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1 Zootaxa 4286 (4): Copyright 2017 Magnolia Press Article Dysaphis (Dysaphis) kadyrovi sp. nov. a new aphid species (Hemiptera: Aphididae) from Tajikistan ISSN (print edition) ZOOTAXA ISSN (online edition) ŁUKASZ DEPA, MARIUSZ KANTURSKI 1, ARTUR TASZAKOWSKI, MARCIN WALCZAK, AGNIESZKA BUGAJ-NAWROCKA & KARINA WIECZOREK Department of Zoology, Faculty of Biology and Environmental Protection, University of Silesia, Bankowa 9, Katowice, Poland. 1 Corresponding author. mariusz.kanturski@us.edu.pl Abstract Dysaphis (Dysaphis) kadyrovi Depa & Kanturski sp. nov., a new aphid species associated with an endemic plant species Anacantha darwasica (C. Winkl.) Soják (Asteraceae) is described and figured from specimens collected in Tajikistan. Taxonomic comments on the morphological resemblance of the new species to closely related Dysaphis (D.) pseudomolli Narzikulov is provided. In view of the finding that the new species shares identical COI sequence with that reported for the morphologically distinct Dysaphis (Dysaphis) lappae lappae (Koch), the reliability of barcoding and molecular data is discussed. Key words: Anacantha, Asteraceae, new species, biodiversity hotspot, barcoding Introduction Central Asia is currently recognized as one of world s hot spots of biological diversity (Myers et al. 2000), comprising, among others, montane grasslands and shrublands of Pamir Alai Mountains. This area still suffers from insufficient recognition of the full richness and diversity of its flora and fauna, which is characterized by high level of endemism (Kadyrov et al. 2016). Within this region, the area of Tajikistan is one of the centers for speciation and plays the vital role in global biodiversity conservation. The geographical location of the country in the middle of Eurasia, at the junction of many floristic and faunal provinces of the northern hemisphere, among the huge mountain systems of Central Eurasia, vast deserts, and complex landscapes, favors concentration of the richest biological diversity, with numerous endemic species, relict communities, ecosystems and genetic resources (Safarov 2003). Taking this into account, the aphid fauna of this area seems to be insufficiently studied, with major review of species made by Narzikulov and co workers (Narzikulov 1963; Narzikulov & Umarov 1969; Narzikulov & Daniyarova 1990). Since that time, there seems to be no detailed studies of this interesting area, except some works of Kadyrbekov, which concern the aphid fauna of neighboring Kazakhstan (Kadyrbekov 2003, 2005; Kadyrbekov & Aoitzhanova 2005). During field studies conducted in June and July 2014, apterous viviparous females of an unknown species were collected. These specimens were similar to Oedisiphum van der Goot, 1917, which is characterized by a very long apical rostrum segment. However, other morphological and biometric characters, such as the combination of short cauda with few setae and tapering siphunculi with an incised proximal constriction and well developed flange, placed this species within the range of morphological characteristics of the genus Brachycaudus van der Goot, On the other hand, the small ( mm) and oval to reniform stigmal pores pointed to the possibility that the specimens belong to the genus Dysaphis Börner, 1931, and this was further confirmed with the mitochondrial marker COI. Accepted by D. Ouvrard: 11 May 2017; published: 4 Jul

2 The Palaearctic genus Dysaphis contains about 110 known species (Blackman & Eastop 2017). In fauna of Tajikistan this genus is represented by 25 species, including 15 species described from this area (Narzikulov & Daniyarova 1990, Blackman & Eastop 2017). After comparison with other representatives of the genus Dysaphis, we described a new species Dysaphis (Dysaphis) kadyrovi Depa & Kanturski sp. nov. and discussed its taxonomic affiliation and the reliability of existing barcode sequence. Material and methods Study area. The collection site of the new aphid species, Sarikhosor Darvaz, is situated in the Central Tajikistan