Cleridae, Latreille, 1802
publication ID |
https://doi.org/ 10.5281/zenodo.5178603 |
publication LSID |
lsid:zoobank.org:pub:4FA1390B-9417-47FC-A616-ACF93197FA3B |
persistent identifier |
https://treatment.plazi.org/id/03E87A62-FFDB-FFDE-FF39-F8D74FB7586F |
treatment provided by |
Felipe |
scientific name |
Cleridae |
status |
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Cleridae View in CoL View at ENA
Isoclerinae Kolibáč: Ababa Casey , sp. ( Brazil). Alimentary canal: As in previous species ( Fig 3 View Figures 1–12 ).
Hydnocerinae Spinola : Callimerus Gorham , sp. (South India). Alimentary canal: 6 cryptonephridial malpighian tubules. Reproductive organs: Female-Spermathecal capsule well sclerotized; spermathecal gland attached to middle of capsule; bursa copulatrix not saccular; ovariole comprising 6 follicles ( Fig. 4 View Figures 1–12 ).
Eurymetopum maculatum Blanchard. Reproductive organs: Male-One pair of accessory glands; seminal vesicle well developed; testis comprising 12 follicles ( Fig. 5 View Figures 1–12 ). Female-Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix not saccular ( Fig. 6 View Figures 1–12 ).
Eurymetopum Blanchard , species a-e ( Chile). Species a. Reproductive organs: Male-Two pairs of accessory glands, lateral pair bifurcated distally and much shorter than medial pair; testis comprising 6 follicles ( Fig. 7 View Figures 1–12 ). Species b. Reproductive organs: Male-Two pairs of accessory glands, lateral pair much longer than medial pair ( Fig. 8 View Figures 1–12 ). Species c-e. Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix not saccular; ovary comprising 6 follicles ( Figs. 9 – 11 View Figures 1–12 ).
Isohydnocera aegra (Newman) . Reproductive organs: Male-One pair of accessory glands, glands swollen at base and particularly long; testis comprising 6 follicles ( Fig. 12 View Figures 1–12 ). Female- Spermathecal capsule well sclerotized; spermathecal gland attached to apex of capsule; bursa copulatrix not saccular ( Fig. 13 View Figures 13–27 ).
Isohydnocera tabida (Leconte) . Reproductive organs: Male-As in previous species except testis comprising 12 follicles ( Fig. 14 View Figures 13–27 ). Female- As in previous species, except spermathecal gland notably shorter ( Fig. 15 View Figures 13–27 ).
Isohydnocera Chapin , species a (Arizona), species b ( México). Reproductive organs: Males-As in previous species except testis comprising 6 follicles. Species a ( Fig. 16 View Figures 13–27 ). Species b ( Fig. 17 View Figures 13–27 ).
Lemidia nitens (Newman) . Reproductive organs: Male- One pair of accessory glands; testis comprising 12 follicles ( Fig. 18 View Figures 13–27 ). Female- Spermathecal capsule well sclerotized; spermathecal gland attached near middle of capsule; bursa copulatrix not saccular ( Fig. 19 View Figures 13–27 ).
Lemidia Spinola , species a-g ( Australia). Reproductive organs: Species a. Male- One pair of accessory glands, glands diverticulated at base; testis comprising 12 follicles ( Fig. 20 View Figures 13–27 ). Female- As in previous species ( Fig. 25 View Figures 13–27 ). Species b. Male- One pair of accessory glands, glands particularly long; testis comprising 12 follicles ( Fig. 21 View Figures 13–27 ). Species c-g. Female- As in L. nitens ( Figs. 22, 23, 24, 26, 27 View Figures 13–27 ).
Phyllobaenus pallipennis (Say) . Reproductive organs: Male- One pair of accessory glands, glands trilobed; seminal vesicle bipartite; vas deferens particularly long; testis comprising 6 follicles ( Fig. 28 View Figures 28–35 ). Female- Spermathecal capsule well sclerotized; spermathecal gland attached to apex of capsule; bursa copulatrix not saccular ( Fig. 29 View Figures 28–35 ).
