029203 MK055904 Jogocerus concavus China: Yunnan MK064039 MK063963 MK055910 Amritodus atkinsoni India – – HQ268819 Amritodus brevis China: Yunnan MK063994 MK063926 MK055876 Amritodus brevistylus Sri Lanka – – HQ268817 Amritodus saeedi India – – – Paramritodus introflexus sp. nov. China: Yunnan – – – Paramritodus flavocapitatus comb. nov. China: Zhejiang MK063995 MK063927 MK055877 Paramritodus pistacious comb. nov. China: Taiwan – – – Paramritodus podocarpus comb. nov. China: Guizhou – – – Paramritodus spatiosus sp. nov. China: Yunnan – – – Paramritodus triangulus sp. nov. China: Yunnan MK064057 MK063977 MK055926 ‘–’ indicates no sequence available. Open in new tab Table 1. GenBank accession numbers for the respective DNA fragments Taxon Location 28S D2 16S COI Chunra quadrispinosa China: Hainan MK064017 MK063942 MK055892 Busonia albilateralis China: Yunnan MK064007 MK063938 MK055888 Anidiocerus brevispinus China: Shaanxi MK064000 MK063932 MK055881 Busoniomimus polydoros Australia: QLD MK064009 MK063939 MK055889 Hyalinocerus flavoscutatus comb. nov. China: Guizhou MK063996 MK063928 MK055878 Hyalinocerus nigrimaculatus China: Shaanxi MK064025 MK063950 MK055899 Idioscopus nagpurensis China: Yunnan MK064032 MK063956 MK055903 Idioscopus nitidulus China: Yunnan MK064033 NC029203 MK055904 Jogocerus concavus China: Yunnan MK064039 MK063963 MK055910 Amritodus atkinsoni India – – HQ268819 Amritodus brevis China: Yunnan MK063994 MK063926 MK055876 Amritodus brevistylus Sri Lanka – – HQ268817 Amritodus saeedi India – – – Paramritodus introflexus sp. nov. China: Yunnan – – – Paramritodus flavocapitatus comb. nov. China: Zhejiang MK063995 MK063927 MK055877 Paramritodus pistacious comb. nov. China: Taiwan – – – Paramritodus podocarpus comb. nov. China: Guizhou – – – Paramritodus spatiosus sp. nov. China: Yunnan – – – Paramritodus triangulus sp. nov. China: Yunnan MK064057 MK063977 MK055926 Taxon Location 28S D2 16S COI Chunra quadrispinosa China: Hainan MK064017 MK063942 MK055892 Busonia albilateralis China: Yunnan MK064007 MK063938 MK055888 Anidiocerus brevispinus China: Shaanxi MK064000 MK063932 MK055881 Busoniomimus polydoros Australia: QLD MK064009 MK063939 MK055889 Hyalinocerus flavoscutatus comb. nov. China: Guizhou MK063996 MK063928 MK055878 Hyalinocerus nigrimaculatus China: Shaanxi MK064025 MK063950 MK055899 Idioscopus nagpurensis China: Yunnan MK064032 MK063956 MK055903 Idioscopus nitidulus China: Yunnan MK064033 NC029203 MK055904 Jogocerus concavus China: Yunnan MK064039 MK063963 MK055910 Amritodus atkinsoni India – – HQ268819 Amritodus brevis China: Yunnan MK063994 MK063926 MK055876 Amritodus brevistylus Sri Lanka – – HQ268817 Amritodus saeedi India – – – Paramritodus introflexus sp. nov. China: Yunnan – – – Paramritodus flavocapitatus comb. nov. China: Zhejiang MK063995 MK063927 MK055877 Paramritodus pistacious comb. nov. China: Taiwan – – – Paramritodus podocarpus comb. nov. China: Guizhou – – – Paramritodus spatiosus sp. nov. China: Yunnan – – – Paramritodus triangulus sp. nov. China: Yunnan MK064057 MK063977 MK055926 ‘–’ indicates no sequence available. Open in new tab The specimens used in this study are deposited in the following archives: the National Zoological Museum of China, Institute of Zoology, Chinese Academy of Sciences, Beijing, China (IOZ); the Entomological Museum, Northwest A&F University, Yangling, China (NWAFU); the Queen Sirikit Botanical Garden, Chiang Mai, Thailand (QSBG); the Shanghai Entomological Museum, Chinese Academy of Sciences, Shanghai, China (SEMCAS); the Department of Entomology, University of Agricultural Sciences, Bangalore, India (UASB); and the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA (USNM), as indicated in the text. DNA sequencing Adult specimens were preserved in 95% or anhydrous ethanol before molecular analyses. We selected one nuclear gene, 28S D2 (~700 bp), and two mitochondrial genes, 16S (~550 bp) and COI (658 bp) gene fragments. These genes have been used previously to reconstruct phylogenetic relationships among Membracoidea (e.g. Dietrich et al., 2001; Zahniser & Dietrich, 2010; Krishnankutty & Dietrich, 2011; Wang et al., 2016, 2017; Evangelista et al., 2017). Total genomic DNA of individual specimens was extracted from either the abdomen or thoracic muscles using Qiagen DNEasy Kits (Qiagen, Valencia, CA, USA). Fragments of 28S D2 were amplified by PCR in 25 μL reaction volumes with the following cycling protocol: 94 °C for 3 min, then 30 cycles of 94 °C for 1 min, 52–57 °C for 1 min, 72 °C for 1.5–2.0 min, and one final extension at 72 °C for 7 min. COI was amplified as follows: 3 min at 94 °C; five cycles of 1 min at 94 °C, 1.5 min at 45 °C and 1.5 min at 72 °C; 35 cycles of 1 min at 94 °C, 1 min at 52–55 °C and 1 min at 72 °C; and 5 min at 72 °C. 16S was amplified as follows: 5 min at 94 °C; 11 cycles of 92 °C for 1 min, 48 °C for 1 min and 72 °C for 1.5 min; 30 cycles of 92 °C for 1 min, 54–56 °C for 35 s and 72 °C for 2 min; and 72 °C for 7 min. The PCR primers are listed in Table 2. Total genomic DNA was stored at −20 °C before PCR and Sanger sequencing using the same primer pairs. Table 2. Primers used in this study Primer Primer sequence (5′–3′) Source 28SD2F AGTCGKGTTGCTTGAKAGTGCAG Dietrich et al. (2001) 28SD2R TTCAATTTCATTKCGCCTT Dietrich et al. (2001) 16SF(LR-J-12887) CCGGTYTGAACTCARATCAWGT Dietrich et al. (1997) 16SR(LR-N-13398) CTGTTTAWCAAAAACATTTC Dietrich et al. (1997) COIF(LCO1490) GGTCAACAAATCATAAAGATATTGG Folmer et al. (1994) COIR(HCO2198) TAAACTTCAGGGTGACCAAAAAATCA Folmer et al. (1994) Primer Primer sequence (5′–3′) Source 28SD2F AGTCGKGTTGCTTGAKAGTGCAG Dietrich et al. (2001) 28SD2R TTCAATTTCATTKCGCCTT Dietrich et al. (2001) 16SF(LR-J-12887) CCGGTYTGAACTCARATCAWGT Dietrich et al. (1997) 16SR(LR-N-13398) CTGTTTAWCAAAAACATTTC Dietrich et al. (1997) COIF(LCO1490) GGTCAACAAATCATAAAGATATTGG Folmer et al. (1994) COIR(HCO2198) TAAACTTCAGGGTGACCAAAAAATCA Folmer et al. (1994) Open in new tab Table 2. Primers used in this study Primer Primer sequence (5′–3′) Source 28SD2F AGTCGKGTTGCTTGAKAGTGCAG Dietrich et al. (2001) 28SD2R TTCAATTTCATTKCGCCTT Dietrich et al. (2001) 16SF(LR-J-12887) CCGGTYTGAACTCARATCAWGT Dietrich et al. (1997) 16SR(LR-N-13398) CTGTTTAWCAAAAACATTTC Dietrich et al. (1997) COIF(LCO1490) GGTCAACAAATCATAAAGATATTGG Folmer et al. (1994) COIR(HCO2198) TAAACTTCAGGGTGACCAAAAAATCA Folmer et al. (1994) Primer Primer sequence (5′–3′) Source 28SD2F AGTCGKGTTGCTTGAKAGTGCAG Dietrich et al. (2001) 28SD2R TTCAATTTCATTKCGCCTT Dietrich et al. (2001) 16SF(LR-J-12887) CCGGTYTGAACTCARATCAWGT Dietrich et al. (1997) 16SR(LR-N-13398) CTGTTTAWCAAAAACATTTC Dietrich et al. (1997) COIF(LCO1490) GGTCAACAAATCATAAAGATATTGG Folmer et al. (1994) COIR(HCO2198) TAAACTTCAGGGTGACCAAAAAATCA Folmer et al. (1994) Open in new tab Sequences were assembled and edited with MEGA v.6 (Tamura et al., 2013). Sequences were aligned in MAFFT v.7.037 (Katoh & Standley, 2013) using the G-INS-i algorithm. Molecular and morphological datasets were merged using SEQUENCEMATRIX v.1.7.8 (Vaidya et al., 2011). GenBank accession numbers are provided in Table 1. Morphological characters The male abdomen was removed from the specimen and treated with 8–10% NaOH for 24 h, rinsed with water and then transferred to glycerol for further dissection and examination. After examination, the dissected genitalia were stored in a microvial with fresh glycerol and pinned below the specimen from which the abdomen was removed. Photographs and drawings were prepared using a Microvision system and CARTOGRAPH v.8.0.6 automontage software, a Nikon SMZ1500 dissecting microscope, an Olympus BH-2 stereoscopic microscope and a Zeiss stereoscopic microscope, and then adjusted in Adobe Photoshop. Morphological terminology mainly follows Zhang (1990) and Dietrich (2005), except for the apodemes of the abdomen, which follows Hamilton (1980). Fourty-seven morphological characters were used in the phylogenetic analysis, composed of 41 binary and six multistate