Province, which occupies the central part of Tajikistan.The climate of the Sarikhosor Darvaz region is variable, mild continental and cool. The annual average temperature is 11 C, precipitation mm/m 2 a year. Here, the largest rivers of Tajikistan Vakhsh, Surkhob, Yakhsu and Obikhingou and major glaciers, including the Pamir glaciation knot with 40% of Central Asian glaciers, are formed and the moraine lakes occur. The flora and the vegetation cover are diverse, with mesophyllic forests, high grass semi savannas, xerophytic light forests, mountain steppes and alpine meadows prevailing. The floristic composition is estimated at 4000 flowering and spore bearing plant species. The most valuable plant communities and endemic species are assigned to this region (Safarov 2003). Methods. The material was preserved in 70% ethanol and prepared after Kanturski & Wieczorek (2012). The measurements were done according to Ilharco & van Harten (1987) and Blackman & Eastop (2006). The slides were examined using a Nikon Ni U light microscope and photographed with a Nikon DS Fi2 camera. Measurements are given in mm (Table 1). The following abbreviations are used in the descriptions and table (partly after Blackman & Eastop 1994): BL body length (from anterior border of the head to the end of cauda); HW greatest head width across compound eyes; ANT antennae or their lengths; ANT I, II, III, IV, V, VI antennal segments I, II, III, IV, V, VI or their lengths (ratios between antennal segments are simply given as e.g. VI:III ); LS ANT III length of longest setae of ANT III; BD III basal articular diameter of ANT III; BASE basal part of last antennal segment or its length; PT processus terminalis of last antennal segment or its length; ARS apical segment of rostrum (IV + V) or its length; III FEMUR hind femur length; III TIBIA hind tibia length; HT I first segment of hind tarsus or its length, HT II second segment of hind tarsus or its length; SIPH siphunculi or their length; ABD I VIII abdominal tergites I VIII, A few specimens were collected to 99.8% ethanol and the DNA was extracted using DNeasy Tissue Kit by Qiagen, following manufacturer instructions, for molecular analysis of two specimens (samples C1 and C2). The standard COI primers (LCO1490, HCO2198) were applied (Folmer et al. 1994). Phylogenetic analysis of obtained sequences was performed with MEGA version 6 software (Tamura et al. 2013) and the Neighbor Joining (NJ) (Saitou & Nei 1987) method was applied. Sequence of Brachycaudus (Prunaphis) cardui (Linnaeus, 1758), obtained from Genbank, was used as an outgroup. Sequences of Dysaphis species were also obtained from Genbank and their accession numbers are given in Fig. 1. For morphological comparison with Dysaphis (D.) lappae lappae (Koch, 1854), the following material was examined: FRANCE: Oraison (B. A) 01 IX 1959, Lappa sp. Hille Ris Lambers leg., 4 apterous viviparous females, BM , Natural History Museum (BMNH), London (UK). The following publications concerning genus Dysaphis s. lat. were studied for the purpose of proper recognition of the collected specimens: Narzikulov 1961, 1967; Stroyan 1963, 1981, 1985; Shaposhnikov 1986; Chakrabarti & Medda 1993; Stekolshchikov Results. Molecular analysis of two samples, from two different specimens of the studied sample indicated their unambiguous taxonomic affiliation within the genus Dysaphis (Fig. 1). However, received sequences were in 100% identical to the sequences deposited in Genbank under taxonomic affiliation to D. (D.) lappae lappae, from samples collected in Russian part of north-western Altai Mts. (Voronova et al. 2011). We checked the specimens of D. (D.) lappae lappae from the BMNH collection for morphological similarity, to exclude the possibility of wrong determination of our samples. Dysaphis (D.) lappae lappae is characterized by the presence of spinal tubercles on ABD I and (VI)VII VIII and by large marginal tubercles on ABD I V (Fig. 2), whereas our samples were without any marginal and spinal tubercles in all specimens. 574 Zootaxa 4286 (4) 2017 Magnolia Press DEPA ET AL.