Phyllobaenus longus (Leconte) . Reproductive organs: Male- As in P. pallipennis except base of accessory gland highly diverticulated ( Fig. 30 View Figures 28–35 ). Female- As in P. pallipennis ( Fig. 31 View Figures 28–35 ).
Phyllobaenus antillae (Wolcott) . Reproductive organs: Male- As in Isohydnocera tabida except seminal vesicle bipartite and testis comprising 6 follicles ( Fig. 32 View Figures 28–35 ). Female- Female-As in P. pallipennis ( Fig. 33 View Figures 28–35 ).
Phyllobaenus Dejean , sp. ( Guatemala). Reproductive organs: Male- As in P. antillae ( Fig. 35 View Figures 28–35 ). Female- As in P. pallipennis ( Fig. 36 View Figures 36–44 ).
Wolcottia sobrina (Fall) . Reproductive organs: Male- One pair of accessory glands, glands particularly long, swollen at base; testis multifollicular ( Fig. 34 View Figures 28–35 ). Female- Spermathecal capsule well sclerotized, elongated; spermathecal gland attached to apex of capsule; bursa copulatrix not saccular ( Fig. 39 View Figures 36–44 ).
Clerinae Latreille : Aulicus femoralis Schaffer. Reproductive organs: Male- Two pairs of accessory glands, both glands biramous, medial pair transformed into spermatophoral gland ( Fig. 37 View Figures 36–44 ).
Aulicus nigriventris Schaeffer. Reproductive organs: Female- Spermathecal capsule not well sclerotized, bipartite; spermathecal gland attached to middle deflexed branch of capsule; bursa copulatrix not saccular; ovary comprising 13 follicles ( Fig. 38 View Figures 36–44 ).
Axina conspicua Schenkling. Alimentary canal: With 6 cryptonephridial malpighian tubules. Reproductive organs: Male- Two pairs of biramous accessory glands, medial gland much shorter than lateral glands, inner branch of medial gland with central longitudinal dark streak ( Fig. 40 View Figures 36–44 ). Female- Spermathecal capsule slightly sclerotized; spermathecal gland attached to middle of capsule; bursa copulatrix saccular ( Fig. 41 View Figures 36–44 ).
Axina bifasciata (Chevrolat) . Reproductive organs: Male. With two pairs of biramous accessory glands, medial pair much shorter than lateral pair, inner branch of medial branch with central longitudinal dark streak; testis comprising 44 follicles ( Fig. 42 View Figures 36–44 ).
Axina parcepunctata Schenkling. Reproductive organs. Females- Spermathecal capsule slightly sclerotized; spermathecal gland attached to middle of capsule ( Fig. 44 View Figures 36–44 ).
Axina plagiata Schenkling. Reproductive organs: Male- With 2 pairs of biramous accessory glands, inner branch of medial gland highly coiled distally and with central longitudinal dark streak; testis comprising 49 follicles ( Fig. 43 View Figures 36–44 ).
Caestron concinnus (Gorham) . Reproductive Organs: Male- Two pairs of accessory glands, both glands with basal diverticulum, inner chamber of medial gland with central longitudinal dark streak; testis comprising 11 follicles ( Fig. 45 View Figures 45–51 ). Female- Spermathecal capsule slightly sclerotized; spermathecal gland attached to basal region of capsule; spermathecal duct exceptionally long; bursa copulatrix saccular ( Fig. 46 View Figures 45–51 ).
Caestron contractus (Gorham) . Reproductive Organs: Female- As in previous species ( Fig. 47 View Figures 45–51 ). Notes: The extraordinarily long spermathecal duct may be apotypic for Caestron Spinola.