characters, including 11 from the head, six from the forewing and hindleg and 30 from the male genitalia. Morphological data were compiled using MESQUITE v.3.31 (Maddison & Maddison, 2017). Inapplicable characters are indicated as ‘–’. All characters were treated as unordered and of equal weight. The data matrix is given in Table 3. Table 3. Morphological character matrix Species 01 02 03 04 05 06 07 08 09 10 11 12 13 14 15 16 17 18 19 02 21 22 23 24 Chunra quadrispinosa 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 - - 0 0 0 0 Busonia albilateralis 0 0 0 1 0 1 - - 0 0 0 1 0 1 0 1 0 0 - - 0 0 0 1 Anidiocerus brevispinus 1 1 0 0 0 0 1 1 1 1 0 0 1 1 1 1 1 1 2 0 0 0 0 0 Busoniomimus polydoros 1 1 0 0 0 0 0 1 1 1 0 0 1 1 1 1 1 1 2 1 1 0 1 0 Hyalinocerus flavoscutatus comb. nov. 1 0 1 1 1 0 1 1 1 0 1 0 1 1 1 1 1 1 2 0 0 0 0 0 Hyalinocerus nigrimaculatus 1 0 1 1 0 0 0 0 1 0 0 0 1 1 1 1 1 0 - - 0 0 1 1 Idioscopus nagpurensis 1 1 0 0 0 0 0 0 0 1 0 0 1 1 1 1 1 0 - - 0 0 0 0 Idioscopus nitidulus 1 1 0 0 0 0 1 1 1 1 1 0 1 0 1 1 2 0 - - 0 0 0 0 Jogocerus concavus 1 0 1 1 0 0 0 1 0 0 0 0 1 0 1 1 0 1 2 1 0 1 1 0 Amritodus atkinsoni 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Amritodus brevis 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Amritodus brevistylus 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Amritodus saeedi 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Paramritodus introflexus sp. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 1 0 0 0 1 0 Paramritodus flavocapitatus comb. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Paramritodus pistacious comb. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Paramritodus podocarpus comb. nov. 1 1 0 0 0 0 ? ? 0 0 0 0 0 1 1 1 ? 1 0 0 0 0 1 0 Paramritodus spatiosus sp. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Paramritodus triangulus sp. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Species 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 Chunra quadrispinosa 0 1 0 0 0 0 0 0 0 1 2 0 0 0 0 0 0 0 - - - 0 0 Busonia albilateralis 0 1 0 0 0 0 0 0 1 - - - - 0 0 0 0 1 - - - 1 - Anidiocerus brevispinus 0 0 2 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 - - - 1 - Busoniomimus polydoros 0 1 3 0 0 0 0 1 1 - - - - 0 0 0 1 0 - - - 0 0 Hyalinocerus flavoscutatus comb. nov. 1 1 2 1 1 0 0 0 1 - - - - 0 0 0 1 0 - - - 1 - Hyalinocerus nigrimaculatus 1 1 3 0 1 0 0 0 0 2 0 0 1 0 0 0 0 0 - - - 1 - Idioscopus nagpurensis 0 0 2 0 0 0 0 0 0 1 1 1 1 0 0 0 1 0 - - - 0 0 Idioscopus nitidulus 0 0 2 0 0 0 0 0 0 1 0 1 1 0 0 0 1 0 - - - 0 0 Jogocerus concavus 0 1 2 0 0 0 1 0 0 0 1 0 1 0 0 0 1 0 - - - 0 0 Amritodus atkinsoni 0 0 1 0 1 1 0 0 1 - - - - 1 0 0 1 1 1 1 0 0 0 Amritodus brevis 0 2 1 0 1 0 0 0 1 - - - - 1 0 1 1 1 1 1 0 0 0 Amritodus brevistylus 0 0 1 0 1 0 0 0 0 0 2 0 0 1 0 0 1 1 1 1 0 0 0 Amritodus saeedi 0 0 1 0 1 0 0 0 1 - - - - 1 0 0 1 1 1 1 0 0 0 Paramritodus introflexus sp. nov. 0 1 3 0 0 0 0 0 0 0 2 0 0 1 1 0 2 1 0 1 1 0 1 Paramritodus flavocapitatus comb. nov. 0 1 3 0 0 0 0 0 1 - - - - 1 1 0 2 1 0 0 0 0 1 Paramritodus pistacious comb. nov. 0 1 3 0 0 0 0 0 1 - - - - 1 1 0 2 1 0 0 1 0 1 Paramritodus podocarpus comb. nov. 0 1 3 0 0 0 0 0 1 - - - - 1 1 0 2 1 0 1 0 1 - Paramritodus spatiosus sp. nov. 0 1 3 0 0 0 0 0 0 0 2 1 0 1 1 0 1 1 0 1 1 0 0 Paramritodus triangulus sp. nov. 0 1 3 0 0 0 0 0 0 0 2 0 0 1 1 0 2 1 0 1 1 0 1 Species 01 02 03 04 05 06 07 08 09 10 11 12 13 14 15 16 17 18 19 02 21 22 23 24 Chunra quadrispinosa 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 - - 0 0 0 0 Busonia albilateralis 0 0 0 1 0 1 - - 0 0 0 1 0 1 0 1 0 0 - - 0 0 0 1 Anidiocerus brevispinus 1 1 0 0 0 0 1 1 1 1 0 0 1 1 1 1 1 1 2 0 0 0 0 0 Busoniomimus polydoros 1 1 0 0 0 0 0 1 1 1 0 0 1 1 1 1 1 1 2 1 1 0 1 0 Hyalinocerus flavoscutatus comb. nov. 1 0 1 1 1 0 1 1 1 0 1 0 1 1 1 1 1 1 2 0 0 0 0 0 Hyalinocerus nigrimaculatus 1 0 1 1 0 0 0 0 1 0 0 0 1 1 1 1 1 0 - - 0 0 1 1 Idioscopus nagpurensis 1 1 0 0 0 0 0 0 0 1 0 0 1 1 1 1 1 0 - - 0 0 0 0 Idioscopus nitidulus 1 1 0 0 0 0 1 1 1 1 1 0 1 0 1 1 2 0 - - 0 0 0 0 Jogocerus concavus 1 0 1 1 0 0 0 1 0 0 0 0 1 0 1 1 0 1 2 1 0 1 1 0 Amritodus atkinsoni 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Amritodus brevis 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Amritodus brevistylus 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Amritodus saeedi 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Paramritodus introflexus sp. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 1 0 0 0 1 0 Paramritodus flavocapitatus comb. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Paramritodus pistacious comb. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Paramritodus podocarpus comb. nov. 1 1 0 0 0 0 ? ? 0 0 0 0 0 1 1 1 ? 1 0 0 0 0 1 0 Paramritodus spatiosus sp. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Paramritodus triangulus sp. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Species 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 Chunra quadrispinosa 0 1 0 0 0 0 0 0 0 1 2 0 0 0 0 0 0 0 - - - 0 0 Busonia albilateralis 0 1 0 0 0 0 0 0 1 - - - - 0 0 0 0 1 - - - 1 - Anidiocerus brevispinus 0 0 2 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 - - - 1 - Busoniomimus polydoros 0 1 3 0 0 0 0 1 1 - - - - 0 0 0 1 0 - - - 0 0 Hyalinocerus flavoscutatus comb. nov. 1 1 2 1 1 0 0 0 1 - - - - 0 0 0 1 0 - - - 1 - Hyalinocerus nigrimaculatus 1 1 3 0 1 0 0 0 0 2 0 0 1 0 0 0 0 0 - - - 1 - Idioscopus nagpurensis 0 0 2 0 0 0 0 0 0 1 1 1 1 0 0 0 1 0 - - - 0 0 Idioscopus nitidulus 0 0 2 0 0 0 0 0 0 1 0 1 1 0 0 0 1 0 - - - 0 0 Jogocerus concavus 0 1 2 0 0 0 1 0 0 0 1 0 1 0 0 0 1 0 - - - 0 0 Amritodus atkinsoni 0 0 1 0 1 1 0 0 1 - - - - 1 0 0 1 1 1 1 0 0 0 Amritodus brevis 0 2 1 0 1 0 0 0 1 - - - - 1 0 1 1 1 1 1 0 0 0 Amritodus brevistylus 0 0 1 0 1 0 0 0 0 0 2 0 0 1 0 0 1 1 1 1 0 0 0 Amritodus saeedi 0 0 1 0 1 0 0 0 1 - - - - 1 0 0 1 1 1 1 0 0 0 Paramritodus introflexus sp. nov. 0 1 3 0 0 0 0 0 0 0 2 0 0 1 1 0 2 1 0 1 1 0 1 Paramritodus flavocapitatus comb. nov. 0 1 3 0 0 0 0 0 1 - - - - 1 1 0 2 1 0 0 0 0 1 Paramritodus pistacious comb. nov. 0 1 3 0 0 0 0 0 1 - - - - 1 1 0 2 1 0 0 1 0 1 Paramritodus podocarpus comb. nov. 0 1 3 0 0 0 0 0 1 - - - - 1 1 0 2 1 0 1 0 1 - Paramritodus spatiosus sp. nov. 0 1 3 0 0 0 0 0 0 0 2 1 0 1 1 0 1 1 0 1 1 0 0 Paramritodus triangulus sp. nov. 0 1 3 0 0 0 0 0 0 0 2 0 0 1 1 0 2 1 0 1 1 0 1 Open in new tab Table 3. Morphological character matrix Species 01 02 03 04 05 06 07 08 09 10 11 12 13 14 15 16 17 18 19 02 21 22 23 24 Chunra quadrispinosa 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 - - 0 0 0 0 Busonia albilateralis 0 0 0 1 0 1 - - 0 0 0 1 0 1 0 1 0 0 - - 0 0 0 1 Anidiocerus brevispinus 1 1 0 0 0 0 1 1 1 1 0 0 1 1 1 1 1 1 2 0 0 0 0 0 Busoniomimus polydoros 1 1 0 0 0 0 0 1 1 1 0 0 1 1 1 1 1 1 2 1 1 0 1 0 Hyalinocerus flavoscutatus comb. nov. 1 0 1 1 1 0 1 1 1 0 1 0 1 1 1 1 1 1 2 0 0 0 0 0 Hyalinocerus nigrimaculatus 1 0 1 1 0 0 0 0 1 0 0 0 1 1 1 1 1 0 - - 0 0 1 1 Idioscopus nagpurensis 1 1 0 0 0 0 0 0 0 1 0 0 1 1 1 1 1 0 - - 0 0 0 0 Idioscopus nitidulus 1 1 0 0 0 0 1 1 1 1 1 0 1 0 1 1 2 0 - - 0 0 0 0 Jogocerus concavus 1 0 1 1 0 0 0 1 0 0 0 0 1 0 1 1 0 1 2 1 0 1 1 0 Amritodus atkinsoni 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Amritodus brevis 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Amritodus brevistylus 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Amritodus saeedi 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Paramritodus introflexus sp. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 1 0 0 0 1 0 Paramritodus flavocapitatus comb. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Paramritodus pistacious comb. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Paramritodus podocarpus comb. nov. 1 1 0 0 0 0 ? ? 0 0 0 0 0 1 1 1 ? 1 0 0 0 0 1 0 Paramritodus spatiosus sp. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Paramritodus triangulus sp. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Species 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 Chunra quadrispinosa 0 1 0 0 0 0 0 0 0 1 2 0 0 0 0 0 0 0 - - - 0 0 Busonia albilateralis 0 1 0 0 0 0 0 0 1 - - - - 0 0 0 0 1 - - - 1 - Anidiocerus brevispinus 0 0 2 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 - - - 1 - Busoniomimus polydoros 0 1 3 0 0 0 0 1 1 - - - - 0 0 0 1 0 - - - 0 0 Hyalinocerus flavoscutatus comb. nov. 1 1 2 1 1 0 0 0 1 - - - - 0 0 0 1 0 - - - 1 - Hyalinocerus nigrimaculatus 1 1 3 0 1 0 0 0 0 2 0 0 1 0 0 0 0 0 - - - 1 - Idioscopus nagpurensis 0 0 2 0 0 0 0 0 0 1 1 1 1 0 0 0 1 0 - - - 0 0 Idioscopus nitidulus 0 0 2 0 0 0 0 0 0 1 0 1 1 0 0 0 1 0 - - - 0 0 Jogocerus concavus 0 1 2 0 0 0 1 0 0 0 1 0 1 0 0 0 1 0 - - - 0 0 Amritodus atkinsoni 0 0 1 0 1 1 0 0 1 - - - - 1 0 0 1 1 1 1 0 0 0 Amritodus brevis 0 2 1 0 1 0 0 0 1 - - - - 1 0 1 1 1 1 1 0 0 0 Amritodus brevistylus 0 0 1 0 1 0 0 0 0 0 2 0 0 1 0 0 1 1 1 1 0 0 0 Amritodus saeedi 0 0 1 0 1 0 0 0 1 - - - - 1 0 0 1 1 1 1 0 0 0 Paramritodus introflexus sp. nov. 0 1 3 0 0 0 0 0 0 0 2 0 0 1 1 0 2 1 0 1 1 0 1 Paramritodus flavocapitatus comb. nov. 0 1 3 0 0 0 0 0 1 - - - - 1 1 0 2 1 0 0 0 0 1 Paramritodus pistacious comb. nov. 0 1 3 0 0 0 0 0 1 - - - - 1 1 0 2 1 0 0 1 0 1 Paramritodus podocarpus comb. nov. 0 1 3 0 0 0 0 0 1 - - - - 1 1 0 2 1 0 1 0 1 - Paramritodus spatiosus sp. nov. 0 1 3 0 0 0 0 0 0 0 2 1 0 1 1 0 1 1 0 1 1 0 0 Paramritodus triangulus sp. nov. 0 1 3 0 0 0 0 0 0 0 2 0 0 1 1 0 2 1 0 1 1 0 1 Species 01 02 03 04 05 06 07 08 09 10 11 12 13 14 15 16 17 18 19 02 21 22 23 24 Chunra quadrispinosa 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 - - 0 0 0 0 Busonia albilateralis 0 0 0 1 0 1 - - 0 0 0 1 0 1 0 1 0 0 - - 0 0 0 1 Anidiocerus brevispinus 1 1 0 0 0 0 1 1 1 1 0 0 1 1 1 1 1 1 2 0 0 0 0 0 Busoniomimus polydoros 1 1 0 0 0 0 0 1 1 1 0 0 1 1 1 1 1 1 2 1 1 0 1 0 Hyalinocerus flavoscutatus comb. nov. 1 0 1 1 1 0 1 1 1 0 1 0 1 1 1 1 1 1 2 0 0 0 0 0 Hyalinocerus nigrimaculatus 1 0 1 1 0 0 0 0 1 0 0 0 1 1 1 1 1 0 - - 0 0 1 1 Idioscopus nagpurensis 1 1 0 0 0 0 0 0 0 1 0 0 1 1 1 1 1 0 - - 0 0 0 0 Idioscopus nitidulus 1 1 0 0 0 0 1 1 1 1 1 0 1 0 1 1 2 0 - - 0 0 0 0 Jogocerus concavus 1 0 1 1 0 0 0 1 0 0 0 0 1 0 1 1 0 1 2 1 0 1 1 0 Amritodus atkinsoni 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Amritodus brevis 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Amritodus brevistylus 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Amritodus saeedi 1 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 2 1 0 1 0 0 0 0 Paramritodus introflexus sp. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 1 0 0 0 1 0 Paramritodus flavocapitatus comb. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Paramritodus pistacious comb. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Paramritodus podocarpus comb. nov. 1 1 0 0 0 0 ? ? 0 0 0 0 0 1 1 1 ? 1 0 0 0 0 1 0 Paramritodus spatiosus sp. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Paramritodus triangulus sp. nov. 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 1 0 Species 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 Chunra quadrispinosa 0 1 0 0 0 0 0 0 0 1 2 0 0 0 0 0 0 0 - - - 0 0 Busonia albilateralis 0 1 0 0 0 0 0 0 1 - - - - 0 0 0 0 1 - - - 1 - Anidiocerus brevispinus 0 0 2 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 - - - 1 - Busoniomimus polydoros 0 1 3 0 0 0 0 1 1 - - - - 0 0 0 1 0 - - - 0 0 Hyalinocerus flavoscutatus comb. nov. 1 1 2 1 1 0 0 0 1 - - - - 0 0 0 1 0 - - - 1 - Hyalinocerus nigrimaculatus 1 1 3 0 1 0 0 0 0 2 0 0 1 0 0 0 0 0 - - - 1 - Idioscopus nagpurensis 0 0 2 0 0 0 0 0 0 1 1 1 1 0 0 0 1 0 - - - 0 0 Idioscopus nitidulus 0 0 2 0 0 0 0 0 0 1 0 1 1 0 0 0 1 0 - - - 0 0 Jogocerus concavus 0 1 2 0 0 0 1 0 0 0 1 0 1 0 0 0 1 0 - - - 0 0 Amritodus atkinsoni 0 0 1 0 1 1 0 0 1 - - - - 1 0 0 1 1 1 1 0 0 0 Amritodus brevis 0 2 1 0 1 0 0 0 1 - - - - 1 0 1 1 1 1 1 0 0 0 Amritodus brevistylus 0 0 1 0 1 0 0 0 0 0 2 0 0 1 0 0 1 1 1 1 0 0 0 Amritodus saeedi 0 0 1 0 1 0 0 0 1 - - - - 1 0 0 1 1 1 1 0 0 0 Paramritodus introflexus sp. nov. 0 1 3 0 0 0 0 0 0 0 2 0 0 1 1 0 2 1 0 1 1 0 1 Paramritodus flavocapitatus comb. nov. 0 1 3 0 0 0 0 0 1 - - - - 1 1 0 2 1 0 0 0 0 1 Paramritodus pistacious comb. nov. 0 1 3 0 0 0 0 0 1 - - - - 1 1 0 2 1 0 0 1 0 1 Paramritodus podocarpus comb. nov. 0 1 3 0 0 0 0 0 1 - - - - 1 1 0 2 1 0 1 0 1 - Paramritodus spatiosus sp. nov. 0 1 3 0 0 0 0 0 0 0 2 1 0 1 1 0 1 1 0 1 1 0 0 Paramritodus triangulus sp. nov. 0 1 3 0 0 0 0 0 0 0 2 0 0 1 1 0 2 1 0 1 1 0 1 Open in new tab Characters Head: 1. Crown texture: (0) shagreen; (1) rugose. 2. Coronal suture: (0) obsolete; (1) well developed. 3. Pronotum texture: (0) shagreen; (1) rugose. 4. Anteclypeus surpassing apex of gena: (0) no; (1) yes. 5. Anteclypeus elevated: (0) no; (1) yes. 6. Lateral frontal suture: (0) present; (1) absent. 7. Apex of lateral frontal suture, shape: (0) straight; (1) curved laterad. 8. Lateral frontal sutures extended to ocelli: (0) no; (1) yes. 9. Base of gena margin, shape: (0) straight or convex; (1) concave. 10. Apex of rostrum, shape: (0) tapered or parallel-sided; (1) broadened. 11. Colour of anteclypeus: (0) not black; (1) black. Forewing and hindleg: 12. Crossvein r-m1: (0) present; (1) absent. 13. Crossvein m-cu1: (0) absent; (1) present. 14. Number of subapical cells: (0) three; (1) two. 15. Number of apical cells bordering appendix: (0) three; (1) two. 16. Hind femur macrosetal formula: (0) 2 + 0; (1) 2 + 1. 17. Number of first hind tarsomere platellar setae: (0) two; (1) three; (2) four. Male genitalia: 18. Inner process of pygofer: (0) absent; (1) present. 19. Position of pygofer inner process: (0) arising from base on ventral margin; (1) arising from base on ventral margin and connecting with lateral surface; (2) arising from middle on ventral margin. 20. Apical half of pygofer inner process obviously curved dorsad: (0) no; (1) yes. 21. Basal lobe on pygofer ventral margin: (0) absent; (1) present. 22. Pygofer ventral margin concave: (0) no; (1) yes. 23. Length of pygofer distal lobe: (0) short; (1) elongate, surpassing anal tube. 24. Pygofer dorsal apodeme: (0) present, developed; (1) absent or undeveloped. 25. Extension of pygofer dorsal apodeme: (0) not extended to midline of pygofer side; (1) extended to midline of pygofer side. 26. Length of subgenital plate: (0) longer than pygofer; (1) between 0.5 and 1× as long as pygofer; (2) less than half as long as pygofer. 27. Setae on style dorsal margin: (0) absent; (1) dense and short; (2) fine, sparse; (3) stout and long. 28. Shape of style apical half: (0) not slender and tapered; (1) slender and tapered. 29. Style curved dorsally at nearly 90° angle: (0) no; (1) yes. 30. Apex of style curved dorsad: (0) no; (1) yes. 31. Texture of style ventral margin: (0) smooth; (1) serrate. 32. Aedeagal shaft pustulate: (0) no; (1) yes. 33. Aedeagal process: (0) present; (1) absent. 34. Number of aedeagal processes: (0) one pair; (1) two pairs; (2) more than two pairs. 35. Position of aedeagal process: (0) apical; (1) subapical; (2) basal. 36. Length of aedeagal process: (0) shorter than half-length of shaft; (1) longer than half-length of shaft. 37. Direction of aedeagal process: (0) directed distad; (1) directed basad. 38. Aedeagus S-shaped in lateral view: (0) no; (1) yes. 39. Apical half of aedeagal shaft in ventral view, shape: (0) parallel-sided or broadened; (1) slender and tapered. 40. Triangular lobe on aedeagus dorsal margin near apex: (0) absent; (1) present. 41. Position of gonopore: (0) apical; (1) subapical; (2) near middle of shaft. 42. Preatrium: (0) absent or undeveloped; (1) present, developed. 43. Length of preatrium: (0) shorter than shaft; (1) as long or longer than shaft. 44. Shape of preatrium in lateral view: (0) straight; (1) curved. 45. Preatrium much broader than shaft in lateral view: (0) no; (1) yes. 46. Presence of aedeagal dorsal apodeme: (0) present, developed; (1) absent or undeveloped. 