Diorhabda carinulata ( Desbrochers, 1870 )

Tracy, James L. & Robbins, Thomas O., 2009, Taxonomic revision and biogeography of the Tamarix-feeding Diorhabda elongata (Brullé, 1832) species group (Coleoptera: Chrysomelidae: Galerucinae: Galerucini) and analysis of their potential in biological control of Tamarisk, Zootaxa 2101 (1), pp. 1-152 : 86-102

Diorhabda carinulata ( Desbrochers, 1870)

northern tamarisk beetle

( Figs. 7, 8 , 17 , 22 , 27 , 32 , 37 , 42 )

Galeruca carinulata Desbrochers, 1870:134 (Type locality: Sarepta , Russia).

Diorhabda elongata var. sublineata: Weise, 1890:484 (part, central Mongolia).

Diorhabda elongata var. carinata: Weise, 1893:635 (part, Sarepta, Russia).

Diorhabda elongata ab. carinata: Weise, 1924:78 (part; world catalog; Sarepta, Russia); Winkler, 1924 –1932:1307 (part; Palearctic catalog; southern Russia)

Diorhabda elongata ab. sublineata: Weise, 1924:78 (part; world catalog; Mongolia); Laboissière, 1934:54 (part, Mongolia); Winkler, 1924 –1932:1307 (part, Palearctic catalog, Mongolia); Medvedev and Voronova, 1977b:238 (part, keys, Mongolia); Medvedev, 1982:259 (part, keys to adults and larvae, Mongolia).

Diorhabda koltzei ab. basicornis Laboissière, 1935:324 (Type locality: Khotan, Taklamakan Desert, Turkestan [Hotan, Xinjiang Uygur Zizhiqu, China]) (NEW SYNONYM); Wilcox 1971:63 (world catalog, Karakorum [western China]).

Diorhabda elongata: Ogloblin, 1936:79 (part, Russia, transcaucasus, Iran, central Asia, central Mongolia); Rusanov, 1949:118 (part, Central Asia, as Diorrhabda); Kyrzhanovskiy, 1952:198 (part, Turkmenistan), 1965:392 (part, middle Asia); Pavlovskii and Shtakelberg, 1955:566 (part, southwest Russia, transcaucasus, central Asia, Iran, Mongolia, as Diorrhabda); Yakhontov and Davletshina, 1955:58 (part, biology, Amu Darya Delta, northern Uzbekistan); Sinadsky, 1957:950, 1963:84, 1968:64 (part, biology, Amu Darya Valley, Uzbekistan, as Diorrhabda); Yakhontov, 1959:338 (part, biology, Uzbekistan); Medvedev, 1959:118 (part, Turkmenistan); Kulinich, 1962:73 (part, biology, Tajikistan); Kryzhanovsky, 1965:392 (part, middle Asia); Pripisnova, 1965:83 (part, biology, Tajikistan); Kulenova 1968:171 (part, southeastern Kazakhstan); Lopatin and Tadzhibaev, 1972:591 (part, Tajikistan); Lopatin, 1977a:282 (part, keys, Asia); Medvedev and Voronova, 1977a:339 ( Mongolia), 1977b:238 (part, keys, Mongolia), 1979:119 (catalog for Mongolia, hosts); Davletshina et al., 1979:79 (part, southwest Kyzyl–Kum Desert, central Uzbekistan); Ogloblin and Medvedev, 1971:83, 89 (part, key to larva, European Russia); Medvedev, 1982:99 (part, keys to adults and larvae, Mongolia); Samedov and Mirzoeva, 1985:712 (part, Azerbaijan); Lopatin and Kulenova, 1986:129 (part, Kazakhstan); Kovalev, 1995:78 (part, south central Palearctic); Myartseva, 1995:4, 1999:1; 2001:1 (part, biology, Turkmenistan); DeLoach et al., 1996:253 (part, China); Ishkov, 1996:30 (part, Ili River Valley, Kazakhstan); White, 1996:392 (part, China); Mityaev and Jashenko, 1997:4, 1998:7, 1999:6, 2000:14, 2001:9, 2007:3 (part, biology, ecology; southeast Kazakhstan); Gruev and Tomov, 1998:70 (part, Transcaucasus, mid-Asia, Mongolia); USDA- APHIS, 1999:13395 (Fukang, China; approval of initial introduction into U.S.); Anonymous, 2001:52 N (part, Kazakhstan, China); Gould and DeLoach, 2002:302 (part, biological control, North America); Dudley et al. 2000:346 (part, biological control, North America), 2001:260 (part, biological control, North America); Stenquist, 2000:492 ( Kazakhstan); Jolivet, 2001:134 (part, U.S. introduction); Vail et al. 2001:37 (part, U.S. introduction); DeLoach et al., 2003a:230 (part, China, Kazakhstan), 2003b:126 (part, host range; Uzbekistan, Turkmenistan, Iran); Khamraev, 2003:11 (part, Uzbekistan); Petroski, 2003:3234 (synthesis of aggregation pheromone components); Milbrath et al., 2003:225 (part, China, Kazakhstan); Riley et al., 2003:69,189 (part, catalog, North America [introduced]); Warchalowski, 2003:328 (part, taxonomic keys, Caucasus, Central Asia, and Mongolia); Bieṅkowski, 2004:76 (part, keys, eastern Europe (part, southern Russia, Caucasus, Iran, Iraq, Central Asia, Kazakhstan, Mongolia); DeLoach and Carruthers, 2004b:311 (part, China and Kazakhstan); Dudley and DeLoach, 2004:1542 (part, biological control, North America); Lopatin et al., 2004:127 (part, central Asia); Cossé et al., 2005:657 (aggregation pheromone, Fukang, China, population), 2007:2695 (Fukang, China; attraction to green leaf volatiles); Dudley, 2005a:13, 2005b:42 N (part, biological control, ex: China and Kazakhstan); Herrera et al., 2005:775 (part, developmental rates, Fukang, China, population); Carruthers et al. 2006:71, 2008:258 (part, biological control, ex: China and Kazakhstan); Milbrath and DeLoach, 2006a:32 (part, host specificity, Turpan, China, populations); Geraci et al. (in press) (remote sensing of defoliation, Nevada); Milbrath et al. (2007) (part, biology, Fukang, China, population); Dudley et al., 2006:137 (part, releases in Nevada, Fukang, China; host range, Fukang and Turpan, China); Longland and Dudley (2008) (ecology, North America); Dudley et al. (in prep.) (host range, Nevada); Thompson et al. (in prep.) (part, laboratory hybridization, Fukang, China).

Diorhabda rybakowi: Mityaev, 1958:86 (part, biology, Kazakhstan, as rybakovi).

Diorhabda elongata deserticola Chen, 1961 (Type locality: Yuli (= Wei–li), Xinjiang Uygur Zizhiqu, China) (NEW SYNONYM); Gressitt and Kimoto, 1963b:930 ( China); Wilcox, 1971:63 (world catalog, Xinjiang Uygur Zizhiqu, China); Tian et al., 1988:24 (biology, ecology; Ningxia Province, China); Bao, 1989:45 (biology and control as pest; Nei Mongol Zizhiqu, China); Xiao, 1992:537 (biology, China); Sha and Yibulayin, 1993:7 (biology, ecology, control as pest, China, Xinjiang Prov.); Chen et al., 2000:44 (control as pest; Alashan Meng, Nei Mongol Zizhiqu, China); USDA-APHIS, 2005:1 (Fukang, China; large-scale introduction to 13 states in northwestern U.S.); Li et al., 2000:48 (biology, ecology; China, Xinjiang Prov., Fukang); Eberts et al., 2001:3 (Fukang, China, establishment in Colorado); Lair and Eberts, 2001:3 ( China, potential introduction into north Texas); Zhang 2002:1 (biology and ecology, Xinjiang Uygur Zizhiqu, China); DeLoach et al., 2003a:229 ( China, Kazakhstan), 2003b:117 (host specificity; Turpan and Fukang, China and Chilik, Kazakhstan; host range; China, Kazakhstan, Mongolia), 2004:505 (field establishment in North America from populations in Fukang, China and Shelek, Kazakhstan), (2008, in prep.) (part, China); Lewis et al., 2003a:148 (host specificity; Fukang, China and Chilik, Kazakhstan), 2003b:101 (developmental and reproductive biology; Fukang, China and Chilik, Kazakhstan); Meng and Baoping, 2003:99, 2005:192 (biological control; Xinjiang Uygur Zizhiqu, China), 2006:189 (ecology; Xinjiang Uygur Zizhiqu, China); DeLoach and Carruthers, 2004a:10 (field establishment in North America from populations in Fukang, China and Shelek, Kazakhstan); Ding, 2004:57 (control as pest; Anxi County, Gansu Sheng, China); Ming et al. 2004:283 (Xinjiang Uygur Zizhiqu, China); Lopatin et al., 2004:127 (east Kazakhstan, northwest China); Aber et al., 2005:63 (field monitoring in Colorado; ex: Fukang, China); Dudley and Kazmer, 2005:265 (host specificity, Fukang, China, population); Peng et al., 2005:63 (control as pest; Gansu Sheng, China); Zheng et al., 2005:136 ( China); Ding et al., 2006:1442 (biological control, ex: Kazakhstan, China); Li and Wang, 2006:27 (biology, control as pest; Gansu Sheng, China); Bean et al., 2007a:15 (diapause, Fukang, China); Mityaev and Jashenko, 2007:8 (biological control, ex: Kazakhstan, China); Bean et al., 2007b:531 (diapause, Fukang, China); Hudgeons et al., 2007a:158 (biological control; Fukang, China, Chilik, Kazakhstan), 2007b: 215 (part, tamarisk damage in Nevada, ex: Fukang, China); Dalin et al. (in press) (host range; Turpan, China).

Diorhabda elongata sublineata: Gressitt and Kimoto, 1963a:407 (part; keys; northwestern China and Mongolia); Lopatin, 1968:214 ( Mongolia); 1970:254 ( Mongolia, on Tamarix ); 1975:219 ( Mongolia); 1977b:154 ( Mongolia).

Diorhabda carinulata carinulata: Berti and Rapilly, 1973:881 (restored species; Sarepta, Russia); DeLoach et al., 2003b:126; Warchalowski, 2003:328 (taxonomic keys, southern Russia).

Diorhabda deserticola: Yu et al., 1996:94 (taxonomic keys and descriptions, China); Meng et al., 2005:27 (ecology, Xinjiang Uygur Zizhiqu, China); Zhang and Baoping, 2006:109 (behavior and reproductive biology, Xinjiang Uygur

Zizhiqu, China).

Male. Genitalia. Male D. carinulata can be distinguished from all other species in the D. elongata group by two unique characteristics of the palmate endophallic sclerite (PES): (1) the distal margin is acutely rounded and generally smooth, with one or two small subdistal spines, and (2) the PES has a length to width ratio of 0.61–1.02 ( Figs. 17 , 32 ; Table 3). In contrast, the distal margin of the PES is truncate or more broadly rounded with larger distal or subdistal spines in D. elongata , D. carinata , and D. sublineata ( Figs. 14–16 , 29–31 ), and the PES is narrowly rounded with a length to width ratio of 0.35–0.63 in D. meridionalis ( Figs. 18 , 33 ). The length of the spined area (SL) of the elongate endophallic sclerite (EES) is greater than or equal to 0.33 times (or greater than about one third) the length of the EES in D. carinulata ( Figs. 17 , 22 ), compared to the spined area being less than or equal to 0.16 times (or less than about one fifth) the length of the EES in D. elongata ( Table 3; Figs. 14 , 19 , 24 , 48 ). In D. carinulata , the blade of the EES extends less than 2/3 the total length of the sclerite ( Figs. 17 , 22 ), and the EES lacks a strong hook at the apex in dorsal aspect ( Fig. 27 ). In contrast, the blade of the EES extends for more than 2/3 the total length of the EES in D. meridionalis ( Figs. 18 , 23 ), and the apex of the EES is strongly hooked in dorsal aspect ( Fig. 28 ).

Measurements. See Tables 2 and 3.