Calendyma chilensis (Castelnau) . Alimentary canal: Stomodaeum with well- developed proventriculus; ventricular papillae well developed; with 6 cryptonephridial malpighian tubules; proctodaeum well developed ( Fig. 48 View Figures 45–51 ). Reproductive organs: Female- Spermathecal capsule slightly sclerotized, capsule elongated and tapered; spermathecal gland attached to apex of capsule; spermathecal duct extraordinarily long; with large saccular bursa copulatrix ( Fig. 49 View Figures 45–51 ).
Cardiostychus gabonicus (Thomson). Alimentary canal: As in previous species ( Fig. 50 View Figures 45–51 ). Reproductive organs: Male- Two pairs of very long accessory glands, medial gland biramous, lateral gland uniramous; testis comprising 20 follicles ( Fig. 51 View Figures 45–51 ). Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 52 View Figures 52–59 ).
Cardiostychus Quedenfeldt, sp. a ( Democratic Republic of the Congo). Reproductive organs: Male- Two pairs of accessory glands, medial gland biramous, lateral gland not branched ( Fig. 53 View Figures 52–59 ). Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 54 View Figures 52–59 ).
Chiloclerus mimus Solervicens. Alimentary canal: Stomodaeum with well- developed proventriculus; ventricular papillae well developed; 6 cryptonephridial malpighian tubules; proctodaeum well developed ( Fig. 55 View Figures 52–59 ). Reproductive organs: Male- Two pairs of accessory glands, medial pair transformed into spermatophoral glands, lateral pair biramous; testis comprising more than 70 follicles ( Fig. 56 View Figures 52–59 ).
Clerus mutillarius (Fabricius) . Alimentary canal: Stomodaeum with well- developed proventriculus; ventricular papillae well developed; 6 cryptonephridial malpighian tubules; proctodaeum well developed ( Fig. 57 View Figures 52–59 ). Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, lateral pair biramous; testis comprising 26 follicles ( Fig. 59 View Figures 52–59 ). Female- Spermathecal gland well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 58 View Figures 52–59 ).
Eleale Newman , species a-h ( Australia). Reproductive organs. Species a. Male- Two pairs of accessory glands, medial pair not branched, diverticulated at base, lateral pair biramous; testis comprising 32 follicles ( Fig. 60 View Figures 60–72 ). Species b. Male- Two pairs of accessory glands, lateral pair not branched, medial pair extraordinarily long, biramous and diverticulated at base; testis small and spheroid ( Fig. 61 View Figures 60–72 ). Species c. Male- Two pairs of accessory glands, medial pair not branched, medial gland biramous; testis comprising 15 follicles ( Fig. 62 View Figures 60–72 ). Female- Spermathecal gland well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix not saccular ( Fig. 67 View Figures 60–72 ). Species d. Male- Two pairs of accessory glands, medial pair not branched, diverticulated at base, lateral glands biramous ( Fig. 63 View Figures 60–72 ). Female- Spermathecal gland well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix not saccular ( Fig. 66 View Figures 60–72 ). Species e. Male- Two pairs of accessory glands, medial pair not branched, diverticulated at base, lateral pair biramous; testis multifollicular ( Fig. 64 View Figures 60–72 ). Species f-h. Females- Spermathecal capsule well sclerotized; bursa copulatrix not saccular (sp. f, Fig. 65 View Figures 60–72 ; sp. g, Fig. 68 View Figures 60–72 ; sp. h, 69).
Epiclines basalis Blanchard. Reproductive organs: Male- Two pairs of accessory glands, medial gland not branched, highly diverticulated at base, lateral gland biramous; testis multifollicular ( Figs. 70, 72 View Figures 60–72 ). Female- Spermathecal capsule well sclerotized, annulated; spermathecal gland attached near apex of capsule; bursa copulatrix saccular ( Fig. 71 View Figures 60–72 ).