47. Shape of aedeagal dorsal apodeme in lateral view: (0) not broadened; (1) broadened. Phylogenetic analysis Combined Bayesian inference (BI) analysis (16S, 28S D2, COI and morphological data) and molecular-only BI analysis were carried out in MrBayes v.3.2.6 (analyses on CIPRES Science Gateway; Miller et al., 2010). To determine the best-fitting nucleotide model for each gene, we used Partitionfinder v.2 (Lanfear et al., 2016) implemented in CIPRES. The GTR+G model was specified for 16S and 28S D2, and TVM+I+G for COI. The morphology dataset was run under the standard discrete model. Six chains were included in two runs of 10 million generations, sampled every 1000 generations, with a burn-in of 0.25. The average standard deviation of split frequencies was < 0.01, suggesting that runs converged. After the first 25% of trees were discarded as burn-in, posterior probability (PP) values were calculated for the majority-rule consensus tree. Combined maximum likelihood (ML) analysis (16S, 28S D2, COI and morphological data) and molecular-only ML analysis were carried out in IQ-TREE v.1.6.5. For the ML analysis, IQ-TREE selected the model for each gene using the Bayesian information criterion (BIC). The GTR+F+G4 model was specified for 16S, TIM3+F+R2 for 28S D2, TVM+F+I+G4 for COI, and the MK model for morphology. IQ-TREE executed the following tests to determine node support for the ML analysis: 5000 replicates for ultrafast bootstrap (UFB) approximation and 1000 replicates for the Shimodaira–Hasegawa approximate likelihood ratio test (SH-aLRT). The morphological data were analysed under parsimony carried out in TNT v.1.5 (Goloboff et al., 2003) using the traditional search approach, with 100 replicates followed by tree bisection and reconnection (TBR) branch swapping, and 100 trees saved per replication. Bremer support was calculated using TNT and obtained by TBR swapping on the most parsimonious trees. Character optimization and mapping were conducted with WinClada v.1.00.08 (Nixon, 2002). RESULTS The topologies resulting from combined molecular and morphological data are partly incongruent, but both trees recover Amritodus as non-monophyletic. Most nodes of the BI tree (Fig. 1) and the ML tree (Supporting Information, Fig. S1) are strongly supported by PP, SH-aLRT and UFB scores. The eight species of Amritodus are placed in three independent lineages. Amritodus flavoscutatus is recovered as more closely related to H. nigrimaculatus than to species of Amritodus with high support (PP = 1, SH-aLRT = 96.4, UFB = 96). Amritodus flavocapitatus, A. pistacious and A. podocarpus group together in a well-supported clade (PP = 1, SH-aLRT = 99.6, UFB = 100) with the three undescribed species (described below as Paramritodus introflexus, Paramritodus spatiosus and Paramritodus triangulus). The other four species [Amritodus atkinsoni (Lethierry, 1889), A. brevis, A. brevistylus and A. saeedi] of Amritodus group together in a separate clade with high support (PP = 1, SH-aLRT = 99.6, UFB = 99). The relationships among species within clades are poorly resolved. Figure 1. Open in new tabDownload slide Bayesian consensus tree based on analysis of three genes and 47 morphological characters for Amritodus and related idiocerines. Numbers below branches are the Bayesian posterior probabilities. Figure 1. Open in new tabDownload slide Bayesian consensus tree based on analysis of three genes and 47 morphological characters for Amritodus and related idiocerines. Numbers below branches are the Bayesian posterior probabilities. The BI (Supporting Information, Fig. S2) and ML (Supporting Information, Fig. S3) topologies based on molecular data only are congruent with those obtained from the combined data, but relationships among clades remain unresolved. The analysis of morphological characters carried out in TNT v.1.5 using the traditional search finds two equally most parsimonious trees (length = 99, consistency index = 55, retention index = 72). The resulting strict consensus tree (length = 100, consistency index = 55, retention index = 71) and the Bremer support are shown in Figure 2. In this analysis, Amritodus is also recovered as non-monophyletic. The clade A. flavoscutatus + H. nigrimaculatus is supported by a Bremer value of three and by three synapomorphic characters: pygofer dorsal apodeme extended to midline of pygofer side (25: 1), style curved dorsally at nearly 90° angle (29: 1) and aedeagal dorsal apodeme undeveloped (46: 1). The clade of three Amritodus species (A. flavocapitatus, A. pistacious and A. podocarpus) plus the three new species is supported by a Bremer value of two and by two synapomorphic characters: pygofer distal lobe surpassing anal tube (23: 1) and apical half of aedeagal shaft slender and tapered in ventral view (39: 1). The clade {A. brevistylus + [A. atkinsoni + (A. brevis + A. saeedi)]} is supported by a Bremer value of three and by the following synapomorphic characters: forewing without cross-vein r-m1 (12: 1), first hind tarsomere with four platellar setae (17: 2) and style curved dorsally at nearly 90° angle (29: 1). Figure 2. Open in new tabDownload slide Strict consensus of the two equally most parsimonious trees found by TNT. Numbers above the circles refer to characters and those below refer to character states. Black circles represent non-homoplasious changes; white circles represent homoplasious changes. Red numbers below the branches are Bremer support values. Figure 2. Open in new tabDownload slide Strict consensus of the two equally most parsimonious trees found by TNT. Numbers above the circles refer to characters and those below refer to character states. Black circles represent non-homoplasious changes; white circles represent homoplasious changes. Red numbers below the branches are Bremer support values. DISCUSSION Our phylogenetic analysis confirms that Amritodus, as previously defined, is not monophyletic. This supports the observations of Viraktamath (1997), who suggested removing A. pistacious from Amritodus based on differences from the type species in forewing venation, male genitalia, host plant and distribution. Our phylogenetic results also indicate that A. flavoscutatus was misclassified to genus and support the transfer of this species to Hyalinocerus. Morphological evidence supporting this transfer includes the fine transversely rugose crown and pronotum, anteclypeus surpassing the apex of the gena, forewing with r-m1 and m-cu1 cross-veins, pygofer dorsal apodeme extended to the midline of the pygofer side, style curved dorsally at a nearly 90° angle, undeveloped aedeagal dorsal apodeme and short preatrium. Our analyses also consistently placed the remaining previously described Amritodus species in two separate clades. One clade, including three previously described species, Amritodus flavocapitatus, A. pistacious and A. podocarpus, is supported by the following morphological features: crown finely rugose, pronotum shagreen, forewing with r-m1 crossvein, without m-cu1 vein, hind basitarsus with three platellae, pygofer distal lobe surpassing anal tube, style with stout and long setae on dorsal margin, apical half of aedeagal shaft slender and tapered in ventral view, and aedeagal preatrium shorter than shaft. This clade is formally recognized below as the new genus Paramritodus. The remaining Amritodus species, including the type species of the genus, were placed in the other clade, also well supported by morphological and molecular characters. The current classification of this group is revised to conform to these new phylogenetic results (see below). TAXONOMY Genus Amritodus Anufriev, 1970 AmritodusAnufriev, 1970: 376. Type species: Idiocerus atkinsoni Lethierry, 1889; by original designation. Diagnosis: Body generally yellowish brown or brown. Crown with a pair of black spots on either side of midline. Face yellowish or brownish. Pronotum with a pair of black markings on anterior margin. Head wider than pronotum. Crown finely rugose. Head wider than long, ocelli closer to eyes than to each other; lora broad; anteclypeus apex wider than base; rostrum