3 TABLE 1. Measurements (in mm) of apterous viviparous females (n = 9) of Dysaphis (D.) kadyrovi Depa & Kanturski sp. nov. Measurements of the holotype in bold. Character Apterous viviparous female BL HW ANT ANT III ANT IV ANT V BASE PT ROSTRUM ARS III FEMUR III TIBIA HT I HT II SIPH SIPH base width CAUDA length CAUDA width FIGURE 1. A phylogenetic tree (Neighbor Joining method) of Dysaphis based on COI mitochondrial markers with the studied samples (C1, C2) of Dysaphis (D.) kadyrovi Depa & Kanturski sp. nov. A NEW APHID SPECIES FROM CENTRAL ASIA Zootaxa 4286 (4) 2017 Magnolia Press 575

4 Such features of the morphology of the studied specimens clearly distinguished them from other representatives of the subgenus Dysaphis, but were accordant with the characteristics of the former subgenus Neodysaphis Narzikulov, 1961, now treated as a synonym of Dysaphis (Stroyan 1985), but characterised as the D. pseudomolli species group by Stroyan (1985). After comparison of collected specimens with the species of this group Dysaphis (D.) pseudomolli Narzikulov, 1961 and Dysaphis (D.) cousiniae cousiniae Narzikulov, 1967 (and its subspecies: D. (D.) cousiniae minor Narzikulov, 1967) we decided that it is a new species. Another interesting result of the molecular part of the study was the placement of D. affinis, of uncertain taxonomic position in a clade with representatives of the subgenus Pomaphis. Dysaphis (Dysaphis) kadyrovi Depa & Kanturski sp. nov. (Figs 2b, d, f; 3 7; Table 1) Type material: HOLOTYPE: TAJIKISTAN: valley of the Vakhsh River, 38 51'52.7"N, 70 01'32.3"E, 11 VII 2014, on Anacantha darwasica (C. Winkl.) Soják; Taszakowski & Walczak leg., 1 apterous viviparous female, marked as holotype on the slide, 2014/07/1 UŚ. Paratypes: the same data as holotype: 2 apterous viviparous females marked as paratype, 2014/07/01, UŚ; 2 apterous viviparous females, 2014/07/02 UŚ; 2 apterous viviparous females 2014/07/03, UŚ; 2 apterous viviparous females, 2014/07/04, UŚ. Holotype and paratypes of the new species are deposited in the Department of Zoology, University of Silesia in Katowice (UŚ). Paratypes will be also deposited in the Natural History Museum (BMNH), London (UK), 2 apterous viviparous females, 2014/07/02 UŚ; Muséum national d'histoire naturelle (MNHN), Paris (France), 2 apterous viviparous females, 2014/07/03 UŚ; and United States Department of Agriculture (Smithsonian insect collection), Agricultural Research Service (USDA), Beltsville (USA), 2 apterous viviparous females, 2014/07/04 UŚ. Description. Apterous viviparous female (based on nine specimens) Colour: in life, greyish green to dark green, not covered by wax. Head and prothorax green to light brown, ANT and legs brownish. End of abdomen brownish, SIPH brown (Figs 3, 4). In mounted specimens: head sclerotized pale to light brown, ANT pale to light brown. When ANT light brown then ANT III basal part and PT lighter. Pronotum sclerotized, pale to light brown. Legs pale to light brown; femora pale or uniformly light brown, tibiae pale or light brown with darker anterior and posterior part, tarsi pale to light brown. SIPH and cauda pale to light brown (Figs 5 7). Morphology. Dorsal side of head with 6 pairs of spindle shaped, blunt or slightly blunt setae (Fig. 6a); ventral side with 4 pairs of rather fine and slightly pointed to slightly blunt setae. Head setae mm long. HW times ANT. ANT (Fig. 6b) times BL. ANT III shorter or longer than ANT VI, times BL. ANT IV times ANT V. ANT V times BASE. ANT VI with PT times BASE, with rounded primary rhinarium and 3 4 small accessory rhinaria tightly adjoining each other directly to the major rhinarium (Fig. 6c). Other antennal ratios: ANT VI:ANT III ; ANT V:ANT III ; ANT IV:ANT III Antennal chaetotaxy: ANT I with 5 7 setae, ANT II with 4 5 setae, ANT III with 5 11 setae, ANT IV with 3 7 setae, ANT V with 2 4 setae; ANT VI with 4 basal, 4 apical and 1 2 setae on basal