Eunatalis Schenkling , species a-b ( Australia). Species a. Alimentary canal: Stomodaeum with well- developed proventriculus; ventricular papillae well developed; 6 cryptonephridial malpighian tubules; proctodaeum well developed ( Fig. 73 View Figures 73–83 ). Reproductive organs: Male- Two pairs of uniramous accessory glands; testis multifollicular ( Fig. 74 View Figures 73–83 ). Female- Spermathecal capsule feebly sclerotized; spermathecal gland attached to apex of capsule; bursa copulatrix saccular, long and slender ( Fig. 75 View Figures 73–83 ). Species b. Reproductive organs. Male- Two pairs of uniramous accessory glands ( Fig. 77 View Figures 73–83 ). Female- Spermathecal capsule feebly sclerotized, exceptionally elongated; spermathecal gland attached near middle of capsule; bursa copulatrix saccular, long and slender ( Fig. 76 View Figures 73–83 ). Notes: The long narrow condition of the bursa copulatrix may be taxonomically significant in Eunatalis Schenkling.
Metademius Schenkling , species a ( Australia). Reproductive organs: Female- Spermathecal capsule not well sclerotized; spermathecal gland attached near middle of capsule; bursa copulatrix saccular ( Fig. 78 View Figures 73–83 ).
Neoscrobiger patricius (Klug) . Alimentary canal: Stomodaeum short, with well- developed proventriculus; ventricular papillae well developed; 6 cryptonephridial malpighian tubules; proctodaeum well developed ( Fig. 83 View Figures 73–83 ). Reproductive organs: Male- Two pairs of accessory glands, medial gland with spheroid lobe at base, lateral pair not branched; testis multifollicular ( Fig. 81 View Figures 73–83 ). Female- Spermathecal capsule not well sclerotized, bifid; spermathecal gland attached to base of inner chamber; bursa copulatrix saccular and twisted at extremity ( Fig. 82 View Figures 73–83 ).
Omadius semicarinatus Chevrolat. Reproductive organs. Male- Two pairs of accessory glands, medial pair not branched, lateral pair with basal vesicle; testis multifollicular ( Fig. 79 View Figures 73–83 ). Female- Spermathecal capsule well sclerotized, sinuous; spermathecal gland attached to base of capsule ( Fig. 80 View Figures 73–83 ).
Omadius Castelnau , species a-p (New Guinea). Species a. Reproductive organs: Two pairs of accessory glands, medial pair biramous, much longer than biramous lateral gland; testis multifollicular ( Fig. 84 View Figures 84–95 ). Species b. Male- Two pairs of accessory glands, medial pair biramous, much longer than uniramous lateral gland; testis comprising 12 follicles ( Fig. 85 View Figures 84–95 ). Species c. Male- Two pairs of accessory glands, medial pair biramous, dorsal branch very short, lateral glands uniramous ( Fig. 86 View Figures 84–95 ). Species d. Male- Two pairs of accessory glands, medial pair biramous and much longer than uniramous lateral pair; testis comprising 18 follicles ( Fig. 87 View Figures 84–95 ). Species e. Male- Two pairs of accessory glands, medial pair biramous and much longer than uniramous lateral pair; testis comprising 12 follicles ( Fig. 88 View Figures 84–95 ). Species f. Male- Two pairs of accessory glands, medial pair biramous; testis comprising 8 follicles ( Fig. 89 View Figures 84–95 ). Species g-i. Males- Two pairs of accessory glands, medial pair biramous and much longer than uniramous lateral pair; testis comprising 12 follicles (species g Fig. 90 View Figures 84–95 , species i Fig. 92 View Figures 84–95 ). Species h. Male- Testis comprising 7 follicles ( Fig. 91 View Figures 84–95 ). Female-Spermathecal capsule well sclerotized; spermathecal gland attached near apex of capsule; bursa copulatrix not saccular ( Fig. 95 View Figures 84–95 ). Species j. Male- Two pairs of uniramous accessory glands, lateral pair vesicular; testis comprising 12 follicles ( Fig. 93 View Figures 84–95 ). Female- Spermathecal capsule well sclerotized, elongated; spermathecal gland attached to middle of capsule; bursa copulatrix not saccular ( Fig. 94 View Figures 84–95 ). Species k. Female- Spermathecal capsule well sclerotized; spermathecal gland attached to middle of capsule; bursa copulatrix highly reduced ( Fig. 96 View Figures 96–105 ). Species l-o. Females- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular (species l Fig. 97 View Figures 96–105 , species m Fig. 98 View Figures 96–105 , species n Fig. 99 View Figures 96–105 , species o Fig. 100 View Figures 96–105 ). Species p. Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix highly reduced ( Fig. 101 View Figures 96–105 ).