broadened apically. Combined length of mesoscutum and scutellum longer than pronotum, shagreen. Forewing with m-cu1 cross-vein, without r-m1 and m-cu2 cross-veins. Hind femur with 2 + 1 apical setae. Hind tibiae with 18–20 setae on row PD, six setae on AD and six to eight setae on row AV. Hind basitarsus with four platellae. Male pygofer longer than height, with a pair of inner processes on the ventral margin; apical half of inner processes obviously curved dorsad. Subgenital plate with hair-like setae on dorsal and ventral margin. Style curved dorsally, with dense and short setae on dorsal margin. Aedeagus S-shaped, preatrium well developed, as long as or longer than shaft; dorsal apodeme developed; gonopore subapical on ventral margin. Female seventh sternite wider than long, with caudal margin obviously concave. Ovipositor extending beyond pygofer. First valvulae apical one-third to one-quarter curved dorsad, apex attenuated, with sculpture strigate. Second valvulae curved dorsally; dorsal margin with more than ten teeth. Distribution: Bangladesh, China, India, Myanmar, Pakistan, Sri Lanka and Thailand. Remarks: Amritodus is similar to Idioscopus, but differs from the latter in having an elongated aedeagal preatrium, and the aedeagal shaft short and S-shaped without a long process near the apex. Key to species of Amritodus (males) (modified from Viraktamath, 1997) 1. Subgenital plate shorter than half length of pygofer (Fig. 13A) A. brevis Subgenital plate longer than pygofer (Fig. 12A) 2 2. Aedeagus with basal pair of short processes on ventral surface (Fig. 14E) A. brevistylus Aedeagus without process (Fig. 12E) 3 3. Style apex strongly curved dorsad (Fig. 12G) A. atkinsoni Style apex not strongly curved dorsad (Viraktamath, 1997: fig. 9) A. saeedi Amritodus atkinsoni (Lethierry, 1889) (Figs 3, 12) Idiocerus atkinsoni Lethierry, 1889: 252. Amritodus atkinsoni (Lethierry) – Anufriev, 1970: 376, figs 1–7. – Viraktamath, 1997: 113, figs 1–6, 28. Diagnosis: Yellowish brown; pronotum, mesoscutum and scutellum with median brown stripe (Fig. 3A). Subgenital plate with dense, fine and long setae (Fig. 12A). Style apex curved dorsad, with dense, short and stout setae on dorsal surface (Fig. 12G). Aedeagal shaft smooth; preatrium much longer than shaft; dorsal apodeme oval in ventral view (Fig. 12E–F). Figure 3. Open in new tabDownload slide Amritodus atkinsoni. A, habitus of male, dorsal view. B, habitus of male, lateral view. C, head and thorax of male, dorsal view. D, face of male. E, head and thorax of female, dorsal view. F, face of female. G, habitus of female, dorsal view. H, habitus of female, lateral view. I, female abdomen, ventral view. J, first valvula. K, apex of second valvula. L, second valvula. Scale bars: 1.0 mm (A, B, G–I); 0.5 mm (C–F, J, L); 0.2 mm (K). Figure 3. Open in new tabDownload slide Amritodus atkinsoni. A, habitus of male, dorsal view. B, habitus of male, lateral view. C, head and thorax of male, dorsal view. D, face of male. E, head and thorax of female, dorsal view. F, face of female. G, habitus of female, dorsal view. H, habitus of female, lateral view. I, female abdomen, ventral view. J, first valvula. K, apex of second valvula. L, second valvula. Scale bars: 1.0 mm (A, B, G–I); 0.5 mm (C–F, J, L); 0.2 mm (K). Materials examined: Ten ♂ and ten ♀, India, Bhunga, 25 April 1967, coll. A. S. Sohi (USNM). Distribution: Bangladesh, India, Myanmar and Pakistan. Remarks: This species is similar to A. brevistylus in coloration and male genitalia, but can be distinguished from the latter by the aedeagus without process. Amritodus brevis Viraktamath, 1997 (Figs 4, 13) Amritodus brevisViraktamath, 1997: 115, figs 32–41. Diagnosis: Body robust and large. Rostrum much broadened on apical half (Fig. 4D). Subgenital plate shorter than half of pygofer, extended ventrally, without hair-like setae (Fig. 13A). Style with dense, short and fine setae on dorsal surface (Fig. 13G). Aedeagal shaft with a pair of short triangular processes subapically; preatrium slightly longer than shaft (Fig. 13E, F). Figure 4. Open in new tabDownload slide Amritodus brevis. A, habitus, dorsal view. B, habitus, lateral view. C, head and thorax, dorsal view. D, face. Scale bars: 1.0 mm. Figure 4. Open in new tabDownload slide Amritodus brevis. A, habitus, dorsal view. B, habitus, lateral view. C, head and thorax, dorsal view. D, face. Scale bars: 1.0 mm. Materials examined: One ♂, China, Yunnan Province, Daluo, 679 m, 23 May 2011, coll. Lin Lu (NWAFU); one ♂, Thailand, Phetchabun Khao Kho, NP, mixed deciduous forest near office, 16°39.55′N, 101°8.123′E, 230 m, Pan trap, 7–8 February 2007, coll. Somchai Chachumnan & Saink (QSBG). Distribution: India and new records for China and Thailand. Remarks: This species can be distinguished from the other species of Amritodus by the large body, short subgenital plate, and aedeagus with a pair of short triangular processes subapically. Amritodus brevistylus Viraktamath, 1976 (Figs 5, 14) Amritodus brevistylus Viraktamath, 1976: 234, figs 1–5. – Viraktamath, 1997: 115, figs 15–21, 27. Diagnosis: Rostrum broadened on apical half (Fig. 5D). Subgenital plate with dense, long and fine setae on dorsal margin and a few setae on ventral margin (Fig. 14A). Style apex and dorsal margin with dense, short and fine setae, serrate medially on ventral margin (Fig. 14G). Aedeagus with basal pair of short processes on ventral surface; preatrium much longer than shaft; dorsal apodeme fan shaped in ventral view (Fig. 14E, F). Figure 5. Open in new tabDownload slide Amritodus brevistylus. A, habitus of male, dorsal view. B, habitus of male, lateral view. C, head and thorax of male, dorsal view. D, face of male. E, head and thorax of female, dorsal view. F, face of female. G, habitus of female, dorsal view. H, habitus of female, lateral view. I, forewing. J, sternite VII of female, ventral view. K, first valvula. L, second valvula. M, apex of first valvula. N, apex of second valvula. Scale bars: 1.0 mm (A, B, G–J); 0.5 mm (C–F, K, L); 0.2 mm (M, N). Figure 5. Open in new tabDownload slide Amritodus brevistylus. A, habitus of male, dorsal view. B, habitus of male, lateral view. C, head and thorax of male, dorsal view. D, face of male. E, head and thorax of female, dorsal view. F, face of female. G, habitus of female, dorsal view. H, habitus of female, lateral view. I, forewing. J, sternite VII of female, ventral view. K, first valvula. L, second valvula. M, apex of first valvula. N, apex of second valvula. Scale bars: 1.0 mm (A, B, G–J); 0.5 mm (C–F, K, L); 0.2 mm (M, N). Materials examined: Five ♂ and five ♀, Sri Lanka, Kan., Dist., Kandy 1800 feet, Peak View Motel, 15–24 January 1970, coll. Davis & Rowe (USNM). Distribution: India and Sri Lanka. Remarks: This species is an economically problematic pest, feeding on mango and other fruit crops. It is close to A. atkinsoni, having the same colour and similar male genitalia; see remarks under A. atkinsoni for differences. Amritodus saeedi Ahmed, Naheed & Ahmed, 1980 Amritodus saeedi Ahmed, Naheed & Ahmed, 1980: 221, fig. 1. – Viraktamath, 1997: 114, figs 7–13, 22–26. Diagnosis: Pronotum, mesoscutum and scutellum without median brown stripe. Style apex not strongly curved dorsad. Aedeagus simple, without process; preatrium slightly longer than shaft; dorsal apodeme broadened in ventral view. Materials examined: One ♂, India, Karnataka, Jog Falls, 534 m, 18 November 1976, coll. C. A. Viraktamath (UASB); one ♂, India, Mangalore, 16 May 1986, coll. C. A. Viraktamath (UASB). Distribution: India and Pakistan. Remarks: This species is similar to A. atkinsoni in the aedeagus, but can be distinguished from the latter by the pronotum without a median stripe and style apex not strongly curved dorsad. Genus Paramritodus Xue & Zhang gen. nov. Type species (here designated): Paramritodus triangulus Xue & Zhang, sp. nov. LSID:http://zoobank.org/urn:lsid:zoobank.org:act:381ABAC6-4DA9-40F3-9FF6-18AAF8368C1A Diagnosis: Body small. Crown