Female. Genitalia. Female D. carinulata can be distinguished from all other members of the D. elongata group by the following combination of characters in the vaginal palpi (VP) and internal sternite VIII (IS VIII): (1) the vaginal palpi are broadly rounded and about as long as wide or longer with a width to length ratio (LP/ WP) of 0.94–1.36 ( Fig. 37 ; Table 4), (2) if the width to length ratio of the vaginal palpi is 0.94, then the width of the widest lobe of the stalk (WLS) of IS VIII is greater than or equal to 0.11 mm ( Fig. 37 ), and (3) the width of the stalk (WST) of IS VIII is 0.36–0.57 mm and this width is 0.49–0.77 times the width of the apical lobe (WAL) ( Figs. 37 , 42 ; Table 4). In contrast, the vaginal palpi are triangulate and wider than long with a width to length ratio of 0.46–0.89 in D. carinata and D. sublineata ( Figs. 35–36 ). In D. elongata the vaginal palpi are also broadly rounded but are wider than long with a length to width ratio of 0.52–0.94, and when the width to length ratio of the vaginal palpus is 0.94, the width of the widest lobe of IS VIII is less than or equal to 0.10 mm ( Fig. 34 ). In D. meridionalis , the width of the stalk of IS VIII is narrower than in D. carinulata , measuring 0.22–0.33 mm and this width is from 0.33–0.48 times the width of the apical lobe ( Fig. 38 , 43 ). In addition, the width of the widest lobe of the stalk of IS VIII is usually greater than 0.10 mm in D. carinulata ( Figs. 37 , 42 ); whereas, in D. meridionalis the width of the widest lobe of the stalk is from 0.04–0.09 mm ( Fig. 38 , 43 ; Table 4).

Measurements. See Tables 2 and 4.

Coloration. Coloration and elytral vittae of dead ( Fig. 7 ) and living ( Fig. 8 ) D. carinulata is very similar to that discussed above for D. sublineata ( Figs. 5, 6 ). Chen (1961) also noted the similar appearance of D. carinulata (as D. e. deserticola ) and D. sublineata (as D. e. ab. sublineata ) from North Africa.

Type material. According to Berti and Rapilly (1973), the Desbrochers type material for D. carinulata , consisting of a male holotype, is deposited in the Demaison Collection at MNHN. As discussed above (see D. elongata – Type material), we were unable to obtain type materials from MNHN. We studied the original description by Desbrochers (1870), which he based on a single individual 5.5 mm in length, the illustrations of the endophallus of the male holotype by Berti and Rapilly (1973), and topotypes from Sarepta, Russia and the neighboring area.

Chen’s (1961) type material for D. e. deserticola consists of a holotype male and allotype female from Wei–li (=Yuli), Xinjiang Uygur Zizhiqu, China, and 25 paratypes (8 ♂♂, 17 ♀♀) in Xinjiang from Akesu (= Aksu), Atushen (= Atux), Baicheng, K'u–ch'e (Kuqa), Miquan, Turtiaogou, Shajingzi, Yecheng (= Kargilik), and Wei–li. These are deposited at the Institute of Zoology, Chinese Academy of Sciences, Beijing, China ( IZAS). We studied four paratypes (2 ♂♂, 2 ♀♀) from Wei–li County, Xinjiang Uygur Zizhiqu, China (from IZAS), and the original description of D. e. deserticola by Chen (1961).

We studied the type material of D. koltzei ab. basicornis Laboissière from Khotan (= Hotan ), Turkestan (now in Xinjiang Uygur Zizhiqu, China) which consists of a single female according to Laboissière (1935) that Ron Beenen kindly located for us in the Zoological Museum Amsterdam ( ZMAN) .