Opilo taeniatus (Klug) . Reproductive organs: Male- Two pairs of biramous accessory glands, inner branch of medial glands with central longitudinal dark streak; testis comprising 11 follicles ( Figs. 102, 104, 105 View Figures 96–105 ). Female- Spermathecal capsule bifid, not well sclerotized; spermathecal gland attached at middle of secondary chamber; bursa copulatrix not saccular ( Fig. 103 View Figures 96–105 ).
Phloeocopus Spinola , species. ( Democratic Republic of the Congo). Alimentary canal: Stomodaeum with well-developed proventriculus; ventricular papillae well developed; 6 cryptonephridial malpighian tubules ( Fig. 106 View Figures 106–112 ). Reproductive organs: Male- Two pairs of biramous accessory glands, inner branch of medial pair with central longitudinal dark streak; testis multifollicular ( Fig. 107 View Figures 106–112 ). Female- Spermathecal capsule feebly sclerotized; spermathecal gland particularly long and attached to middle of capsule; bursa copulatrix saccular and small, spinous plates near bursa and median oviduct ( Figs. 108, 109 View Figures 106–112 ).
Phlogistus imperialis (Gorham) . Reproductive organs: Female- Spermathecal gland bifid, not well sclerotized; spermathecal gland attached to secondary chamber; bursa copulatrix not saccular ( Fig. 110 View Figures 106–112 ).
Priocera castanea (Newman) . Reproductive organs: Male- Two pairs of accessory glands, medial gland biramous, lateral gland vesicular; testis multifollicular ( Fig. 111 View Figures 106–112 ). Female- Spermathecal capsule partially sclerotized; spermathecal gland attached to apex of capsule; bursa copulatrix saccular ( Fig. 112 View Figures 106–112 ).
Priocera hypocrita Chevrolat. Reproductive organs: Male- medial glands biramous, lateral glands vesicular, round; testis comprising 25 follicles ( Fig. 113 View Figures 113–122 ). Female- Spermathecal capsule partially sclerotized; spermathecal gland attached to apex of capsule; bursa copulatrix saccular ( Fig. 114 View Figures 113–122 ).
Priocera variegata Kirby. Reproductive organs: Female- Spermathecal capsule partially sclerotized, annulated; spermathecal gland attached to apex of capsule; bursa copulatrix not saccular ( Fig. 116 View Figures 113–122 ).
Priocera Kirby , species. ( Brazil). Alimentary canal: Ventricular papillae well developed; 6 cryptonephridial malpighian tubules. Reproductive organs: Female- Spermathecal capsule partially sclerotized; spermathecal gland attached to apex of capsule; bursa copulatrix saccular ( Fig. 115 View Figures 113–122 ). Notes. The condition of having the spermathecal capsule partially sclerotized may have phylogenetic significance in the genus. There is some question regarding the extent of sclerotization of the capsule in P. variegata .
Stigmatium gilberti White. Reproductive organs: Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; spermathecal duct with small papilla at base; bursa copulatrix saccular ( Fig. 118 View Figures 113–122 ).
Stigmatium Gray , species ( Australia). Reproductive organs: Female-As in previous species, except spermathecal duct without basal papilla ( Fig. 117 View Figures 113–122 ).
Thanasimus dubius (Fabricius) . Reproductive organs: Male- Two pairs of accessory glands, medial pair biramous, lateral pair not branched; testis comprising 12 follicles ( Fig. 119 View Figures 113–122 ). Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix reduced to two small chambers; ovary comprising 12 follicles ( Fig. 120 View Figures 113–122 ).