part of PT. Antennal setae short, mm long and pointed; LS ANT III times BD III. Rostrum, reaching from middle coxae to ABD V. ARS times ANT III, times BASE, times ANT VI and times HT II, with 2 3 accessory setae (Fig. 7a). Thorax with short to medium rigid, spindle shaped setae, on pronotum mm long and placed antero-marginally and spinally, mm long on mesonotum and mm long on metanotum. Mesothoracic furca transparent, separated; metathoracic furca short, weakly developed. III FEMORA times BL, III TIBIAE times BL, posterior seta on hind trochanter times diameter of trochantro-femoral suture. HT II times HT I. First segments of tarsi with 3:3:3 ventral setae (Fig. 7b). HT II times ANT III, times BASE and times ANT VI. Empodial setae fine and pointed. ABD I VIII covered by short or medium rigid and blunt setae in spinal, pleural and marginal positions, setae mm long on ABD I V, 6 10 setae mm long on ABD VI VII and 6 (rarely 5) setae mm long on ABD VIII. Dorsal setae on ABD III times BD III. Dorsal surface of 576 Zootaxa 4286 (4) 2017 Magnolia Press DEPA ET AL.

5 ABD segments in form of polygons with irregular edges. ABD I IV membranous, ABD V with spinal or spino pleural scleroites and sclerites. Sclerotic cross bars of ABD VI VIII imbricated or covered by very short spinules. SIPH (Fig. 7c) times CAUDA, times BL and times ANT III, slightly imbricated with poorly developed flange and subapical constriction. Subgenital plate well developed with 2 anterior and posterior fine and pointed setae. Cauda with 5 fine and pointed setae (Fig. 7d). Etymology. The authors have the pleasure to name the species to honor Prof. Abdusalom Hafizovich Kadyrov Entomologist from the Faculty of Biology, Tajik State National University in Dushanbe and long term friend of the Department of Zoology, University of Silesia. Diagnosis. The new species differs from most of other species of Dysaphis by lack of marginal and spinal tubercles. Due to this character and slender and elongated ARS, the new species is similar to D. pseudomolli and D. cousiniae from which it differs by the following characters: Antennal setae long, LS ANT III times BD III, while in D. (D.) pseudomolli LS ANT III are no longer than 0.5 times BD III. Setae on ABD I V mm, while in D. (D.) pseudomolli they are mm long. 4 setae on the apex of PT, among which 3 are apical whereas in D. (D.) pseudomolli there are only 3 setae, with 2 apical. There are 6 setae on ABD VIII while in D. (D.) pseudomolli there are 4. Marginal tubercles always absent while in D. (D.) pseudomolli they are small and only rarely absent. From D. (D.) cousiniae cousiniae it differs by: Lack of sclerotized abdominal shield on ABD I V. Lack of marginal and spinal tubercles on all body segments. From D. (D.) cousiniae minor it differs by: Lack of marginal tubercles on all abdominal tergites Slender, stiletto shaped apical segment of rostrum FIGURE 2. Comparison of morphological features of D. (D.) lappae lappae: (a) ARS; (c) margin of abdomen with big marginal tubercles; (e) end of abdomen with spinal tubercles; and D. (D.) kadyrovi Depa & Kanturski sp. nov. (b) ARS; (d) margin of abdomen without any tubercles; (f) end of abdomen without any tubercles. A NEW APHID SPECIES FROM CENTRAL ASIA Zootaxa 4286 (4) 2017 Magnolia Press 577

6 FIGURE 3. Coloration and host plant of Dysaphis (D.) kadyrovi Depa & Kanturski sp. nov.: (a) host plant Anacantha darwasica; (b) apterous viviparous females on upper surface of leaf; (c) colony of apterous viviparous females on inflorescence and stem attended by ants. Host plant and biology: The species is associated with Anacantha darwasica (C. Winkl.) Soják (syn.: Modestia Char. & Tamamsch.) (Fig. 3). The host plant is endemic to the Pamiroalai Mountains in Tajikistan (Gissarskij Chrebet, Darvaskij Chrebet, Karategin, approx. 38 N, E). Habitat: Subalpine meadows, screes and gravel embankments of rivers between 1300 and 3200 m (Fig. 8) (Soják 1982; Häffner 2000). The aphids were feeding on upper sides of leaves and on upper shoots near inflorescence and were attended by ants from the genus Plagiolepis Mayr, 1861 (Formicinae), det. Ł. Depa (after key to subfamilies and genera in Hölldobler & Wilson 1990). Locality '52.7"N, 70 01'32.3"E; valley of the Vakhsh River, Central Tajikistan province, Sarikhosor subdarvaz region; 1160 m asl; dried river bed, overgrown by sparse herbaceous vegetation (Fig. 8). 578 Zootaxa 4286 (4) 2017 Magnolia Press DEPA ET AL.