Material examined. 120♂♂ dissected (diss.), 52♀♀ diss., 199♂♂, 230♀♀, 128 unsexed specimens. AZERBAIJAN: 1♂ diss., Gobustan [Qobustan; 40.08416°N, 49.41583°E], 9-10-VII-2001, V. Dolin, MBPF [2003-07]; 2♀♀ diss., Ordubat [Ordubad; 38.9081°N, 46.0278°E], [19]14, Dr. Veselý, NMPC [2004-26, 2005-37]; 1♀ diss., Ordubad, Araxes [Aras River] Road, Dr. J. Veselý, 9131, NMPC [2004-18]; CHINA: Nei Mongol Zizhiqu (Inner Mongolia): 1♂ diss., Erjina County [specific location not given], 15-VII-1960, Forest Department, Forest Science Institute, Erjina Co., on Tamarix, IZAS [2005-05]; Gansu Sheng: 1♂ diss., 1♀ diss., 1♀, Jiayuguan, 3 km southeast [39.7900°N, 98.3200°E], north side road, 40,238 km odom., 29-VII-1993, C.J. DeLoach, on T. ramosissima , Diorhabda elongata deserticola det. I. Lopatin 2000, 6682–6684, GSWRL [2003-53, 54]; 1♂ diss., 9♀♀, Lanzhou, 4 km east [36.05639°N, 103.84000°E], 39,249 km odom., 26-VII-1993, C.J. DeLoach, on T. hohenackeri [= T. smyrnensis ]/ T. ramosissima , D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 6♀♀], 6677–6681, 6729, 8980, GSWRL [2003-48]; Xinjiang Uygur Zizhiqu: 1♂ diss., Bogda Shan Mtns., 80 km northeast Urumqi [Urumchi; 43.81850°N, 87.97160°E]; 4-VI-1993, Mityaev, on saksaul [ Haloxylon ammondendron (C.A. Meyer) Bunge (Chenopodiaceae) , apparently incidental occurrence], USNM [2003-15]; 1♀ diss., 5♂♂, 8♀♀, Fukang [44.1667°N, 87.9833°E], 8-VIII- 1995 [not 13-VIII-1995 as in some records], Lu Qing-guang, M.T. Liu & Jiang Xi Liang, on T. ramosissima , D. e. deserticola Chen det. I. Lopatin 2000 [1♀], 8584–8586, 8588, 8591–8593, GSWRL( CJD)-1995-25, GSWRL [2005-20] [11 specimens from Lot GSWRL( CJD)-1995-9 identified as prob. D. e. deserticola by R.E. White on 15-XI-1995 at USDA SEL, SEL Lot # 95-10233]; 1♂ diss., 1♂, 2♀♀, Fukang, 8-IX-1996, Li Bao Ping, on T. ramosissima , voucher shipment GSWRL( CJD)-1996-27, GSWRL [2005-90] [9 specimens from Lot GSWRL( CJD)-1996-4 identified as D. elongata by S.M. Clark on 28-X-1996 at West Virginia Dept of Agric.]; 1♂ diss., 1♂, 1♀, Fukang, 8-VI-1996, Li Bao Ping, on Tamarix sp. prob. ramosissima, GSWRL [2006-02]; 1♀ diss., Fukang, southwest, 9-VI-1996, P.E. Boldt, on T. karelinii [ T. hispida var. karelinii ], GSWRL [2006-01]; 1♂ diss., 11♂♂, 7♀♀, Fukang, Tamarix Park , 1-IX-1997, Li Bao Ping, on Tamarix sp. , voucher shipment GSWRL( CJD)-1997-22, GSWRL [2005-100]; 1♂ diss., 1♀ diss., 15♂♂, 10♀♀, Fukang, Tamarisk Park, 18-IV-1998 [not 11-V-1998 as on some shipment records and labels], Li Bao Ping, on T. ramosissima , 805, 806, 808, 811, 812, 814–815, 820, 821, 823–825, voucher shipment GSWRL-1998-05, GSWRL [2005-67, 68] [parasite Erynniopsis antennata Rondani ( Diptera : Tachinidae ) emerged from 21 adult specimens in Lot GSWRL-2000-04 identified by N. E. Woodley on 23-II-2000 at USDA SEL, SEL Lot # 0001060]; 1♂ diss., 4♂♂, 8♀♀, Fukang, 13-VII-1999, Li Bao Ping, on Tamarix spp. , voucher shipment BCW/WRRC-99-04 received 21-VII-1999, GSWRL [2005-75] [7 specimens from Lot GSWRL-1999-2 identified as D. e. deserticola by S. Lingafelter on 23-VII-1999 at USDA SEL, SEL Lot # 9904542] [ USDA lab colony at Albany, CA] [introduced into cages in Nevada, Colorado, and Wyoming in 1999; released in 2001] [used in biological studies of DeLoach et al. (2003b), Lewis et al. (2003 a, 2003b), Petroski (2003), Cossé et al. (2005), Herrera et al. (2005), Milbrath and DeLoach (2006a), Herr et al. (in prep.), Bean and Keller (in prep.), and Thompson et al. (in prep.)] [in field studies by DeLoach et al. (2004), Dudley and Kazmer (2005), Aber et al. (2005), USDA-APHIS (2005), and Dudley et al. (in prep.)]; 1♂ diss., 2♀♀ diss.; Fukang, 13-VII-1999, Li Bao Ping, on Tamarix spp. , voucher shipment BCW/WRRC-99-04, USDA/ARS lab colony at Temple, Texas (ex Lovell, WY, 2002, J.L. Tracy), GSWRL [2003-28, 29, 32]; 1♂ diss., 1♂, 3♀♀, Fukang, ca. 3 km west [44.14240°N, 87.92806°E], km marker 748, 15-VII-1992, R.Wang & C.J. DeLoach, on T. arceuthoides [1♂ diss], on T. hispida var. karelinii – no flowers [1♂, 2♀♀], on T. laxa – no flowers [1♀], D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 1♂, 2♀♀], 8974, GSWRL [2005-18]; 2♀♀ diss., Fukang, ca. 3 km west, km marker 749, 15-VII-1992, R. Wang & C.J. DeLoach, on T. ramosissima very few flowers [1♀ diss.], on T. ramosissima foliage [1♀ diss.], D. e. deserticola det. I. Lopatin 2000, 6728, 6659, GSWRL [2005-12,13]; 1♂ diss., 3♂♂, 2♀♀, Fukang, 7 km west [44.13658°N, 87.87737°E], 10,406 km odom., 7-VII- 1992, R. Wang & C.J. DeLoach, on T. ramosissima [1♂ diss., 2♂♂, 1♀], on T. laxa - no flowers [1♀], D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 2♂♂, 1♀], 6646–6649, GSWRL [2005-17]; 4♂♂ diss., 6♂♂, 6♀♀, Fukang, 18 km west [44.11252°N, 87.77169°E], 14,731 km odom., 9-VIII-1993, C.J. DeLoach, on T. leptostachya [1♂ diss., 2♂♂, 1♀], on T. karelinii [ hispida var. karelinii ]-some flowers (red) [1♂ diss., 1♂, 2♀♀], on T. ramosissima -some flowers [1♂ diss., 1♂, 1♀], on T. elongata [1♂ diss., 1♂, 1♀], T. laxa – no flowers [1♂, 1♀], D. e. deserticola det. I. Lopatin 2000 [4♂♂ diss., 3♂♂, 4♀♀], 6662–6669, 6693, 6694, GSWRL [2003-47, 2005-07, 19, 22]; 1♂ diss., Jiashi [39.4883°N, 76.7125°E], 11-VII-1992, R.Wang & C.J. DeLoach, on T. ramosissima , D. e. deserticola det. I. Lopatin 2000, 6515, GSWRL [2005-04]; 1♀ diss., Jiashi, 9 km west [39.45198°N, 76.61615°E], 11-VII-1992, R. Wang and C.J. DeLoach, on T. ramosissima , D. e. deserticola det. I. Lopatin 2000, 6658, GSWRL [2005-05]; 1♂ diss., Khotan [Hotan; 37.09972°N, 79.92694°E], D. elongata ab. sublineata Lucas, HNHM [2003-14]; 1♂ diss., 1♂, 1♀, Khotan [Hotan], Kashgar, 13-VI-1890, B. Grombchevskii, ZIN [2005-02]; 1♀, Khotan [Hotan], 6-12-V-1930, J.A. Sillem, Diorhabda koltzei Weise ab. basicornis Laboissière, TYPE [ Laboissière 1935], Nederlandsche Karakorum Expeditie, ZMAN [2008-01]; 1♂ diss., 2♂♂, 4♀♀, Kuldsha [Yining; 43.9000°N, 81.35000°E], Daharkent, NHMB [2003-27]; 1♂ diss., Sanchakou, 50 km west [39.93778°N, 77.8600°E], 10-VII-1992, R. Wang & C.J. DeLoach, on T. ramosissima , D. e. deserticola det I. Lopatin 2000, 6657, GSWRL [2003-57]; 2♂♂ diss., 1♂, 4♀♀, Shanshan [42.8667°N, 90.1667°E], edge of, 41,571 km odom., 5-VIII-1993, C.J. DeLoach, on T. laxa , D. e. deserticola det I. Lopatin 2000, 6671, 6673, 6674, 6676, 6689, 6690, 6730, GSWRL [2005-10, 96]; 1♂ diss., 4♂♂, 1♀, Turpan [Turpan Botany Station; 10 km SE Turpan; 42.863°N, 89.222°E], 17-VII-1992, C.J. DeLoach, on T. arceuthoides , D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 1♂], 8975, 8978, voucher shipment GSWRL( CJD)-1992-21, GSWRL [2005-15] [12 specimens from Lot GSWRL( CJD)-1993-11 identified as prob. D. e. deserticola by R.E. White on 15-X-1993 at USDA SEL, SEL Lot # 93-10349]; 1♂ diss., 2♂♂, 1♀, Turpan Botany Station [10 km SE Turpan], 17-VII-1992, R. Wang & C.J. DeLoach, on T. ramosissima , D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 1♂, 1♀,], 6660–6662, voucher shipment GSWRL( CJD)-1992-21, GSWRL [2005-16] [used in biological studies of DeLoach et al. (2003b)]; 3♂♂ diss., 2♀♀ diss., Turpan Botanical Garden, 10 km SE Turpan, 19-VI-2002, Li Bao Ping, on Tamarix spp. , voucher USDA/ARS lab colony at Temple, Texas (4-IX-2002, J.L. Tracy; 1♂, 1♀, 25-26-VIII-2003, L. Milbrath [nos. 1265, 1268]), shipment EIWRU-2002-1005, GSWRL [2003-21, 31, 33, 2004-04, 2005-84] [released in Colorado in 2004] [used in biological studies of Milbrath and DeLoach (2006a, 2006b), Herr et al. (in prep.), and Bean and Keller (in prep.)]; 1♂, Turpan, ca. 30 km east, 41,751 k m odom., 5-VIII-1993, C. J. DeLoach, 20 sweeps on T. laxa, GSWRL ; 1♀, Turpan, 74 km northwest [43.20415°N, 88.46102°E], White Poplar Valley, rd. km 687, 7-VIII-1993, C.J. DeLoach, on T. arceuthoides , D. e. deserticola det. I. Lopatin 2000, 6691, GSWRL; 1♂ diss., 1♂, 4♀♀, Turpan, 76 to 164 km northwest, 7-VIII-1993, C.J. DeLoach, on T. ramosissima , D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 1♂, 3♀♀], 8981–8984, voucher shipment GSWRL( CJD)-1993-15, GSWRL [2005-76] [11 specimens from Lot GSWRL( CJD)-1993-11 identified as prob. D. e. deserticola by R.E. White on 15-X-1993 at USDA SEL, SEL Lot # 93-10349] [used in biological studies of DeLoach et al. (2003b)]; 1♂ diss., 5♂♂, 9♀♀, Turpan, 74 to 164 km northwest, White Poplar Valley, 7-VIII-1993, C.J. DeLoach, on T. arceuthoides and T. ramosissima, GSWRL [2004-14]; 1♂ diss., Urumqi [43.8000°N, 87.58333°E], water reservoir in suburb, VII-1998, Li Bao Ping, on Tamarix sp. , voucher shipment GSWRL-1998-12, received 4-VIII-1998, GSWRL [2005-24]; 1♂ diss., 2♂♂, 2♀♀, Urumqi, 12 km northeast [43.89406°N, 87.61073°E], by fertilizer factory, 42,403 km odom., 8-VIII-1993, C.J. DeLoach, T. hispida , D. e. deserticola I. Lopatin 2000 [1♂ diss., 2♂♂, 1♀], 6685–6688, GSWRL [2005-01]; 1♂ diss., 1♀, Urumqi, 47 km southeast [43.54156°N, 87.96419°E], 10,525 km odom., 8-VII-1992, R.Wang & C.J. DeLoach, on T. ramosissima - flowers, D. e. deserticola det. I. Lopatin 2000, 8973, GSWRL [2005-11]; 1♂ diss., 2♂♂, 4♀♀, Urumqi, 57 km east [43.56707°N, 88.07503°E], 8-VII-1992, R. Wang & C.J. DeLoach, on T. ramosissima , D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 2♂♂, 3♀♀], 6645, 6650, 6651, 6653–6656, GSWRL [2005-06]; 1♂ diss., Urumqi, 47 km southwest [43.44176°N, 87.23066°E], 10,525 km odom., 8-VII- 1992, C.J. DeLoach, on T. ramosissima, GSWRL [2005-37]; 1♀ diss., Wei Lei Co. Farm 6 [near Yuli: 41.33060°N, 87.25780°E], 14-V-1960, Wang Shuyong, D. e. deserticola det. Chen, PARATYPE, IZAS [2005-04]; 1♂ diss., Wei Lei Co. Farm 6 [near Yuli], 14-V-1960, Zhang Facai, D. e. deserticola det. Chen, PARATYPE, IZAS [2005-03]; 1♀ diss., Wei Lei Co. Farm 4 [near Yuli], 17-V-1960, Wang Shuyong, D. e. deserticola det. Chen, PARATYPE, IZAS [2005-02]; 1♂ diss., Wei Lei Co. Farm 4 [near Yuli], 17-V-1960, Zhang Facai, D. e. deserticola det. Chen, PARATYPE, IZAS [2005-01]; IRAN: 3♂♂ diss., 4♀♀ diss., 1♂, 3♀♀, 124 specimens [ NMPC], Abaregh [Abareq; 29.31667°N, 57.91667°E; irrigation channel in semi-desert; T. leptopetela (= T. kotschyi ) present ( Hoberlandt 1981)], 53 km northwest Bam, 25-IV-1973, Exp. Nat. Mus. Praha Loc. No. 178, NMPC [2004-30, 31, 40, 2005-04, 05], USNM [2003-06, 2004-04]; 1♂ diss., 1♀ diss., Bareng [Berang] to Nasratabad [Nosratabad] [app. location: 30.23670°N, 61.00170°E], Sistan, 11-V-1898, N. Zarudnyl, ZIN [2004-18, 21]; 1♂ diss., Bendun [Bandan] [to] Nayzar [Naizar marsh] [app. location: 31.3033°N, 60.8292°E], Sistan, east Persia, 8-V-1898, N. Zarudnyl, 17585, ZIN [2005-05]; 2♂♂ diss., 5♂♂, Kashan, 25 km toward Natanz [Natenz] [33.81444°N, 51.74361°E], 3,200 m elev. 2-V-2000, J. Gaskin, on Tamarix sp. , #772, GSWRL [2003-06, 37]; 1♂ diss., 1♀, Khatunabad [29.46667°N, 57.80000°E; stony semidesert; Tamarix sp. not recorded ( Hoberlandt 1981)], 70 km northwest Bam, 25-IV-1973, Exp. Nat. Mus. Praha Loc. No. 179, NMPC [2004-35b]; 1♂ diss., Khorassan [Ostan-e Khorasan Province; on northeast border of Iran; specific locality not given], 12-13-VIII-1901, N Zarudnyl, ZIN [2004-01]; 1♂ diss., 1♀, Minu Dasht, 60 km east [37.39°N, 55.85°E], MRSW Park [Mohammed Reza Shah Wildlife Park (= Golestan Biosphere Reserve)], 28-VIII-1972, [on T. ramosissima (Gerling and Kugler 1973) ], D. elongata det. S.L. Shute 1973, P27-29, TAUI [2003-03]; 1♂ diss., 1♀ diss., Shakhrud K. [Sharud Kola; 36.41670°N, 52.85000°E], north of Gorgan [southeast of Astrabad], Christof, ZIN [2004-17, 20]; KAZAKHSTAN: 1♂ diss., 2♂♂, 2♀♀, Chilik [Shelek], near [2♂♂, 2♀♀], or 25 km east [43.6715°N, 78.5050°E], Buryndysu environs [1♂ diss.], 6-VI-1999, Mityaev & Jashenko, on T. ramosissima , voucher shipment GSWRL-1999-8, GSWRL [2004-03] [introduced near Delta, Utah in 1999] [28 specimens from Lot GSWRL-1999-1 identified as D. elongata by A. Konstantinov on 20-VII-1999 at USDA Systematic Entomology