finely rugose. Head wider than long; ocelli closer to eyes than to each other; lora broad; rostrum slight or not broadened apically. Pronotum shagreen, shorter than combined length of mesoscutum and scutellum. Forewing with r-m1 cross-vein, without m-cu1 cross-vein. Hind femur with 2 + 1 apical setae. Hind tibiae with 14–18 setae on row PD, five or six setae on AD and five to seven setae on row AV. Hind basitarsus with three platellae. Male pygofer elongate, distal lobe surpassing anal tube, with pair of inner processes; apex of inner processes often hooked. Subgenital plate shorter than pygofer, with hair-like setae on dorsal and ventral margin. Connective short. Style sickle shaped, with stout and long setae in single row on dorsal margin. Aedeagus S-shaped, without process or with pair of basal processes on ventral surface; aedeagal shaft slender and tapered in ventral view; preatrium shorter than shaft; gonopore subapical or near middle of shaft. Female seventh sternite wider than long, caudal margin produced, concave in middle. First valvulae slightly curved, with sculpture strigate. Second valvulae with several irregular dorsal teeth near apex. Etymology: This masculine generic name refers to the similarity of this new genus to Amritodus. Distribution: China (Guizhou, Shanxi, Taiwan, Yunnan and Zhejiang). Remarks: This new genus is closely related to Amritodus based on its similar body size and coloration, but can be distinguished by the forewing with cross-vein r-m1 and without cross-vein m-cu1, pygofer distal lobe surpassing the anal tube, style with long setae, and aedeagal preatrium shorter than the shaft. Key to species of Paramritodus (males) 1. Aedeagal shaft with basal pair of processes on ventral surface (Fig. 16E) 2 Aedeagus without process (Fig. 15E) 4 2. Aedeagal process longer than half length of shaft; gonopore subapical (Fig. 18E) P. spatiosus Aedeagal process shorter than half length of shaft; gonopore near middle of shaft (Fig. 16E) 3 3. Aedeagal shaft distad of process, concave ventrally in lateral view (Fig. 16E) P. introflexus Aedeagal shaft distad of process, not concave ventrally in lateral view (Fig. 19E) P. triangulus 4. Pronotum with two triangular black markings on anterior margin (Zhang & Li, 2010: fig. 1); aedeagal preatrium ventral margin curved in lateral view (Zhang & Li, 2010: fig. 8) P. podocarpus Pronotum without black markings on anterior margin (Fig. 6A); aedeagal preatrium ventral margin straight in lateral view (Fig. 15E) 5 5. Pygofer caudal margin rounded; segment X with long slender caudal process (Fig. 15A) P. flavocapitatus Pygofer caudal margin angular; segment X with short wide caudal process (Fig. 17A) P. pistacious Paramritodus flavocapitatus (Cai & He, 2001) comb. nov. (Figs 6, 15) Amritodus flavocapitatus Cai & He – Cai et al., 2001: 199, figs 54–61. Diagnosis: Crown and pronotum mainly brownish. Face yellowish. Mesoscutum brown, basal triangles black; scutellum brown (Fig. 6C). Face with short seta adjacent to corresponding eye; rostrum apex slightly broad, not reaching hind coxae. Figure 6. Open in new tabDownload slide Paramritodus flavocapitatus. A, habitus of male, dorsal view. B, habitus of male, lateral view. C, head and thorax of male, dorsal view. D, face of male. E, head and thorax of female, dorsal view. F, face of female. G, habitus of female, dorsal view. H, habitus of female, lateral view. I, forewing. J, female abdomen, ventral view. K, first valvula. L, second valvula. M, apex of first valvula. N, apex of second valvula. Scale bars: 1.0 mm (A, B, G, H–J); 0.5 mm (C–F, K, L); 0.2 mm (M, N). Figure 6. Open in new tabDownload slide Paramritodus flavocapitatus. A, habitus of male, dorsal view. B, habitus of male, lateral view. C, head and thorax of male, dorsal view. D, face of male. E, head and thorax of female, dorsal view. F, face of female. G, habitus of female, dorsal view. H, habitus of female, lateral view. I, forewing. J, female abdomen, ventral view. K, first valvula. L, second valvula. M, apex of first valvula. N, apex of second valvula. Scale bars: 1.0 mm (A, B, G, H–J); 0.5 mm (C–F, K, L); 0.2 mm (M, N). Materials examined: One ♂, China, Zhejiang Province, Mt. Tianmu, at light, 26 July 2011, coll. Lin Lu (NWAFU); one ♂, China, Shanxi Province, Hengqu, Lishan, 19 July 2006, coll. Zhaofu Yang (NWAFU). Distribution: China (Shanxi and Zhejiang). Remarks: This species can be distinguished from the other species of Paramritodus by the following features: crown without black spot; pygofer elongate, caudal margin rounded; pygofer inner processes curved apically, hook-like; style flat, broad subapically; aedeagal shaft basal part straight. Paramritodus introflexus Xue & Zhang sp. nov. (Figs 7, 16) LSID:http://zoobank.org/urn:lsid:zoobank.org:act:5A2F9CE8-334D-40AC-97B2-49F8BB9891EA Diagnosis: Crown with pair of black spots. Pronotum with pair of triangular black markings on anterior margin. Rostrum slightly broadened apically. Pygofer ventral margin with inner process elongate. Aedeagus with a pair of basal processes on ventral surface. Aedeagal shaft concave medially. Description: Length (including wings): male 3.3–3.4 mm. Crown yellowish, with black spot on either side of midline close to eyes; frontoclypeus and anteclypeus brownish; gena and lora yellowish (Fig. 7D). Pronotum greenish, with black markings; anterior margin with dark brown spot on either side of midline. Mesoscutum yellowish, basal triangles small, black; scutellum yellowish (Fig. 7C). Forewing mainly green, apex infuscate, base of costal margin green (Fig. 7E). Face with short seta adjacent to corresponding eye; rostrum apex slightly broad, rhombus shaped, not reaching hind coxae. Figure 7. Open in new tabDownload slide Paramritodus introflexus. A, habitus, dorsal view. B, habitus, lateral view. C, head and thorax, dorsal view. D, face. E, forewing. Scale bars: 1.0 mm (A, B, E); 0.5 mm (C, D). Figure 7. Open in new tabDownload slide Paramritodus introflexus. A, habitus, dorsal view. B, habitus, lateral view. C, head and thorax, dorsal view. D, face. E, forewing. Scale bars: 1.0 mm (A, B, E); 0.5 mm (C, D). Male abdomen with tergal apodemes truncate, slightly less than two segments long, separated mesally by V-shaped notch; sternal apodemes small, less than one segment long (Fig. 16B, C). Male genitalia: Pygofer elongate, obviously surpassing anal tube; inner processes elongate, almost reaching caudal margin, apex curved and pointed (Fig. 16A). Style curved dorsally, with row of long setae on dorsal margin (Fig. 16G). Connective short. Aedeagal shaft much curved, tapering apically in lateral view, concave basally, with basal pair of slender processes on ventral surface, dorsal apodeme and preatrium developed (Fig. 16E, F). Materials examined: Holotype: ♂, China, Yunnan Province, Baoshan, 12 May 2012, coll. Yanghui Cao (NWAFU). Paratypes: three ♂, same data as holotype (NWAFU). Etymology: The specific name is a Latin adjective, which refers to the strongly concave ventral margin of the aedeagal shaft. Remarks: This species is similar to P. triangulus in colour and shape, but can be distinguished from the latter by the elongate pygofer inner process almost reaching the caudal margin and aedeagal shaft strongly concave medially. Paramritodus pistacious (Huang & Maldonado-Capriles, 1992) comb. nov. (Figs 8, 17) Amritodus pistaciousHuang & Maldonado-Capriles, 1992: 2, fig. 1. Diagnosis: Crown and face yellowish. Face with short seta adjacent to corresponding eye; rostrum parallel-sided apically, not reaching hind coxae (Fig. 8D). Pygofer caudal margin angular; segment X with short, wide caudal process (Fig. 17A). Figure 8. Open in new tabDownload slide Paramritodus pistacious. A, habitus of male, dorsal view. B, habitus of male, lateral view. C, head and thorax of male, dorsal view. D, face of female. E, head and thorax of female, dorsal view. F, face of female. G, habitus of female, dorsal view. H, habitus of female, lateral