7 FIGURE 4. Sclerotization and size variation of Dysaphis (D.) kadyrovi Depa & Kanturski sp. nov. Apterous viviparous female in: (a) dorsal; (b) lateral; (c) ventral view; (d) sclerotization of head and pronotum; (e) sclerotization of ABD V VIII; (f) differences in body size among representatives. Discussion Taxonomical comments. The COI data place the new species firmly within the subgenus Dysaphis. Representatives of this subgenus are characterized by mesothoracic furca separated, which is concordant with features of D. (D.) kadyrovi Depa & Kanturski sp. nov., but also well developed marginal tubercles and the presence of at least one spinal tubercle on ABD VIII, which D. (D.) kadyrovi lacks. In D. lappae the spinal and marginal tubercles are particularly well-developed and numerous. The existence of such morphological differences between samples with identical COI sequence is astounding. There are only two explanations: marginal and spinal tubercles are an extremely variable taxonomical character in the subgenus Dysaphis, or the sequence deposited in Genbank is wrongly assigned to D. (D.) lappae. A NEW APHID SPECIES FROM CENTRAL ASIA Zootaxa 4286 (4) 2017 Magnolia Press 579

8 The morphological characterization of the genus Dysaphis is a subject of vast aphidological literature. Detailed investigations and discussions by Stroyan (1963,1981), Shaposhnikov (1986) and Stekolshchikov (1998) show significant variability of the feature of marginal and spinal tubercles not only between the species but also between morphs within a single species. The new species may be assigned to the D. (D.) pseudomolli group of species (Stroyan 1985), a group endemic to Central Asia, associated with Asteraceae-Cynareae and having reduced marginal and spinal tubercles diagnostic features of the subgenus Neodysaphis (Narzikulov, 1961). However, the synonymy of Neodysaphis Narzikulov with Dysaphis s. str. may be confirmed by two facts: molecular positioning of the new species well within the subgenus Dysaphis and presence of separated, although very small mesothoracic furca. An interesting feature of the new species is a disparity in body size among collected specimens (Fig. 4f). Bigger specimens are also more pigmented than the small ones, but their morphological features (e.g. morphometric ratios) overlap. The sample was collected after the blossom period of A. darwasica, so smaller body size of some specimens may be linked to the worsening physiological condition of their host plant. This raises the question of the taxonomic position of Narzikulov s (1967) D. (D.) cousiniae minor, which might be interpreted simply as small specimens of D. (D.) cousiniae. However, even the smallest specimens of D. (D.) kadyrovi Depa & Kanturski sp. nov. reprise all morphological features (qualitative and quantitative) of the biggest specimens. D.(D.) cousiniae minor on the other hand differs from the nominate subspecies by elongated body, lack of spinal tubercles, shorter antennae and shorter and wider apical segment of rostrum. These are in our opinion features significant enough and Dysaphis (D.) cousiniae minor may be treated as full status species, despite sharing the same host plant Cousinia radians. FIGURE 5. Dysaphis (D.) kadyrovi Depa & Kanturski sp. nov. apterous viviparous female holotype, general view. 580 Zootaxa 4286 (4) 2017 Magnolia Press DEPA ET AL.