Laboratory ( SEL), SEL Lot # 9904498]; 1♂ diss., 4♂♂, 5♀♀, Bzandy-Su [Buryndysu] Campsite, 3-4 km north [43.70650°N, 78.68490°E], 186 km east northeast Almaty, 99,921 km odom., 4-VI-1995, C.J. DeLoach, on T. ramosissima , D. e. deserticola det I. Lopatin 2000 [2♂♂, 1♀], 8989–8891, GSWRL [2003-56]; 1♂ diss., 9♂♂, 6♀♀, Chilik [Shelek; 43.60030°N, 78.29630°E], near, 106 east of Almaty, 8-VIII-1995, I.D. Mityaev, on T. ramosissima , D. elongata det. I.D. Mityaev [1995; 3♂♂, 4♀♀], voucher shipment GSWRL( CJD)-1995-23, GSWRL [2005-91]; 1♂ diss., Chilik, near, 4-VI-1997, I.D. Mityaev and P.E. Boldt, on T. sp. prob. ramosissima , voucher shipment GSWRL( CJD)-1997-13, GSWRL [2005-92]; 1♂ diss., 5♂♂, 8♀♀, Chilik [Shelek], near, 1-VIII-1999, I. Mityaev & R. Jashenko, on T. ramosissima , voucher shipment GSWRL-1999- 15, GSWRL [2005-98]; 1♂ diss., 3♂♂, 1♀, Chilik [Shelek], environs, 21-V-2000, Mityaev & Jashenko, on T. ramosissima , voucher shipment GSWRL-2000-9, GSWRL [2005-80, 97] [used in biological studies by DeLoach et al. 2003b and Lewis et al. (2003a)]; 2♂♂ diss., 8♂♂, 8♀♀, Chilik [Shelek], 7 km before [southwest; 43.49000°N, 78.25000°E], 130 km east Almaty, site 1, 99,865 km odom., 3-VI-1995, C.J. DeLoach, on T. ramosissima , D. e. deserticola det. I. Lopatin 2000 [6♂♂, 2♀♀], 8595–8664, 8985, 8987, 8988, voucher shipment GSWRL( CJD)-1995-11, GSWRL [2003-45, 2005-14] [10 specimens from Lot GSWRL( CJD)-1995-9 identified as prob. D. e. deserticola by R.E. White on 15-XI-1995 at USDA SEL, SEL Lot # 95-10233]; 2♂♂ diss., 2♂♂, 5♀♀, Chilik [Shelek], 38 km east [Buryndysu; 43.69710°N, 78.66300°E], 15-VI-1996, P.E. Boldt, on T. ramosissima , D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 2♀♀], 9013 [1♂ diss.], 9015, 9016, voucher shipment GSWRL( PEB)-1996-9, GSWRL [2003-83, 2005-23]; 1♂ diss., Chilik [Shelek], 23 km east [odom. reading 9,465 km] and 38 km east [odom. reading 9,480 km; Buryndysu; not west Chilik as in some shipment records], 15-VI-1996, I.D. Mityaev & P.E. Boldt, on T. ramosissima , voucher shipment GSWRL( PEB)-1996-9, GSWRL [2005-99] [8 specimens from Lot GSWRL( CJD)-1996-4 identified as D. elongata by S.M. Clark on 28-X-1996 at West Virginia Dept of Agric.] [8 specimens (no. 9205) from Lot GSWRL( CJD)-1997-1 identified as D. e. prob. elongata by A. Konstantinov on 21-V-1997 at USDA SEL, SEL Lot # 9704023]; 2♂♂ diss., 5♀♀ diss., 3♂♂, 3♀♀, Chundzha, 40 km west [43.47035°N, 79.02773°E], desert near Boguty Mtns., 9-VI-1999, Mityaev & Jashenko, on T. ramosissima , voucher shipment GSWRL-1999-9, GSWRL [2003-62, 63, 64, 2005-02, 48, 73, 74] [subject of field studies by Mityaev and Jashenko (1999, 2007)]; 2♂♂ diss., 3♂♂, 3♀♀, Chundzha, 40 km west, desert near Boguty Mtns., 1-VIII-1999, I.D. Mityaev & R. Jashenko, on T. ramosissima , voucher shipment GSWRL-1999-15, GSWRL [2005-93, 95]; 1♂ diss., Dzhulek [Zhulek; 44.28330°N, 66.43330°E], Syrdar’indkaya Oblast, VII- 1910, Kozchanchikov, ZIN [2004-15]; 1♂ diss., Indersk [Ynderbor; 48.55000°N, 51.78330°E], Reitter, D. e. ab. sublineata, HNHM [2003-10]; 7♂♂ diss., 2♂♂, 1♀, Kerbulak [43.9311°N, 77.6019°E], ca. 150 km north northeast Almaty, Ili River Valley, 2-VI-1999, Mityaev & Jashenko, on Tamarix sp. , voucher shipment GSWRL-1999-7, GSWRL [2004-19; 2008-06, 07, 08, 09,10, 11] [subject of field studies by Mityaev and Jashenko (1999, 2007)]; 1♂ diss., Lavar [43.56860°N, 78.13360°E], 10 km west Chilik [Shelek], 21-V-2000, Mityaev & Jashenko, on T. ramosissima , voucher shipment GSWRL-2000-9, GSWRL [2004-21] [introduced near Delta, Utah in 2000]; 1♂ diss., 10♂♂, 3♀♀, Masaq [43.62028°N, 78.30556°E] [not Marak as in some shipment records and labels], near, Ili Desert, 4-VI-1997, I.D. Mityaev and P.E. Boldt, on Tamarix sp. , voucher shipment GSWRL( CJD)-1997-13 [1♂], GSWRL [2004-18]; 1♂ diss., Sarytogay, Charyn Canyon [43.43240°N, 79.15040°E], 13-VI-1998, Mityaev, on Tamarix sp. , voucher shipment GSWRL-1998-10, GSWRL [2004-20]; 1♂ diss., Shelek, near, Shelek River, 6-VI-1999, Mityaev & Jashenko, on T. ramosissima , voucher shipment GSWRL-1999-8, GSWRL [2004-02] [introduced near Delta, Utah in 1999] [used in biological studies of DeLoach et al. (2003b) and Lewis et al. (2003b); subject of field studies by Mityaev and Jashenko (1999, 2007)]; KYRGYZSTAN: 1♂ diss., 1♀ diss., Dostuk [41.36972°N, 75.63667°E], environs, At–Bash River bed, Naryn Region, 20-VIII-1995, S. Saluk, USNM [2003-14, 16]; 1♂ diss., Naryn, 80 km west [41.36972°N, 75.3799°E], Central Tian Shan Mts., 3-VII-1966, E. Gurieva, ZIN [2004-14]; MONGOLIA: 1♂ diss., Mongolia [specific locality not given], Potanin, Coll[ection] Weise, [D.] e. var. sublineata [1♂ diss.], ZMHB [2006-05] [probably from series collected by G.N. Potanin from 22-VII to 3- VIII-1886 from Central Mongolia listed as D. e. var. sublineata by Weise (1890)]; 1♂ diss. 1♀ diss., southwestern Mongolia [specific locality not given], 17-VI [1♂ diss.], 18-VI [1♀ diss.] [circa 1928 ( Hedin 1943)], Söderbom, D. elongata [1♂ diss.], Sven Hedins Expedition Central Asia, NHRS [2003-02, 03]; Bayanhongor Aymag: 1♂ diss., Tooroyn Bulag spring [42.76745°N, 98.96116°E], 13 km east Tsagan Bulag, Bayanhongor Aymag, 16-VIII-1969, Zaitsev, ZIN [2004-16]; Hovd Aymag: 1♂ diss., Baruun Hurray [Barun Khurai Tract; 45.66667°N, 91.66667°E], 50 km southwest of Uench [Uyonch], 3-4-VIII-1976, L.N. Medvedev, D. elongata det. L. Medvedev, MRSN [2003-01] [from series listed as D. elongata by Medevedev and Voronova (1979)]; 2♂♂ diss., 2♀♀, Bulgan Somon [Burenhayrhan; 45.99394°N, 91.55267°E], 10 km south southwest, Chovd Aimak [Hovd Aymag], 4-5-VII-1966, Exp. Dr. Z. Kaszab, Nr. 628, D. elongata sublineata det. K. Lopatin [1♂ diss., EGRC], USNM [2003-08], EGRC [2005-05] [from series listed as D. e. sublineata by Lopatin (1968)]; 1♂ diss., Yelkhon, 20 km southeast [sic; should be southwest along Bodochiin Gol from Altai] Altai [Bor–Udzuur], on Bodonchiyn [Bodochiin Gol stream] [45.7008°N, 92.0836°E], 27- VII-1970, Emeljanov, ZIN [2004-05]; RUSSIA: 1♂ diss., 1♂, 1♀, Astrachan [Astrakhan; 46.34944°N, 48.04917°E], coll[ection] Koltze, 10883, DEI [2003-25]; 1♀ diss., signata [designated] Astrachan, Becker, coll[ection] L.V. Heyden, [ Galeruca ] carinulata Desbr., DEI [2003-04]; 1♀ diss., 2♂♂, 1♀, Astrachan, Coll[ection] Kraatz, [ Galeruca ] carinulata Desbr. [1♀ diss.], DEI [2005-11]; 1♂, 1♀, Astrakan [Astrakhan], Becker, D. e. ab. carinata det. Le Moult, IRSNB; 1♂ diss., 1♀, Astrakhan, Letzner, coll[ection], Gal [eruca] carinulata Desbr. [1♂ diss.], DEI [2003-03]; 1♂ diss., 1♂, 2♀♀, Astrakhan, Becker, coll[ection] Reitter, D. elongata, HNHM [2003-01]; 1♂ diss., 1♀, Astrakhan. [region; specific locality not given], Bekker, ZIN [2005-04]; 1♂ diss., Derbent [42.06278°N, 48.29583°E], Coll[ection] Koltze, 10884, DEI [2003-05]; 1♀ diss., Derbent, Becker, DEI [2004-01]; 1♂ diss., 1♂, 1♀, Malaya Areshkeva [44.02083°N, 46.82305°E], northern Daghestan [Respublika Dagestan], 31-V-1925, A.N. Kiritschenko, ZIN [2004-03]; 1♂ diss., 1♀ diss., Sarepta [48.52778°N, 44.48361°E], G. carinulata Desbrochers [1♂ diss.], Coll[ection] Stierlin, DEI [2003-07, 26]; 1♀ diss., Sarepta, Becker, [G.] carinulata, NHRS [2003-01]; 1♀ diss., Sarepta, D. Becker, ex. coll. J. Weise, ZMHB [2003-01]; 1♀ diss., Sarepta. Schioda [?], 896, Collectie C. et O. Vogt Acq. 1960, ZMAN [2008-13]; 2♂♂ diss., 1 ♀ diss., Ikrianoe [Ikryanoye; 46.09440°N, 47.73470°E] Village, Astrakhan region, 22–28-VIII- 2002, Osipov collection, GSWRL [2003-70, 2004-25, 26]; TAJIKISTAN: 1♂ diss., 1♀ diss., Vakhsh River, Tigrovaya Balka State Reserve [37.10528°N, 68.30389°E], Dzhilikul District, Kurgan–Tube [Kurgan Tyube] Region, 9-V-1992, Osipov collection, GSWRL [2003-77, 2004-08]; TURKMENISTAN: 2♂♂ diss., 2♀♀ diss., Ashgabat, 2-IX-2002, S.N. Myartseva, on T. ramosissima, GSWRL [2003-01, 03, 05, 2006-06]; 1♀ diss., Ashgabat, Dry Sport Lake, 11-V-1997, S. N. Myartseva, on Tamarix ramosissima [prob. T. aralensis J. Gaskin ], voucher shipment GSWRL( CJD)-1997-11, GSWRL [2005-101] [8 specimens (nos. 9202, 9203) from Lot GSWRL( CJD)-1997-1 identified as D. e. prob. elongata by A. Konstantinov on 21-V-1997 at USDA SEL, SEL Lot # 9704023]; 2♀♀ diss., Ashgabat [37.95°N, 58.36667°E], Dry Sport Lake, 11-VI-1997, S. N. Myartseva and P. Boldt, on T. ramosissima [prob. T. aralensis J. Gaskin ], voucher shipment GSWRL( CJD)- 1997-12, D. elongata det. I. Lopatin 1999 [1♀], 9023, GSWRL [2005-08, 2006-07]; 2♂♂ diss., Ashgabat, Dry Sport Lake, 26-IX-2002, S.N. Myartseva, on Tamarix spp. , GSWRL [2006-08, 09]; Ashgabat, near Quaragum Canal, 27-V-1994, C.J. DeLoach, on Tamarix sp. , D. e. deserticola det. I. Lopatin 2000, 6731, GSWRL [2005-03]; 1♀ diss., 2♂♂, Ashgabat, canal [Quaragum], 11-VI-1997, S.N. Myartseva and P. Boldt, on Tamarix sp. , D. elongata det. I. Lopatin 1999, 9017–9019, GSWRL [2005-21]; 3♂♂ diss., 4♀♀ diss., Ashgabat, Karakum [Quaragum] Canal, 12-IX-2002, S. N. Myartseva, on T. ramosissima, GSWRL [2003- 13b, A5, A6, A7, A8, 2005-49, 63]; UNKNOWN COUNTRY: 1♀ diss., Araxesthal [Aras River of Armenia, Azerbaijan and Iran], Reitter, HNHM [2003-11]; 1♂ diss., Caucasus [specific locality not given; possibly Armenia or Azerbaijan], Reitter. Leder., NMPC [2005-41]; 1♀ diss., 1♀, Kammenaja retschka [Kamennaya Rechka, Russia?], 4-VI-1909, Collectie C. et. O. Vogt Acq. 1960, ZMAN [2008-03]; 1♀ diss., Tscholokai [Tschkalow/Orenburg, Russia?], 31-V-1909, Collectie C. et. O. Vogt Acq. 1960, ZMAN [2008-02]; UNITED STATES OF AMERICA (introduced): Colorado: Pueblo Co.: 1♂ diss., 4♂♂, 7♀♀, Arkansas River, Pueblo Reservoir, 38.270°N, - 104.713°W, 7-IX-2005, D. Hosler and F. Nibling, on T. ramosissima , voucher [source: Fukang, China], GSWRL [2006-16]; Nevada: Churchill Co.: 1♂ diss., 5♂♂, 4♀♀, Carson Sink, Trinity Junction, 38.92°N, - 118.75°W, 23-IX-2005, T. Dudley, on T. ramosissima , voucher [source: Fukang, China], GSWRL [2006-12]; Mineral Co.: 1♂ diss., 4♂♂, 5♀♀, Walker Lake, 38.88°N, - 118.79°W, 21-IX-2005, T. Dudley, on T. ramosissima , voucher [source: Fukang, China], GSWRL [2006-13]; Pershing Co.: 1♂ diss., 3♂♂, 6♀♀, Humboldt Sink, 40.13°N, - 118.42°W, 23-IX-2005, T. Dudley, on T. ramosissima , voucher [source: Fukang, China], GSWRL [2006-14]; Utah: Millard Co.: 1♂ diss., 5♂♂, 6♀♀, Sevier River west of Delta, 39.172°N, - 112.903°W, 15-IX-2005, C.J. DeLoach, on T. ramosissima , voucher [source: Shelek, Lavar, and environs Buryndysu, Kazakhstan], GSWRL [2006-17]; Wyoming: Big Horn Co.: 1♂ diss., 4♂♂, 5♀♀, Big Horn Canyon National Recreation Area [44.857°N, - 108.208°W, east of Lovell], 22-VIII-2005, D. Kazmer, on T. ramosissima , voucher [source: Fukang, China], GSWRL [2006-15]; UZBEKISTAN: 2♂♂ diss., 11♂♂, 2♀♀, Buchara [Buxoro], southeast [39.7555°N, 64.4867°E], 3-IV-1999, R. Sobhian, on Tamarix spp. , D. elongata det. I.K. Lopatin 1999 [12♂♂, 2♀♀], 9036–9048, 9050, GSWRL [2003-82, 2008-20]; 2♂♂ diss., 1♀ diss., Buxoro, 5 km northwest [39.8228°N, 64.39134°E], along the road to Gazli, 27-IX-2002, R. Sobhian, on Tamarix, USDA lab [Buchara] colony at Albany, California, voucher (2002–2003, D. Bean), shipment EIWRU-2002-1010, GSWRL [2005-53, 54, 56]; 2♂♂ diss., 1♀ diss., 5♂♂, 2♀♀, Buchara [Buxoro], 10 km west [39.7614°N, 64.3011°E], 3-IV-1999, Kirk & Sobhian, on Tamarix sp. , D. elongata det. I.K. Lopatin 1999 [2♂♂ diss. 4♂♂], 9028–9031, 9033, 9034, GSWRL [2003-80, 2004-05, 2008-21]; 1♂ diss., 1♂, Bukhara [Buxoro], 25 km west [39.69778°N, 64.18315°E], 20-V-1994, Osipov Coll., GSWRL [2004-22]; 1♂ diss., 1♂, Karshi [Qarshi], 10 km west [38.92278°N, 65.73307°E], 7-IV-1999, A. Kirk & R. Sobhian, on Tamarix , D. elongata det. I.K. Lopatin 1999, 9065, 9066, GSWRL [2003-78]; 1♂ diss., Tashkent [Toshkent], 140 km south [approximate location: 40.29057°N, 68.91332°E], 10-IV-1999, Kirk & Sobhian, on Tamarix spp. , GSWRL [2004-17].