Thanasimus formicarius Linnaeus. Reproductive organs: Male- Two pairs of accessory glands, medial pair biramous, lateral pair not branched; testis comprising 12 follicles ( Fig. 121, 122 View Figures 113–122 ). Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix reduced to two small chambers ( Fig. 123 View Figures 123–131 ).
Thanasimus ceylonicus Kraatz. Reproductive organs: Male- Two pairs of accessory glands, both glands biramous; testis comprising 12 follicles ( Figs. 124, 125 View Figures 123–131 ).
Thanasimus rufipes (Brahm) . Reproductive organs: Male- Two pairs of accessory glands, medial pair with small basal branch ( Figs. 126, 127 View Figures 123–131 ).
Notes: The characteristic of having the bursa copulatrix modified into two small chambers may have phylogenetic significance within the genus.
Trichodes alvearius (Fabricius) . Reproductive organs: Male- Two pairs of biramous accessory glands, medial pair short and transformed into spermatophoral gland, lateral pair very long; testis multifollicular ( Fig. 128 View Figures 123–131 ). Female- Spermathecal capsule bifid, not well sclerotized; spermathecal gland attached to secondary chamber; bursa copulatrix modified into spermatophoral chamber-note spermatophore Fig. 129 View Figures 123–131 - ( Fig. 130 View Figures 123–131 ).
Trichodes apiarius (Linnaeus) . Reproductive organs: Male- As in previous species ( Fig. 131 View Figures 123–131 ). Female- As in previous species ( Fig. 132 View Figures 132–137 ).
Trichodes nutalli (Kirby) . Reproductive organs: Female- As in previous species except spermathecal gland shorter ( Fig. 133 View Figures 132–137 ). Notes: I examined the reproductive organs of the following Trichodes Herbst species and found essentially the same design in structure as those I described in the abovementioned species: T. bibalteatus Leconte , T. crabroniformis (Fabricius) , T. favarius Illiger , T. flavocinctus Spinola , T. horni Wolcott , T. leucopsideus (Olivier) , T. octopunctatus (Fabricius) , T. oregonensis Barr, T. oresterus Wolcott, T. punctatus Fischer , T. quadriguttatus Adams , and T. umbellatarum (Olivier) . Notes: Opitz (2003) found the modification of male accessory glands into spermatophoral glands and the bifurcation of the spermathecal capsule phylogenetically significant among various Nearctic and Palaearctic genera.
Trogodendron fasciculatum (Schreibers) . Alimentary canal: Stomodaeum with well-developed proventriculus; ventricular papillae well developed, 6 cryptonephridial malpighian tubules; proctodaeum well developed ( Fig. 136 View Figures 132–137 ). Reproductive organs: Male- Two pairs of accessory glands, medial glands much longer than lateral glands; testis multifollicular ( Figs. 134, 135 View Figures 132–137 ). Female- Spermathecal capsule bifid, poorly sclerotized; spermathecal gland attached to secondary chamber; bursa copulatrix not saccular ( Fig. 137 View Figures 132–137 ).
Tillinae Leach : Callotillus elegans Erichson. Reproductive organs: Male- Two pairs of accessory glands, medial pair uniramous and with central longitudinal dark streak, lateral pair diverticulated at base; testis comprising 1 follicle ( Fig. 138 View Figures 138–146 ).
Cladiscus obeliscus Lewis. Reproductive organs: Male Two pairs of accessory glands, medial pair lobed basally not branched and with central longitudinal dark streak, lateral branch biramous; testis comprising 6 follicles ( Fig. 139 View Figures 138–146 ).
Cladiscus sanguinicollis (Spinola) . Reproductive organs: Female- Spermathecal capsule well sclerotized; spermathecal gland short, attached to apex of capsule; bursa copulatrix saccular; ovary comprising 6 follicles ( Fig. 140 View Figures 138–146 ).