9 FIGURE 6. Dysaphis (D.) kadyrovi Depa & Kanturski sp. nov.: (a) head; (b) antenna; (c) ANT V and ANT VI. Reliability of barcode sequence. The identity of the barcode sequence deposited in Genbank under D. (D.) lappae with the sequence of D. (D.) kadyrovi Depa & Kanturski sp. nov. despite the great morphological dissimilarities between the two species (Fig. 3), puts in question the reliability of taxonomic affiliation of available molecular sequences and their usefulness as barcodes. Wrongly identified source specimens may give rise to taxonomic confusion and make barcoding untrustworthy. Such mistakes may also influence any phylogenetic studies that make use of existing but wrongly ascribed sequences a threat already expressed by some studies (Taylor & Harris 2012, Collins & Cruickshank 2013). It especially concerns groups of species with difficult morphology involving a considerable degree of overlap of morphological features between species, as in the genus Dysaphis. This is a difficult genus, with more than 100 species, significant intraspecific variability and often with host plant affiliations, life cycles and morphs that are not fully recognised, all of which could lead to misidentification and incorrect barcoding. This may lead to further complications with molecular resources. On the other hand, so far conducted studies with molecular markers on Dysaphis also show significant problems with proper barcoding of species (Coeur d acier et al. 2014). This may result from high morphological variability of Dysaphis in response to environmental factors, so strict studies correlating morphological and molecular variability are required in this group, with trusted connection between the barcode and the morphospecies/environment/ hostplant of the sample from which barcode originated. A NEW APHID SPECIES FROM CENTRAL ASIA Zootaxa 4286 (4) 2017 Magnolia Press 581

10 FIGURE 7. Dysaphis (D.) kadyrovi Depa & Kanturski sp. nov.: (a) ARS; (b) hind tarsus; (c) SIPH; (d) cauda with anal plate (ap). Although it is possible to extract DNA from individuals later used for morphological studies (Favret 2005; Miller et al. 2013), it is hardly possible to provide DNA for type specimens of most of the taxa of aphids so far described, due to DNA-deteriorating mounting procedures (contrary to e.g. sponges, where similar problems exist to a significantly higher extent Erpenbeck et al. 2016). It may be necessary, in future taxonomic studies, to carry out not only morphological revisions of taxa but also, to avoid further confusion, molecular revisions of existing sequences. A similar proposal has already been formulated in case of fungi (Hawksworth et al. 2016). Also, any new species should have its typical sequence assigned as a part of the taxonomic description, to provide a point of reference for such revisions, a procedure already applied by some authors (Lis et al. 2014). Acknowledgements The authors are sincerely grateful to Arkadiusz Nowak, University of Opole (Poland) for valuable help with determination of the host plant. Special thanks to Colin Favret, Department for Biological Sciences, University of Montréal (Canada) for his help with collection of the literature. Special thanks go to Łukasz Kajtoch, Institute of Systematics and Evolution of Animals, Polish Academy of Sciences (Poland) for his assistance with molecular study and Wojciech Szczepański, University of Silesia (Poland) for the photograph of the locus typicus. We are very grateful to the anonymous Reviewers, who greatly improved the current version of the manuscript. 582 Zootaxa 4286 (4) 2017 Magnolia Press DEPA ET AL.

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