Distribution. General. The native distribution of D. carinulata ranges from Sarepta, Russia south to Iran and east to China and Mongolia (Map 5). Surveys by our colleague C. J. Deloach reveal that D. carinulata populations are temporally sporadic in China and Kazakhstan, with outbreaks in some years and rarity in other years. Further collections might also reveal D. carinulata in Armenia, Afghanistan, Pakistan and northwestern Iraq.

Confirmed Records. We dissected specimens (see Material examined) from Russia, the single country with a previous literature record for D. carinulata ( Desbrochers 1870) (Map 5).

New Records. We have dissected specimens from the following countries for which we find no previous specific reports of D. carinulata in the literature (Maps 5–6): Azerbaijan, Iran, Turkmenistan, Kazakhstan, Uzbekistan, Tajikistan, Kyrgyzstan, China, Mongolia, and the United States of America (Nevada, Utah, Colorado, Wyoming; introduced; Map 7, see Potential in Tamarisk Biological Control below for additional details). Past reports of D. elongata from Azerbaijan, Turkmenistan, Kazakhstan, Uzbekistan, and the United States should refer, at least in part, to D. carinulata . Past reports of D. e. deserticola from China, Mongolia, Kazakhstan, and the United States should also refer to D. carinulata (see synonymy above). Past reports of D. e. sublineata and D. e. ab. sublineata in China and Mongolia are also D. carinulata .

Unconfirmed Records. We find no country distribution records that remain unconfirmed. We consider reports of Diorhabda from Littlefield, Arizona (L.D. Walker, USDI Bureau of Land Management, St. George, Utah) that spread south from Utah as D. carinulata . All available locations from which specimens were dissected from China (25) and Mongolia (6) were D. carinulata , with the exception of specimens of D. carinata from China near the border of Kazkhstan in the Ili Kazakh Autonomous Prefecture and Yining. Therefore, we consider all literature records of D. elongata , D. e. sublineata and D. e. deserticola from these countries to be D. carinulata , although D. carinata may also be present at sites in far western Xinjiang Uygur Zizhiqu, China. From 11 locations east of 75° E in Kazakhstan, D. carinulata was dissected from ten locations; while, D. carinata was dissected from four locations, two of which were series greatly predominated by D. carinulata . Therefore, we consider all literature reports of D. elongata east of 75°E in Kazakhstan to include D. carinulata , but it is probable that D. carinata occurs in lower frequency at many of these same sites. Only D. carinulata was dissected from all five locations available from north of 42°N in southern Russia and we consider all reports of D. elongata from this area to be D. carinulata .

Below are 33 unconfirmed locality records that we consider as D. carinulata , but are listed as D. elongata in Russia, Kazakhstan and Mongolia, D. e. sublineata in Mongolia ( Lopatin 1970, 1977b), and D. e. deserticola in China (Map 5):

CHINA: Gansu Sheng: Anxi [Yuanquan; 40.500°N, 95.800°E], on Tamarix ; Dunhuang [40.16667°N, 94.68333°E], on Tamarix ; Nei Mongol Zhizhiqu: Dong He River [41.73647°N, 100.85793°E], on Tamarix (Bao Ping, Nanjing Agricultural University, Nanjing, China, pers. comm.); Ningxia Huizu Zizhiqu: Yanchi [37.7869°N, 107.3994°E], on Tamarix (B. Ping, pers. comm.); Xinjiang Uygur Zizhiqu: Akesu [Aksu; 41.1231°N, 80.2644°E]; Atushen [Atux; 36.7061°N, 76.1519°E]; Baicheng [41.7739°N, 81.8689°E] ( Chen 1961); Hami [42.800°N, 93.45000°E] (Sha and Yibulayin 1993); Hotan (Sha and Yibulayin 1993); K'u–ch'e [Kuqa; 41.7278°N, 82.93640°E]; Miquan [43.96667°N, 87.7000°E]; Shajingzi [40.76667°N, 79.96667°E]; Turtiaogou [geocoordinates not locatable] ( Chen 1961); Turpan [ D. carinulata dissected from same location], on Tamarix (Sha and Yibulayin 1993) ; Yecheng [Kargilik; 37.88500°N, 77.4131°E]; Wei–li [Yuli; 41.3306°N, 86.25780°E] ( Chen 1961); KAZAKHSTAN: Bakanas [Baqanas; 44.8081°N, 76.2772°E], Ili River, on Tamarix ( Ishkov 1996) ; Boguty [desert near Boguty Mts., 40 km west of Chundzha; D. carinulata dissected from same location], on Tamarix (Mityaev and Jashenko 1999, 2007); Borokhudzhir [43.9667°N, 79.5833°E], on Tamarix ( Kulenova 1968) ; Burundysu [Chilik, 38 km east; D. carinulata dissected from same location], on Tamarix ; Chilik [Shelek; D. carinulata dissected from same location], near, in Chilik River Valley, on Tamarix ; Kerbulak [ D. carinulata dissected from same location], Ili River Valley, on Tamarix (Mityaev and Jashenko 1999, 2007); Koksu [45.02780°N, 77.9460°E], on Tamarix ( Kulenova 1968) ; Lavar [ D. carinulata dissected from same location], on Tamarix (Mityaev and Jashenko 1999, 2007); Masak [Masaq; D. carinulata dissected from same location] Town environs, Chilik [Shelek] riverbed, 22-VI-1995, Myricaria sp. (Mityaev and Jashenko 1997, 2007); Sarytogay [ D. carinulata dissected from same location], Charyn Canyon, on Tamarix (Mityaev and Jashenko 1999, 2007); MONGOLIA: Bayanhongor Aymag: 2 specimens, Echin Gol Oasis [Ekhiin Gol, 43.30851°N, 98.99646°E], 90 km north of Capanbulag border post, 950 m, 27–28-VI- 1967 (Nr. 855), on Tamarix ( Lopatin 1970) ; Ezhiin Gola [Ekhiin Gol], 900 m, 3–4-IX-1976; Khatan–Sudlyn–Bulak [geocoordinates not locatable], 75 km west north west Tsagan–Bulak [Tsagaan Bulag spring], 1,450 m [elev.], 28-VII-1977 (Medvedev and Voronova 1979); Zxiln Gol oasis [geocoordinates not locatable], 150 km south Shine Dzhinsta [Dzalaa or Shinejinst], 900 m [elev.], 12–15-IX-1976 (Medvedev and Voronova 1977b); Talyn–Belgekh–Bulak spring [geocoordinates not locatable], 40 km south southeast of Ezhiin Gola [Ekhiin Gol], 1,250 m [elev.], 26-VII-1977 (Medvedev and Voronova 1979); Govi– Altay Aymag: Dzahuy [44.950°N, 96.600°E]; Khatan Khaïrkhan environs [Hatan Hayrhan Mt.], 50 km east southeast Altay [geocoordinates not locatable] (Medvedev and Voronova 1977b); Khatan Khayrkhav [Hatan Hayrhan Mt.], north slope, 1,225 m [elev.] [44.9881°N, 96.1792°E], 13–14-VII-1976; Khatan Khayrkhav [Hatan Hayrhan Mt.], 20 km southwest, 1,300 m [elev.] [44.8505°N, 96.0897°E], 15-VII-1976 (Medvedev and Voronova 1977a); Shara–Xulsni–Bulak spring [Shara Hulasanii Bulag; 43.3°N, 97.75°E], 100 km west Ezhiin Gola [Ekhiin Gol], 29-VII-1977; Hovd Aymag: Bodonchiyn Gol [Bodochiin Gol stream], 15 km southeast [sic; should be southwest along Bodochiin Gol from Altai] Altai [Bor–Udzuur] [45.7367°N, 92.1217°E], 4-VII-1976, on Myricaria alopecuroides (Medvedev and Voronova 1979) ; Jarantaj [geocoordinates not locatable], Wuste, 15-V-1974, R. Piechocki ( Lopatin 1977b); Uench Gol [Uyonch Gol stream], 25 km southwest Uencha [Uyonch], 1,300 m [elev.] [45.8861°N, 91.7807°E], 28-VII-1976 (Medvedev and Voronova 1977a); Omnogovi Aymag: Obooto Hural, 50 km west [43.02078°N, 100.90040°E], 1600 m [elev.], 25-VII-1977 (Medvedev and Voronova 1979); RUSSIA: Dosang [46.9044°N, 47.9111°E], Astrakhan Province, VI-2003, K.A. Grebennikov (with photograph, Berlov 2006).

From our habitat suitability index models (Maps 8b and 9a) and our preliminary species distribution models based on 10´climate grids (Tracy and DeLoach, unpublished data), we estimate northern Uzbekistan as much more suitable habitat for D. carinulata than D. carinata . Therefore, we consider reports of D. elongata along the Amu Darya (river) in northern Uzbekistan by Yakhontov and Davletshina (1955, 1959), Sinadsky (1963, 1968) and Khamraev (2003) to probably refer primarily to D. carinulata rather than D. carinata (Map 5). Consequently, we consider the following four unconfirmed distribution records of D. elongata as uncertain records of D. carinulata (Map 5):

UZBEKISTAN: Chertombayskoy Forest dachau [geocoordinates not locatable], Amu Darya Valley, heavy defoliation on Tamarix , VIII-1954; Nazarkhan [42.3333°N, 59.9833°E]; Nukus [42.8333°N, 59.4833°E]; Turtkul [41.5544°N, 61.0111°E] ( Sinadsky 1963, 1968).

Discussion. Taxonomy. Galeruca carinulata ( Desbrochers 1870) was described from Sarepta, Russia and incorrectly synonymyzed under D. elongata var. carinata by Weise (1893). Berti and Rapilly (1973) removed D. carinulata from synonymy with both D. elongata and D. carinata based on their investigation of the endophallus. However, as discussed under D. carinata (above), only Warchalowski (2003) has accepted this taxonomic change.

We dissected a single male topotype specimen (DEI 2003-07) from the type locality of D. carinulata in Sarepta, Russia, with endophalli matching that of Berti and Rapilly’s (1973) illustration of the endophallus of the holotype. This evidently old specimen (possibly from the late 1800’s) bore the following hand written label data: “Sarepta”; “ G. carinulata Desbrochers ”. A total of five males dissected from Astrakhan and Ikryanoye, Russia, ca. 365 km southeast of Sarepta (Map 5), also bore endophalli matching that of the description by Berti and Rapilly (1973), and three specimens of D. carinulata (of DEI) from Astrakhan also bore identification labels of G. carinulata Desbrochers. Three key characters of the endophallus of D. carinulata that distinguish it from other members of the D. elongata group can be seen in the illustrations of the endophallus of the holotype ( Fig. 19b–c of Berti and Rapilly 1973) and our figures ( Figs. 17 , 22 , 27 , 32 ): (1) the acutely rounded distal margin of the palmate endophallic sclerite; (2) elongate endophallic sclerite with blade extending for less than 2/3 the length of the sclerite; and (3) lack of hooked apex of the elongate sclerite. Desbrocher’s (1870) holotype male for G. carinulata was 5.5 mm in length, below that usually found for D. carinata (mean male length is 6.29 mm, minimum male size is 5.12 mm, Table 2) but well within the size range of the 6 males we dissected from Sarepta, Astrakhan, and Ikryanoye (4.98–5.85 mm). We are confident that these specimens form a single species conspecific with D. carinulata . We find additional distinguishing characters in the endophallic sclerites and female genitalia (vaginal palpi and internal sternite VIII) of D. carinulata from the vicinity of the type locality in southern Russia and throughout its range to western China that distinguish it from D. elongata , D. carinata and other members of the D. elongata group (see Male- Genitalia and Female- Genitalia above; Figs. 17 , 22 , 27 , 32 , 37 , 42 ; Map 5). The distinctive genitalic characters of D. carinulata are maintained in the same areas where D. carinata and D. elongata occur and near its abutting range boundary with D. meridionalis , and this is strong evidence for reproductive isolation between these species (see Biogeography below; Map 1, Table 8). Although D. sublineata is allopatric with D. carinulata , their status as reproductively isolated is supported by the number of diagnostic genitalic characters separating D. sublineata from D. carinulata (6 characters) being greater than the number of characters separating the moderately sympatric D. carinata from D. carinulata (5 characters) ( Table 5). Therefore, we firmly support Berti and Rapilly (1973) in restoring D. carinulata to species status, removing it from synonymy with both D. elongata and D. carinata . Habitus drawings of “ D. elongata ” adults from southern Russia ( Fig. 30 of Ogloblin 1936) and Eastern Europe ( Fig. 50 of Bieṅkowski 2004) are probably D. carinulata . Drawings of “ D. elongata ” larval structures from southern Russia ( Fig. 31 of Ogloblin 1936; Fig. 41.7 of Ogloblin and Medvedev 1971) probably also belong to D. carinulata .