view. I, female abdomen, ventral view. J, first valvula. K, second valvula. L, apex of first valvula. M, apex of second valvula. Scale bars: 1.0 mm (A, B, G–I); 0.5 mm (C–F, J, K); 0.2 mm (L, M). Figure 8. Open in new tabDownload slide Paramritodus pistacious. A, habitus of male, dorsal view. B, habitus of male, lateral view. C, head and thorax of male, dorsal view. D, face of female. E, head and thorax of female, dorsal view. F, face of female. G, habitus of female, dorsal view. H, habitus of female, lateral view. I, female abdomen, ventral view. J, first valvula. K, second valvula. L, apex of first valvula. M, apex of second valvula. Scale bars: 1.0 mm (A, B, G–I); 0.5 mm (C–F, J, K); 0.2 mm (L, M). Materials examined: Four ♂ and three ♀ (paratypes), China, Taiwan, Taichung, 26 July 1987, coll. C. T. Yang (USNM); one ♂, China, Taiwan, Duona, 3 June 2011, coll. Xiaolei Huang (IOZ). Distribution: China (Taiwan). Remarks: This species can be distinguished from the other species of Paramritodus by the following features: pygofer triangular, extended well beyond apex of anal tube, apex not rounded; pygofer inner process curved apically, reaching caudal margin; style slightly curved. Paramritodus podocarpus (Zhang & Li, 2010) comb. nov. Amritodus podocarpusZhang & Li, 2010: 730, figs 1–9. Diagnosis: Crown yellow, without black spot. Pronotum with a pair of triangular black markings on anterior margin. Rostrum slightly broadened apically. Pygofer inner process not reaching caudal margin. Style subapical extended. Aedeagal dorsal apodeme undeveloped. Aedeagal preatrium arcuate in lateral view. Distribution: China (Guizhou). Remarks: Specimens of this species were not available for study. This species can be distinguished by the following features: crown without a pair of black spots on either side of midline; style medially concave and extended subapically; aedeagal shaft slender and elongate, and preatrium arcuate in lateral view. Paramritodus spatiosus Xue & Zhang sp. nov. (Figs 9, 18) LSID:http://zoobank.org/urn:lsid:zoobank.org:act:2F91797D-5C95-49AE-A39A-58D29B5AB205 Diagnosis: Crown with a pair of black spots. Pronotum with a pair of triangular black markings on anterior margin. Rostrum parallel-sided apically. Aedeagal process longer than half length of shaft. Aedeagal dorsal apodeme undeveloped. Aedeagal preatrium broadened basally. Gonopore subapical. Description: Length (including wings): male 3.4 mm. Crown with black spot on either side of midline (Fig. 9C). Pronotum with pair of black triangular markings on anterior margin (Fig. 9C). Face without seta adjacent to corresponding eye; rostrum short, not reaching hind coxae, parallel-sided apically. Figure 9. Open in new tabDownload slide Paramritodus spatiosus. A, habitus, dorsal view. B, habitus, lateral view. C, head and thorax, dorsal view. D, face. Scale bars: 1.0 mm (A, B); 0.5 mm (C, D). Figure 9. Open in new tabDownload slide Paramritodus spatiosus. A, habitus, dorsal view. B, habitus, lateral view. C, head and thorax, dorsal view. D, face. Scale bars: 1.0 mm (A, B); 0.5 mm (C, D). Male abdomen with large tergal apodemes, length as long as two segments, separated mesally by a V-shaped notch; sternal apodemes small, less than one segment long (Fig. 18B, C). Male genitalia: Pygofer with rounded caudal margin; inner process arising from base of ventral margin, apex curved mesad. Subgenital plate with dense, long and fine setae on dorsal margin, few fine setae on ventral margin (Fig. 18A). Aedeagus with a pair of basal processes on ventral surface, length longer than half of shaft; preatrium broadened basally in ventral and lateral view, slender medially; dorsal apodeme undeveloped (Fig. 18E, F). Material examined: Holotype: ♂, China, Yunnan Province, Dali, Yangbi, Yangjiang, 17 June 2013, coll. Qingquan Xue (NWAFU). Etymology: The specific epithet, a Latin adjective, refers to the elongate aedeagal process. Remarks: The new species seems to have close affinities with P. introflexus and P. triangulus (see below) based on the similar body colour, but it can be distinguished from these two species by the aedeagal process longer than half the length of the shaft and undeveloped dorsal apodeme. Paramritodus triangulus Xue & Zhang sp. nov. (Figs 10, 19) LSID:http://zoobank.org/urn:lsid:zoobank.org:act:8CD6C406-C95A-4CDE-8F0E-CB4C257F8469 Diagnosis: Crown with a pair of black spots. Pronotum with a pair of triangular black markings on anterior margin. Rostrum parallel-sided apically. Aedeagal process shorter than half length of shaft. Aedeagal shaft not concave medially. Description: Length (including wings): male 3.4–3.5 mm, female 3.4 mm. Crown stramineous, with black spot on either side of midline; frontoclypeus and anteclypeus and upper part of face brownish with black stripe; gena and lora yellowish (Fig. 10D). Pronotum greenish, with black markings, anterior margin with dark brown spot on either side of midline. Mesoscutum yellowish, basal triangles black; scutellum yellowish (Fig. 10C). Forewing infuscate, claval area and base of costal margin and parts of venation greenish. Face with short seta adjacent to corresponding eye; rostrum parallel-sided apically, not reaching hind coxae. Figure 10. Open in new tabDownload slide Paramritodus triangulus. A, habitus of male, dorsal view. B, habitus of male, lateral view. C, head and thorax of male, dorsal view. D, face of male. E, head and thorax of female, dorsal view. F, face of female. G, habitus of female, dorsal view. H, habitus of female, lateral view. I, sternite VII of female, ventral view. J, first valvula. K, apex of first valvula. L, second valvula. M, apex of second valvula. Scale bars: 1.0 mm (A, B, G, H); 0.5 mm (C–F, I, J, L); 0.2 mm (K, M). Figure 10. Open in new tabDownload slide Paramritodus triangulus. A, habitus of male, dorsal view. B, habitus of male, lateral view. C, head and thorax of male, dorsal view. D, face of male. E, head and thorax of female, dorsal view. F, face of female. G, habitus of female, dorsal view. H, habitus of female, lateral view. I, sternite VII of female, ventral view. J, first valvula. K, apex of first valvula. L, second valvula. M, apex of second valvula. Scale bars: 1.0 mm (A, B, G, H); 0.5 mm (C–F, I, J, L); 0.2 mm (K, M). Male abdomen with tergal apodemes truncate, less than two segments long, separated mesally by a U-shaped notch; sternal apodemes small, less than one segment long (Fig. 19B, C). Male genitalia: Pygofer triangular; pygofer inner process S-shaped, apex hook-like; subgenital plate with dense, long setae on dorsal margin and few setae on ventral margin (Fig. 19A). Style with long and stout setae on dorsal margin (Fig. 19G). Connective short. Aedeagal shaft tapering apically in lateral view, with basal pair of slender processes on ventral surface, dorsal apodeme and preatrium developed; gonopore on ventral surface medially (Fig. 19E, F). Materials examined: Holotype: ♂, China, Yunnan Province, Nansan, 29 May 2011, coll. Lin Lu (NWAFU). Paratypes: five ♂ and two ♀, same data as holotype (NWAFU). Etymology: The specific epithet, a Latin adjective, refers to the pygofer being triangular in lateral view. Remarks: This new species is close to P. introflexus, having the same colour and similar male genitalia; see remarks under P. introflexus for morphological comparison. Genus Hyalinocerus Zhang & Li, 2012 Hyalinocerus Zhang & Li, 2012: 204. Type species: Hyalinocerus nigrimaculatus Zhang & Li, 2012; by original designation. Diagnosis: Crown with a pair of large spots. Crown and pronotum finely transversely rugose; frontoclypeus and anteclypeus elevated; anteclypeus surpassing apex of gena. Forewing with r-m1 and m-cu1 cross-vein, without m-cu2 cross-vein. Hind femur with 2 + 1 apical setae. Hind tibiae with 16–25 setae on row PD, six to 14 setae on AD and seven to 14 setae on row AV. Hind basitarsus with three platellae. Pygofer dorsal apodeme present, extended to midline of pygofer lateral surface. Segment X inner