Chen (1961) described D. e. deserticola from the Taklamakan Desert region of Xinjiang Autonomous Region, China. The type locality of D. e. deserticola is Yuli (= Wei–li) (Map 5) and 8 additional localities are listed for paratypes. Yu et al. (1996) elevated D. e. deserticola to species status as D. deserticola without comment, and this change is mostly overlooked in recent biological literature (e.g., Li et al. 2000; DeLoach et al. 2003b; Lewis et al. 2003 a, 2003b; Cossé et al. 2005; Milbrath and DeLoach 2006a), except by Meng et al. (2005) and Zhang and Baoping (2006). Lopatin et al. (2004) extended the range of D. e. deserticola to eastern Kazakhstan. We dissected two male and two female paratypes of D. e. deserticola collected from the type locality of Wei-lei, Xinjiang, China ( Figs. 22 , 27 , 32 —Wei Lei Ta) and all belong to D. carinulata NEW SYNONYMY. We identified as D. carinulata all specimens dissected from Chen’s original given distribution of D. e. deserticola , China and Mongolia (Map 5).

Chen’s description of D. e. deserticola gives ranges in body length of 5–6 mm for males and 5.5–7 mm for females, which is close to our observed size ranges of 4.6–6.1 mm for males and 5–7 mm for females ( Table 2). Lewis et al. (2003b) gives mean lengths for males (5.6 ± 0.2 mm) and females (5.9 ± 0.2 mm) that are slightly larger than we observed in specimens collected over the entire range of D. carinulata ( Table 2).The size range of 4.5–8 mm given for adult D. carinulata in regional keys of Mongolia (as D. e. ab. sublineata ; Medvedev and Voronova 1977b, Medvedev 1982) and China (as D. e. sublineata ; Gressitt and Kimoto 1963a) erroneously includes the larger size of other sibling species that can exceed 7 mm in length ( Table 2), such as D. carinata , but which we have not found in these areas (Map1). An excellent habitus drawing of D. carinulata (as D. deserticola ) from China is provided in Fig. 4-14 of Yu et al. (1996).

Laboissière (1935) described D. koltzei ab. basicornis Laboissière from Khotan (=Hotan), Turkestan (now in Xinjiang Uygur Zizhiqu, China) from a single female. Wilcox (1971) listed D. k. ab. basicornis as a synonym of D. koltzei . However, Ogloblin (1936) had earlier synonymized D. koltzei under D. rybakowi and ranked it as the aberration D. rybakowi ab. koltzei . We dissected the genitalia of Laboissière’s (1935) female type specimen (from ZMAN) and identified it as D. carinulata . We also studied 9 syntypes of D. koltzei (from DEI; label data: Ili [Ili Kazakh Autonomous Prefecture, Xinjiang Uygur Zizhiqu, China] [18]’97; Weise; Coll. Koltze; Syntype) and 7 identified specimens of D. rybakowi (6 from USNM and 1 from DEI), and we concur that D. koltzei is conspecific with D. rybakowi which is very distinct from D. carinulata . Therefore, we conclude that D. k. ab. basicornis is conspecific with D. carinulata NEW SYNONYMY.

The external characters provided by Berti and Rapilly (1973) to separate D. carinulata and D. carinata involve mainly the posterior angles of the pronotum and number of elytral carinae, and these characters are considered by Warchalowski (2003) as separating D. carinulata from D. elongata . However, we find these characters to be interspecifically variable and of no use in species diagnosis (see Discussion - Taxonomy under D. carinata ). Chen (1961) noted that the ventral tarsal pubescence was medially absent, leaving a glabrous median line under the tarsi; in D. carinulata (as D. e. deserticola ), but that the ventral tarsal pubescence is generally distributed in D. sublineata (as D. e. ab. sublineata ) from North Africa. We examined dissected specimens of D. sublineata from North Africa with the same pattern of ventral tarsal pubescence as Chen described for D. e. deserticola , making this character too variable for use in species separation. We have dissected specimens of D. carinulata that were misidentified by taxonomists using external diagnostic characters as D. elongata from six countries and as D. e. sublineata or D. e. var. sublineata from Mongolia (see Material examined).

Common Name. The vernacular name “striped tamarisk leaf beetle” has been applied to D. carinulata in Kazakhstan (as D. rybakowi ; Mityaev 1958, Mityaev and Jashenko 2007) and China (as D. e. deserticola ; Tian et al. 1988, Bao 1989, Sha and Yibulayin 1993). Because stripes can also be prominent in several other species of the D. elongata group (especially D. sublineata ), we have dropped the term “striped” from the name and adopt the name “northern tamarisk beetle”, referring to its northernmost range of all tamarisk beetles ( Fig. 51B , Map 1).

Biology. Host Plants. Sha and Yibulayin (1993) listed the following hosts of D. carinulata (as D. e. deserticola ) in order of preference based on numbers collected from the Turpan Eremophyte Botanical Garden in China: Tamarix laxa (Wildenow) , T. elongata (Ledebour) , T. kansuensis Zhang , T. gracilis Wildenow , T. androssowii Litvinov , T. arceuthoides , T. hispida , T. smyrnensis (as T. hohenackeri ), and T. chinensis ( Table 1). Mityaev and Jashenko (1998, 2007) reported T. leptostachya as a new host record from Kazakhstan among five other Tamarix spp. (see Table 1). The host range of D. carinulata (as D. e. deserticola ) was reviewed by DeLoach et al. (2003b) who reported T. hispida var. karelinii (Bunge) Baum as a new host. DeLoach et al. (2003b) found the highest field populations on T. ramosissima among a total of seven Tamarix spp. on which it was collected in western China (including T. leptostachya , which was omitted from DeLoach et al. 2003b) ( Table 1). Our collaborators R. Jashenko, I. Mityaev, and C. J. DeLoach collected D. carinulata from T. ramosissima at six locations and Tamarix sp. at three locations in Kazakhstan. Tamarix aralensis is a new host record for D. carinulata collected from Dry Sport Lake, Ashgabat, Turkmenistan by S. Myartseva (GSWRL collection). In addition to Tamarix spp. , D. carinulata is feeds upon Myricaria alopecuroides Schrenk ( Tamaricales : Tamaricaceae ) in Mongolia (as D. elongata ; Medvedev and Voronova 1979) and on one occasion was found to completely defoliate a few shrubs of Myricaria sp. by the Shelek River in southeastern Kazakhstan (as D. e. deserticola ; Mityaev and Jashenko 1997, 2007). Sinadsky (1968) reported what is probably D. carinulata (as D. elongata , see above discussion) producing heavy defoliation on T. ramosissima compared to light damage on T. hispida along the Amu Darya in Karakalpakia [Qoraqalpog`iston Respublikasi], northern Uzbekistan. Tamarix kotschyi Bunge (as T. leptopetela Bunge ) was recorded among the vegetation sampled at Abareq, Iran ( Hoberlandt 1981) in the collection of at least 142 specimens of D. carinulata that we examined. Later detailed vegetation surveys in this area map T. kotschyi as the only Tamarix species in the vicinity of Abareq ( Baum 1983), and we consider T. kotschyi a possible host for D. carinulata . Although T. aphylla is not a recorded host of D. carinulata , T. aphylla is common in the Registan-North Pakistan Sandy Desert in eastern Iran ( Browicz 1991), an area from which we dissected D. carinulata from two locations (Map 6). Populations of D. carinulata established in North America are vigorously defoliating T. ramosissima in Nevada, Utah, Wyoming and Colorado and T. chinensis in Colorado (DeLoach et al. 2004). Large populations in western Nevada have fed upon cultivated T. parviflora , although this host is less preferred than T. ramosissima ( Dudley et al. 2006, Dudley et al. in prep.) as Dalin et al. (in press) also observed in multiple-choice field cage studies in California. Tamarix parviflora is a new host record and novel host association in that it is not native where D. carinulata occurs in Asia. Attack on T. parviflora by D. carinulata is predictable from field cage studies in which T. parviflora did not significantly differ from most accessions of T. ramosissima / T. chinensis in terms of either suitability for larval survival or preference for adult oviposition (Milbrath and DeLoach 2006a).

DeLoach et al. (2003b) found that D. carinulata (as D. e. deserticola ) larvae from Turpan and Fukang, China and Shelek, Kazakhstan had highest survival on Tamarix followed by Myricaria and Frankenia . Adults oviposited less on bouquets of T. aphylla compared to T. ramosissima in the laboratory. Lewis et al. (2003a) found larval survival in the laboratory and adult oviposition in both laboratory and field cages on three North American Frankenia spp. ( F.salina , F. johnstonii , and F. jamesii ) generally were significantly lower than that on species of invasive North American Tamarix . But Milbrath and DeLoach (2006a) found that survival of larvae from Turpan was not different among the three North American Frankenia spp. and five invasive Tamarix spp. However, they confirmed previous results that adult oviposition was insignificant on the three Frankenia spp. in a combination of choice and no-choice field cage studies. Dudley and Kazmer (2005) planted alkali heath, Frankenia salina , at a site with large populations of D. carinulata defoliating tamarisk near Lovelock, Nevada. In this open field test, D. carinulata did not lay eggs on the Frankenia and larvae crawling from defoliated tamarisk trees produced <4% leaf damage to Frankenia . In field cage tests, D. carinulata oviposited significantly less on T. aphylla compared to other invasive North American tamarisks, including T. ramosissima , T. chinensis , T. canariensis / T. gallica ( Lewis et al. 2003a, Milbrath and DeLoach 2006a). Tamarix aphylla is at moderate risk of damage by D. carinulata in the field ( DeLoach et al. 2003b), but the degree of damage that D. carinulata might cause to T. aphylla , especially in the absence of other Tamarix spp , is difficult to predict. Frankenia is at very low risk of damage from D. carinulata ( Lewis et al. 2003a, Milbrath and DeLoach 2006a). Risk of damage to both T. aphylla and Frankenia by D. carinulata is probably much lower when these plants are not in the proximity of preferred Tamarix spp. (e.g., Blossey et al. 2001).

Ecology. Diorhabda carinulata (as D. e. deserticola ) is probably the most damaging specialized herbivore of tamarisk in Asia as a result of its abilities to: (1) defoliate large acreages of tamarisk, (2) reach high population densities, (3) produce several overlapping generations per season, (4) aggregate in large numbers, and (5) disperse widely from 2–6 km per day ( Ding 2004). Sporadic outbreaks of D. carinulata (as D. e. deserticola or D. elongata ) can defoliate 90% or more of the tamarisk over wide areas in Kazakhstan ( DeLoach et al. 2003b), Uzbekistan ( Sinadsky 1968), and China, where the beetles are controlled in order to protect tamarisk stands used in soil stabilization ( Bao 1989, Tian et al. 1988, Sha and Yibulayin 1993, Chen et al. 2000, Ding 2004, Peng et al. 2005, Li and Wang 2006). In contrast, D. carinulata (as D. elongata ) have been reported at population levels too low to damage tamarisk in the northern part of its range (ca. 45°N) in Mongolia (Medvedev and Voronova 1977b). Bao (1989) reported 66,666 ha of tamarisk colonized by D. carinulata (as D. e. deserticola ), along the Donge He River (Ping Bao, Alashan Range Extension Station, Nei Mongol Zizhiqu, China, pers. comm.) of Erjina County, Inner Mongolia, China (Map 5). Of the 66,666 ha of colonized tamarisk, Bao reported 40,000 ha, or 60%, was defoliated by D. carinulata in this area (a region known for extensive stands of T. ramosissima ; Kurschner 2004). Factors favoring periodic outbreaks of D. carinulata in Anxi County, Gansu Sheng, China, where up to 20,850 ha of tamarisk have been defoliated in a year, include (1) poor regulation from natural enemies, (2) mild winter temperatures increasing adult overwintering survival, (3) low precipitation during times of summer and fall pupation, reducing drowning of pupae on the ground, and (4) low water tables reducing mortality of ground dwelling pupae and overwintering adults ( Ding 2004). Diorhabda carinulata extensively defoliated tamarisk stands in a 26,300 ha area over a 259 km stretch of the Humboldt River basin by late 2005 near Lovelock, Nevada, where it was introduced in 2001 ( Carruthers et al. 2006, 2008). Large areas of tamarisk also have been defoliated near Lovell, Wyoming (D. Kazmer, pers. comm.), on the Colorado, Green, and Sevier rivers in Utah, and on the Dolores River in Colorado (D. Bean, pers. comm.). Smaller areas of tamarisk have been defoliated near Pueblo, Colorado (Map 7).

Tamarisk defoliation by D. carinulata in Kazakhstan is primarily from feeding of first generation larvae in early summer (Mityaev and Jashenko 1999, 2007). Third instar larvae consume much more tamarisk than earlier stages in Ningxia Province, China ( Tian et al. 1988). In Inner Mongolia, entire tamarisk trees are wilted from desiccation caused by larval feeding upon foliage and bark of tender branches, and tops of trees are discolored and wilted from aggregated feeding by adults ( Chen et al. 2000). Densities per tamarisk bush reached over 10,000 larvae, and averaged 1,000 adults (maximum 4,000) in Inner Mongolia ( Bao 1989). Near Turpan, China, average densities from 100 sampled tamarisk branches can reach 70 adults and 210 larvae per meter (Sha and Yibulayin 1993). Densities reach 270 adults and 534 larvae per meter of branch, and more than 7,000 larvae per individual tamarisk tree in Anxi County ( Ding 2004). Densities of 173 to 247 larvae per meter of branch result in near defoliation of tamarisk near Sarytogay, Kazakhstan (Mityaev and Jashenko 1999, 2007). In northern Uzbekistan, larvae crawl from defoliated trees over ground to seek other tamarisk ( Sinadsky 1968), a phenomenon that R. Carruthers and C.J. DeLoach also observed at Lovelock and at Schurz, Nevada, in 2004. Localized migrations of adult D. carinulata occur following emergence from overwintering, during mating of each generation, and prior to overwintering (Mityaev and Jashenko 1998, 2007).

Diorhabda carinulata (as D. elongata ) damages tamarisk over wide areas in all types of situations, including sand stabilization plantations and in the understory of turanga poplar ( Populus euphratica Olivier ), in northern Uzbekistan ( Sinadsky 1968). Diorhabda carinulata more heavily damages the succulent tamarisk of riparian areas compared to that of drier upland areas in China ( Bao 1989) and Kazakhstan (Mityaev and Jashenko 1998, 1999, 2007). Tamarisks growing in dense monoculture are attacked at a higher incidence compared to those in stands mixed with other shrubs such as Calligonum (Polygonaceae) and Haloxylon (Chenopodiaceae) near Turpan (Sha and Yibulayin 1993). Diorhabda carinulata defoliates tamarisk in the understory of plains cottonwood ( Populus deltoides subsp. monilifera ) at Pueblo, CO, and in extensive tamarisk monocultures at Lovelock, NV (C. Jack DeLoach, USDA/ARS, Temple, TX, pers. comm.).

Tamarisk severely defoliated by D. carinulata completely resprouts in a short time in Kazakhstan (Mityaev and Jashenko 1998, 2007; see DeLoach et al. 2003b, Fig. 2 ). In tamarisk defoliated in the Chertombayskoy area of the Amu Darya, Uzbekistan, bud break is delayed by about 15 days the following spring and some of these bushes exhibit die back in the tops ( Sinadsky 1968). Near Lovelock, Nevada regrowth also followed defoliation by D. carinulata , but it is usually accompanied by severe dieback and tree death is becoming more widespread in trees defoliated for several successive years ( Carruthers et al. 2006, 2008). Tamarisk dieback and death at this site is probably related to observed significant reductions in nonstructural carbohydrates in tamarisk root crowns following one to four years of defoliation by D. carinulata (as D. e. deserticola ) ( Hudgeons et al. 2007b).

Adult D. carinulata tend to aggregate in the field, and over 1000 adults were counted in a 2 x 2 m area in Inner Mongolia ( Bao 1989). Near Chilik Kazakhstan, newly emerged first generation beetles aggregate on certain bushes just prior to mating (Mityaev and Jashenko 1999, 2007). In populations originating from Fukang, China, a male produced aggregation pheromone was identified consisting of two components: (2 E,4 Z)-2,4-heptadienal and (2 E,4 Z)-2,4-heptadien-1-ol ( Cossé et al. 2005). This pheromone is being used to monitor populations near Lovelock, Nevada. A blend of green leaf volatiles is attractive for D. carinulata in the field, and the attraction is synergized when pheromone is added with the green leaf volatiles ( Cossé et al. 2006). Additional observations on biology, including mating behavior, are noted by Tian et al. (1988), Bao (1989), Mityaev and Jashenko (1998, 1999, 2007), Zhang (2002), and Zhang and Baoping (2006) in central Asia.

Phenology. Diorhabda carinulata has four generations from mid-April to mid-September at Turpan, China (Sha and Yibulayin 1993); three generations from April to October in northern Uzbekistan ( Sinadsky 1968); three generations in Fukang ( Li et al. 2000) and Ningxia province ( Tian et al. 1988), China; two to three generations in Inner Mongolia Province, China ( Bao 1989, Chen et al. 2000); and two generations in southern Kazakhstan (Mityaev and Jashenko 1998, 2007) and north of 38° in North America (DeLoach and Carruthers 2004b). Adults enter diapause from mid-August to September in China ( Bao 1989, Tian et al. 1988, Sha and Yibulayin 1993). In southeastern Kazakhstan, adults begin overwintering in mid-September, but, if moisture is favorable and saltcedar has new growth, larvae still can be seen feeding on plants into September and adults are found into early October (Mityaev and Jashenko 1998, 2007). The typical hibernaculum of second generation adults overwintering in Kazakhstan is the boundary layer between the soil and detritus under saltcedar trees (Mityaev and Jashenko 1998, 2007). In Ningxia Province, China, adults overwinter under dead leaves or soil in areas facing the sun and protected from wind ( Tian et al. 1988), At Lovelock, NV overwintering adults were found on the soil beneath the saltcedar litter during the fall and winter. Overwintered adults emerge in April in Ningxia Province ( Tian et al. 1988) and in late April when temperatures rise over 13°C in Inner Mongolia ( Bao 1989) and in May in southern Kazakhstan (Mityaev and Jashenko 1998, 2007).

The Fukang, China D. carinulata climatype did not initially establish at several sites in North America south of 38°N latitude, probably due to asynchrony of the critical photoperiod inducing diapause with the onset of cooler temperatures in these areas ( Bean et al. 2007a). In the south, these populations diapause prematurely in the summer, rather than the fall, leaving them inadequate food reserves for overwintering ( Lewis et al. 2003b).

Development and Reproduction. Diorhabda carinulata has three larval instars and a development time of 34 days from egg to adult at 24.1°C. Fecundity at 28.6°C on T. ramosissima averaged 194 eggs (range 78 to 550 eggs) with a population doubling time of 6.2 days. Fecundity on T. aphylla was lower than that found on T. ramosissima , T. canariensis and T. parviflora ( Lewis et al. 2003b) . Herrera et al. (2005) found 30–35°C is optimal among six constant temperatures from 15–40°C for highest survival and developmental rates for all life stages of D. carinulata . Both Zhang (2002) and Herrera et al. (2005) report temperature developmental thresholds and degree-days for development for various stages of D. carinulata .

Milbrath et al. (2007) found that, at 28°C, D. carinulata (as D. elongata from Turpan and Fukang, China) had a development time of 18.5–20.4 days from egg to adult (with 74–78% survival), a fecundity of 272– 283 eggs, and a population doubling time of 5.2–6.1 days. These values were all very similar to those found for D. elongata (Crete) , D. carinata ( Uzbekistan) , and D. sublineata ( Tunisia) (all as D. elongata ).

Natural Enemies. The parasitoid tachinid fly, Erynniopsis antennata , attacks larvae and emerges from adult D. carinulata from Fukang and Wujiaqu, Xinjiang Uygur Zizhiqu, China ( Zhang 2002). At quarantine in Temple, Texas, we commonly encountered the parasitoid E. antennata emerging from the bloated abdomens of killed overwintered adult D. carinulata that were originally collected alive in April at Fukang, China. Diorhabda carinulata from Fukang, represents both a new host and locality record for E. antennata . Unidentified mymarid egg parasitoids attack as much as 24% or more of D. carinulata eggs in the field near Fukang and Wujiaqu, China ( Zhang 2002, Zhang and Baoping 2006). An unidentified 1mm-long "fly", possibly an eulophid parasitoid hymenopteran, parasitized ca. 14.5–32.5% of the D. carinulata pupae in Inner Mongolia ( Bao 1989). The ubiquitous fungal pathogen Beauveria bassiana (Balsamo) was found infecting adult D. carinulata originating from the following locations (all identified by T. Poprawski): Fukang, China (shipment GSWRL(CJD)-1996-27); near Urumqi, China (shipment GSWRL-1998-12); and near Shelek, Kazakhstan (shipment GSWRL-2000-9).

At Turpan, China, the chief predators of D. carinulata are reduviids and mantids (Sha and Yibulayin 1993). Ants and coccinellids prey upon eggs and larvae of D. carinulata in Inner Mongolia ( Bao 1989). In southeastern Kazakhstan, nymphs and adults of the pentatomid Arma custos Fabricius , feed on larvae of D. carinulata (Mityaev and Jashenko 2000, 2007). Seven species of birds have been reported to feed upon adult D. carinulata . In Inner Mongolia, tree sparrows, Passer montanus (Linnaeus) , and common pheasants, Phasianus colchicus (Linnaeus) , were observed eating adult beetles, and the crop of one common pheasant hunted in late winter contained over 200 overwintering adults along with a few wheat seeds ( Bao 1989). Chen et al. (2000) also noted the crested lark, Galerida cristata (Linnaeus) , as a predator in Inner Mongolia. In Almaty, Kazakhstan, beetle adults were preyed upon by the great tit ( Parus major Linnaeus ), Eurasian blackbird ( Turdus merula Linnaeus ), common myna ( Acridotheres tristis [Linnaeus]), and house sparrow ( Passer domesticus [Linnaeus]) (Mityaev and Jashenko 2000, 2007).

Biogeography. Comparative. Diorhabda carinulata differs from other tamarisk beetles by the following combination of biogeographic characteristics: (1) primarily continental, usually beyond 1,400 km from the ocean at elevations above 400 m; (2) commonly found in temperate cold desert and grassland biomes; and (3) latitudinal range of 29– 49°N and most common at 40– 44°N. Diorhabda carinata and D. carinulata are moderately sympatric and syntopic over portions of their western range (see above discussion under D. carinata - Biogeography) (Map 1, Table 8), and they are the most similar species in the D. elongata group in terms of biomes inhabited ( Tables 9 and 10, Fig. 53 ). Diorhabda carinata differs from D. carinulata in being commonly collected in southern areas from 35– 42°N and in preferring temperate grasslands over deserts. Diorhabda carinulata is marginally sympatric with D. elongata in southern Russia (Dagestan Republic) and allopatric with D. sublineata (Map 1; Table 8). Diorhabda elongata and D. sublineata both differ from D. carinulata in being primarily maritime and most common in the Mediterranean biome. Diorhabda carinulata is parapatric with D. meridionalis in southern Iran. Both D. carinulata and D. meridionalis have a similar preference for deserts, but D. meridionalis differs in being maritime and most common further south at 26– 30°N (Maps 1 and 6; Figs. 51–52 ; Table 8).

Descriptive. Diorhabda carinulata occurs over the central portion of the Palearctic realm (Maps 1, 5 and 6). It has been most commonly collected between 40– 44°N in the Deserts and Xeric Shrublands and Temperate Grasslands and Shrublands biomes of eastern Central Asia. All reports of damage to tamarisk by D. carinulata are from this region of the Palearctic realm from elevations of ca. 30 m (at Turpan, China) to 970 m (Erjina County, Inner Mongolia, China). Diorhabda carinulata also occurs in Montane Grasslands and Shrublands biomes in western China around 44°N (from ca. 550–1850 m). From 35– 40°N it is usually found in the Deserts and Xeric Shrublands biome from 0–1,400 m elevation. From 36– 37°N, D. carinulata is occasionally found in the Temperate Conifer Forests biome in Iran (at ca. 1,400 m elevation). From 29– 34°N in Iran, collections are from two biomes: Montane Grasslands and Shrublands from ca. 1,625 –1,850 m elevation in the Kuh Rud and Eastern Iran Montane Woodlands ecoregion; and the Deserts and Xeric Shrublands biome from ca. 450–700 m elevation in the Registan-North Pakistan Sandy Desert (Map 6). The distribution of D. carinulata broadly coincides with that of the common host T. ramosissima from Mongolia and China to Iran and southern Russia, but its range appears to fall short of following the western distribution of T. ramosissima into Iraq and Turkey (Map 5).