process developed. Style elongate and curved dorsally at nearly a 90° angle. Aedeagal shaft tubular; preatrium short; dorsal apodeme undeveloped; gonopore situated near apex on ventral surface. Female seventh sternite shorter than wide, caudal margin obviously convex. Female first valvulae sculpture strigate, strongly curved, apex pointed. Female second valvulae curved obviously, with several dorsal teeth. Distribution: China (Guizhou, Henan, Hubei, Shaanxi and Sichuan). Remarks: This genus can be distinguished from other genera of Idiocerini by the following features: pronotum rugose; anteclypeus elevated; anteclypeus surpassing apex of gena; pygofer dorsal apodeme extended to midline of pygofer lateral surface; style distinctly curved dorsally. Hyalinocerus flavoscutatus (Cai & Shen, 1998) comb. nov. (Figs 11, 20) Amritodus flavoscutatus Cai & Shen, 1998: 33, fig. 7. Figure 11. Open in new tabDownload slide Hyalinocerus flavoscutatus. A, habitus of male, dorsal view. B, habitus of male, lateral view. C, head and thorax of male, dorsal view. D, face of male. E, head and thorax of female, dorsal view. F, face of female. G, habitus of female, dorsal view. H, habitus of female, lateral view. I, sternite VII of female, ventral view. J, first valvula. K, apex of first valvula. L, second valvula. M, apex of second valvula. N, forewing. Scale bars: 1.0 mm (A, B, G, H, N); 0.5 mm (C–F, I, J, L); 0.2 mm (K, M). Figure 11. Open in new tabDownload slide Hyalinocerus flavoscutatus. A, habitus of male, dorsal view. B, habitus of male, lateral view. C, head and thorax of male, dorsal view. D, face of male. E, head and thorax of female, dorsal view. F, face of female. G, habitus of female, dorsal view. H, habitus of female, lateral view. I, sternite VII of female, ventral view. J, first valvula. K, apex of first valvula. L, second valvula. M, apex of second valvula. N, forewing. Scale bars: 1.0 mm (A, B, G, H, N); 0.5 mm (C–F, I, J, L); 0.2 mm (K, M). Figure 12. Open in new tabDownload slide Amritodus atkinsoni. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm. Figure 12. Open in new tabDownload slide Amritodus atkinsoni. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm. Figure 13. Open in new tabDownload slide Amritodus brevis. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm. Figure 13. Open in new tabDownload slide Amritodus brevis. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm. Figure 14. Open in new tabDownload slide Amritodus brevistylus. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm. Figure 14. Open in new tabDownload slide Amritodus brevistylus. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm. Figure 15. Open in new tabDownload slide Paramritodus flavocapitatus. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm. Figure 15. Open in new tabDownload slide Paramritodus flavocapitatus. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm. Figure 16. Open in new tabDownload slide Paramritodus introflexus. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm (A–C, E–G); 0.1 mm (connective). Figure 16. Open in new tabDownload slide Paramritodus introflexus. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm (A–C, E–G); 0.1 mm (connective). Figure 17. Open in new tabDownload slide Paramritodus pistacious. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm (A–C, E–G); 0.1 mm (D). Figure 17. Open in new tabDownload slide Paramritodus pistacious. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm (A–C, E–G); 0.1 mm (D). Figure 18. Open in new tabDownload slide Paramritodus spatiosus. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm (A–C, E–G); 0.1 mm (D). Figure 18. Open in new tabDownload slide Paramritodus spatiosus. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm (A–C, E–G); 0.1 mm (D). Figure 19. Open in new tabDownload slide Paramritodus triangulus. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm (A–C, E–G); 0.1 mm (D). Figure 19. Open in new tabDownload slide Paramritodus triangulus. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm (A–C, E–G); 0.1 mm (D). Figure 20. Open in new tabDownload slide Hyalinocerus flavoscutatus. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm (A–C, E–G); 0.1 mm (D). Figure 20. Open in new tabDownload slide Hyalinocerus flavoscutatus. A, male pygofer, anal tube and subgenital plate, lateral view. B, abdominal tergal apodemes. C, abdominal sternal apodemes. D, connective, ventral view. E, aedeagus, lateral view. F, aedeagus, ventral view. G, style, lateral view. Scale bars: 0.2 mm (A–C, E–G); 0.1 mm (D). Diagnosis: Crown with a pair of black spots. Pronotum finely rugose. Anteclypeus elevated and surpassing apex of gena. Pygofer inner process reaching caudal margin of pygofer. Style tapering apically. Aedeagal shaft straight in lateral view. Aedeagal dorsal apodeme and preatrium short. Description: Body brown. Base of face with black stripe in middle; ocelli with black markings around; lorum, anteclypeus, antennal pits and apical half of frontoclypeus black; frontoclypeus with several black markings on either side (Fig. 11D). Pronotum with a pair of semicircular black markings on anterior margin, and lateral area black. Mesoscutum mostly black; scutellum yellowish (Fig. 11C). Forewing veins dark brown and brownish, with black markings on subcostal margin (Fig. 11N). Female colour similar to male, but base of face with short brown stripe in middle; apical half of lorum, lateral and anterior margin of frontoclypeus black; pronotum with brown markings (Fig. 11E,F). Male abdomen with large tergal apodemes, length shorter than one segment long, separated mesally by a V-shaped notch; sternal apodemes small, less than one segment long (Fig. 20B, C). Pygofer caudal margin rounded, laterally with transverse hyaline band; with inner process arising from middle on ventral margin, reaching caudal margin. Subgenital plate slender, shorter than pygofer, apex with a few short and fine setae on ventral margin (Fig. 20A). Connective Y-shaped. Materials examined: Two ♂ and one ♀, China, Shaanxi Province, Huoditang, 1600 m, 7 July 2010, coll. Lin Lu (NWAFU); one ♂, China, Guizhou Province, Kuankuoshui, Fenshuiling, 1180 m, 15 August 2012, coll. Yang Wang (NWAFU); one ♂, China, Hubei Province, Xuanen, Chunmu, 30 July 1989, coll. Qingyao Liu (SEMCAS). Distribution: China (Guizhou, Henan, Hubei and Shaanxi). SUPPORTING INFORMATION Additional Supporting Information may be found in the online version of this article at the publisher's web-site: Figure S1. Maximum likelihood (ML) tree estimated from the combined morphological and molecular dataset. Numbers below branches are Shimodaira–Hasegawa approximate likelihood ratio test (SH-aLRT) and ultrafast bootstrap (UFB) from maximum likelihood analysis. Figure S2. Bayesian consensus tree recovered from Bayesian analysis of molecular dataset (without morphology). Numbers below branches are Bayesian posterior probabilities (PP). Figure S3. Maximum likelihood (ML) tree estimated from molecular dataset. Numbers below branches are Shimodaira–Hasegawa approximate likelihood ratio test (SH-aLRT) and ultrafast bootstrap (UFB) from maximum likelihood analysis. [Version of record, published online 21 November 2019; http://zoobank.org/urn:lsid:zoobank.org:pub:7AF64F35-5144- 432F-81C2-D6365C17B6FF] ACKNOWLEDGEMENTS We sincerely thank Professor C. A. Viraktamath (Agricultural Sciences University, India) for checking specimens and C. H. Dietrich (Illinois Natural History Survey, University of Illinois, USA) and J. R. Schrock (Emporia State University, USA) for revising this manuscript. 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Google Preview WorldCat COPAC © 2019 The Linnean Society of London, Zoological Journal of the Linnean Society This article is published and distributed under the terms of the Oxford University Press, Standard Journals Publication Model (https://academic.oup.com/journals/pages/open_access/funder_policies/chorus/standard_publication_model)