Primary ecoregions for D. carinulata in the northern portion of its range (40– 46°N) in the Deserts and Xeric Shrublands include the Taklamakan Desert (30–1,600 m), Junggar Basin Semi-Desert (500–1,200 m), and Alashan Plateau Semi-Desert (900–1600 m) in China and Mongolia (Map 5) . The primary northern ecoregion of D. carinulata in the Temperate Grasslands and Shrublands biome is the Tian Shan Foothill Arid Steppe (500–1,850 m) in southeastern Kazakhstan and northern Tajikistan.

Potential in Tamarisk Biological Control. Summary. The northern tamarisk beetle is highly effective in biological control of T. ramosissima / T. chinensis in temperate cold deserts of the Great Basin Shrub Steppe, Colorado Plateau Shrublands and Wyoming Basin Shrub Steppe (Map 13). Diorhabda carinulata may be the most suitable tamarisk beetle for some warm temperate desert areas also, such as western portions of the Mojave Desert, southern portions of the Colorado Plateau Shrublands and Trans-Pecos Chihuahuan Desert (Map 13). Introduction of southern climatypes of D. carinulata from 29– 34°N in the Registan North Pakistan Sandy Desert (Map 6) may speed its adaptation to corresponding latitudes in North America.

Discussion. A northern D. carinulata climatype from ca. 44°N in China and Kazakhstan has been released and established in North America at several locations north of 37°N (Map 7). Diorhabda carinulata from Fukang, China, were introduced into cages in 1999, and released (under permit as D. elongata ) into the field in 2001 at four sites in Nevada, Colorado, and Wyoming for biological control of T. ramosissima / T. chinensis . Concurrently, populations from Shelek, and Lavar (10 km west Shelek), and Buryndysu environs (25 km east Shelek), Kazakhstan, were released and established in Utah (DeLoach et al. 2004) (Maps 7, 9). In 2005, the USDA-APHIS (2005) began a large-scale introduction program of D. carinulata (as D. e. deserticola ) from Fukang, China into 13 western states north of 38°N latitude. States involved in this program include Colorado, Idaho, Iowa, Kansas, Missouri, Montana, Nebraska, Nevada, North Dakota, Oregon, South Dakota, Washington and Wyoming. By 2005, it had defoliated over 20,000 ha of tamarisk in the Humboldt River basin near Lovelock, Nevada (Map 7). The northern tamarisk beetle has a moderate risk of damaging T. aphylla ( DeLoach et al. 2003b) and a low risk of damaging Frankenia ( Lewis et al. 2003a, Milbrath and DeLoach 2006a), and both these risks are probably much reduced at less proximity to preferred Tamarix spp. (e.g., Blossey et al. 2001).

The introduced northern D. carinulata climatype probably has the greatest potential to defoliate tamarisk in North America in the temperate cold Deserts and Xeric Shrublands and the Temperate Grasslands and Shrublands biomes from 40– 44°N (Map 13, see Biogeography). Elevations above 1,000 m might extend this defoliation further south in some ecoregions. Diorhabda carinulata has already defoliated large acreages of tamarisk at sites in three North American temperate cold desert ecoregions: the Great Basin Shrub Steppe (Lovelock, NV and Delta, UT), the Colorado Plateau Shrublands (Potash, UT), and the Wyoming Basin Shrub Steppe (Lovell, WY) (Map 13). The southern portion of the Snake/Columbia Shrub Steppe is probably also a highly suitable cold desert region (Map 13). The most suitable ecoregions of the Temperate Grasslands and Shrublands biome at 40– 44°N probably include the southern portion of the Northwestern Mixed Grasslands and the northern portions of the Western Short Grasslands (Map 13). Diorhabda carinulata has also established well at 38°N at Pueblo, Colorado in the Western Short Grasslands, but D. carinata may be better suited to the temperate grassland biome at this latitude (Map 13). The potential spread of the introduced parasitoid tachinid fly Erynniopsis antennata from California to Nevada should be monitored as it might severely reduce overwintering survival of northern tamarisk beetles.

The northern climatype of D. carinulata from Fukang, China was initially ill-adapted to areas south of 38°N (DeLoach et al. 2004) because of premature entry into diapause during short summer daylengths ( Lewis et al. 2003b; Bean et al. 2007a, 2007b). Overwintering of the northern Fukang climatype failed at field cages in Big Pine, California, Artesia, New Mexico, and Seymour, Temple, and Lake Thomas, Texas. The lower elevation D. carinulata northern climatype from Turpan, China failed to establish in cages at Lake Thomas, Texas (personal observation), and is establishing poorly at John Martin Reservoir, Colorado (Debra Eberts, USDI/Bureau of Reclamation, Denver, CO, pers. comm.) (Map 7). However, in 2008, populations originating from Delta, Utah, had established south to about 37°N on the Virgin River at Littlefield, Arizona (L.D. Walker, pers. comm.) and in Ute Canyon, Colorado (Dan Bean, pers. comm.). Natural expansion of the population was rapid in 2008, covering over 75 km south of Gateway along the Dolores River (D. Bean, pers. comm.). In 2006, a population of the northern Fukang climatype from Nevada began to increase and defoliated about 0.8 ha of tamarisk in the open field at Artesia, New Mexico (at ca. 33°N), but populations disappeared by 2008 (D. Thompson, pers. comm.). Apparently, the northern Fukang climatype may be gradually adapting further south. Putative climatypes from the extreme southern range limits of D. carinulata occur from 29– 34°N in widely varied habitats of southeastern Iran, such as the Registan-North Pakistan Sandy Desert (at ca. 450–700 m elev.) and Kuhrud Mountains (at ca. 1,625 –1,850 m elev.) (Map 6). In North America, these southern climatypes may more quickly adapt to areas in the southern half of the Colorado Plateau Shrublands, western portions of the Mojave Desert, Trans-Pecos Chihuahuan Desert, and southwestern portions of the Western Short Grasslands ecoregions. According to our HSI models, D. carinulata will probably not establish further south in subtropical deserts such as the Sonoran, Tamaulipan Mezquital and southern Chihuahuan (Map 13).

Initial attempts at establishing the northern Fukang D. carinulata climatype were unsuccessful north of 46°N at Fort Peck, Montana (D. Kazmer, pers. comm.) (Map 7). Putative climatypes of D. carinulata from the extreme northern limits of its native range occur from 46– 48°N at low elevations (ca. 0 m) in the Volga Valley of southern Russia and higher elevations (at ca. 1,200 m) of the Junggar Basin Semi-Desert in western Mongolia (Map 5). These extreme northern climatypes of D. carinulata may be better preadapted to Montana.

Figures

FIGURE 48. Scatter plots of length of blade of elongate endophallic sclerite (A) and length of spined area of elongate endophallic sclerite (B) versus the length of the elongate endophallic sclerite for field collected material of Diorhabda elongata species group and laboratory produced F1 and F2 D. elongata/D. sublineata hybrids (see Figs. 45–47 for illustrations of endophallic sclerites for individual hybrids identified by alphanumeric codes in plots; see Table 3 for sample sizes and statistics for measurements).

FIGURE 51. Schematic box plots for elevations (A) and latitudes (B) from native presence-only field collection data for the Diorhabda elongata species group at 5 minute grid resolution. Box plots for each species depict the mean, median, first quartile (Q1), third quartile (Q3), interquartile range (IQR, Q1–Q3), low whisker (LW; lowest point at or above 1.5*IQR lower than Q1), high whisker (HW, highest point at or below 1.5*IQR higher than Q3), mild outliers (MO, points between the low or high whisker and 3*IQR from Q1 and Q3, respectively), and extreme outliers (EO, points below or above 3*IQR from Q1 and Q3, respectively) (Proc Boxplot; SAS Institute 2005). Sample sizes of field localities (N) and a table summary statistics and plotted values are inset in each chart.

FIGURES 1–9. Dorsal habitus of males (dead - top row, living - bottom row) of the Diorhabda elongata species group (tamarisk beetles): 1–2—D. elongata (Mediterranean tamarisk beetle), 3–4—D. carinata (larger tamarisk beetle), 5–6—D. sublineata (subtropical tamarisk beetle), 7–8—D. carinulata (northern tamarisk beetle), 9—D. meridionalis (southern tamarisk beetle). SSV—subsutural elytral vitta, SMV—submarginal elytral vitta. Scale bar 5.0 mm.

FIGURES 14–18. Male genitalia: median lobe (ML) of aedeagus with everted endophallus (END; uninflated) (lateral and dorsal views).14—Diorhabda elongata, 15—D. carinata, 16—D. sublineata, 17—D. carinulata, 18—D. meridionalis. BF—basal foramen (or basal orifice), CES—connecting endophallic sclerite, DO—dorsal ostium (or apical orifice); EES—elongate endophallic sclerite, GP—gonopore, LA—lateral appendage, LB—length of blade, PES—palmate endophallic sclerite, SL—length of spined area of blade, VM—ventral membrane. Scale bar 1.0 mm.

FIGURES 19–23. Elongate (ventral) endophallic sclerite (lateral view) with connecting (lateral) endophallic sclerite (dorsal view, when present). 19—Diorhabda elongata, 20—D. carinata, 21—D. sublineata, 22—D. carinulata, 23—D. meridionalis. CES—connecting endophallic sclerite, EES—elongate endophallic sclerite, LA—lateral appendage, LB—length of blade, LN—lateral notch, SL—length of spined area of blade. Scale bar 1.0 mm.

FIGURES 24–28. Elongate (ventral) endophallic sclerite (dorsal views and a single dorso-lateral view [DLV]). 24—Diorhabda elongata, 25—D. carinata, 26—D. sublineata, 27—D. carinulata, 28—D. meridionalis. BL—blade, HA —hooked apex, CES—connecting endophallic sclerite (partial), EES—elongate endophallic sclerite, LA—lateral appendage, LB—length blade, LN—lateral notch. Scale bar 1.0 mm.

FIGURES 29–33. Palmate (dorsal) endophallic sclerite (dorsal view). 29—Diorhabda elongata, 30—D. carinata, 31—D. sublineata, 32—D. carinulata, 33—D. meridionalis. CES—connecting endophallic sclerite, DS—distal spines, EES—elongate endophallic sclerite, LA—lateral appendage, LN—lateral notch, PES—palmate endophallic sclerite, SDS—subdistal spines. Scale bar 1.0 mm.

FIGURES 34–38. Female genitalia: internal sternite VIII (IS VIII), vaginal palpi (VP), and spermatheca (SP). 34—Diorhabda elongata, 35—D. carinata, 36—D. sublineata, 37—D. carinulata, 38—D. meridionalis. AL—apical lobe of IS VIII, LP—length vaginal palpus, PA—pointed appendage of SP, ST—stalk of IS VIII, TL—tips of lobes of IS VIII, WAL—width apical lobe of IS VIII, WLS—width lobe of stalk of IS VIII, WP—width vaginal palpus, WST—width stalk of IS VIII. Scale bar 1.0 mm.

FIGURES 39–43. Female internal sternite VIII (IS VIII). 39—Diorhabda elongata, 40—D. carinata, 41—D. sublineata, 42—D. carinulata, 43—D. meridionalis. AL—apical lobe, ST—stalk, TL—tips of lobes of stalk, *—IS VIII distinctly identifiable as D. carinata, **—IS VIII distinctly identifiable as D. sublineata. Scale bar 1.0 mm.

FIGURES 49–50. Genitalic phenograms for the Diorhabda elongata species group. 49–dissimilarity phenogram based on average taxonomic dissimilarity matrix (Table 5), 50—similarity phenogram based on and Pearson product-moment correlation similarity matrix (Table 6). Phenograms produced with NTSYSpc Tree plot module (Rohlf 2006) from clusters formed with unweighted arithmetic average clustering (UPGMA) (NTSYSpc SAHN module).

FIGURES 53–54. Biomic dissimilarity dendrograms based on biomic Bray-Curtis dissimilarity matrices. 53—Diorhabda elongata species group (from Table 10), 54—both the D. elongata group and Tamarix species invasive in North America (from Table 12). Dendrograms produced with NTSYSpc Tree plot module (Rohlf 2006) from clusters formed with unweighted arithmetic average clustering (UPGMA) (NTSYSpc SAHN module). Line connecting D. elongata and T. gallica at left signifies that the positions of these taxa are interchangeable in an alternate dendrogram of equal rcoph value.