Cladiscus Chevrolat , species ( Japan). Reproductive organs: Female- Spermathecal capsule well sclerotized, wrinkled; spermathecal gland attached to apex of capsule; bursa copulatrix saccular ( Fig. 141 View Figures 138–146 ).
Cymatoderella collaris (Spinola) . Alimentary canal: Stomodaeum with proventriculus not well defined externally; ventricular papillae well developed; 6 cryptonephridial malpighian tubules; proctodaeum well developed ( Fig. 145 View Figures 138–146 ). Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, lateral pair biramous; testis comprising 1 follicle ( Fig. 142 View Figures 138–146 ).
Cylidrus abdominalis Klug. Alimentary canal: Stomodaeum long, with proventriculus not well defined externally; ventricular papillae well defined; 4 cryptonephridial malpighian tubules; proctodaeum well developed ( Fig. 146 View Figures 138–146 ). Reproductive organs: Male- One pair of accessory glands; vas deferens particularly elongated; testis comprising 6 follicles ( Fig. 143 View Figures 138–146 ). Female- Spermathecal capsule bifid, not well sclerotized; spermathecal gland attached to secondary chamber; bursa copulatrix not saccular ( Fig. 144 View Figures 138–146 ).
Cymatodera aegra Wolcott. Reproductive organs: Female- Spermathecal capsule bifid, curvate; spermathecal gland attached to base of capsule; bursa copulatrix not saccular, bursal plate as in Fig. 148 View Figures 147–162 ( Fig. 147 View Figures 147–162 ).
Cymatodera undata Spinola. Reproductive organs: Male- Two pairs of accessory glands, medial gland not branched, with central longitudinal dark streak, lateral pair branched; testis comprising 1 very long follicle ( Fig. 149 View Figures 147–162 ). Female- Spermathecal capsule slightly bifid; spermathecal gland attached to base of capsule; bursa copulatrix not saccular; bursal plate as in Fig. 150 View Figures 147–162 ( Fig. 151 View Figures 147–162 ).
Cymatodera antennata Schaeffer. Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral pair branched; testis comprising 1 follicle ( Fig. 154 View Figures 147–162 ). Female- Spermathecal gland partially divided; spermathecal gland attached to base of capsule; bursa copulatrix not saccular; bursal plate as in Fig. 152 View Figures 147–162 ( Fig. 153 View Figures 147–162 ).
Cymatodera bicolor (Say) . Reproductive organs: Female- Spermathecal capsule bifid, poorly sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix not saccular; bursal plate present ( Fig. 158 View Figures 147–162 ).
Cymatodera californica Horn. Reproductive organs: Female- spermathecal capsule bifid; spermathecal gland attached to base of capsule; bursa copulatrix not saccular; bursal plate as in Fig. 159 View Figures 147–162 ( Fig. 160 View Figures 147–162 ).
Cymatodera dietrichi Barr. Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed and very long; testis comprising 1 long follicle (Fig: 155). Female- Spermathecal capsule feeble incised, poorly sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix not saccular; bursal plate as in Fig, 156 ( Fig. 157 View Figures 147–162 ).
Cymatodera fuchsii Schaeffer. Reproductive organs: Female- Spermathecal capsule feebly incised, poorly sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix not saccular; bursal plate as in Fig, 161 ( Fig. 162 View Figures 147–162 ).
Cymatodera horni Wolcott. Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed and very long; testis comprising 1 follicle (Fig: 163). Female- Spermathecal capsule feebly incised, poorly sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix not saccular; bursal plate present ( Fig. 164 View Figures 163–170 ).
Cymatodera inornata (Say) . Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed and very long; testis comprising1 follicle ( Fig. 165 View Figures 163–170 ).
Cymatodera latefascia Schaeffer. Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed and very long; testis comprising 1 follicle ( Fig. 166 View Figures 163–170 ). Female- Spermathecal capsule bifid, poorly sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix not saccular; bursal plate as in Fig. 168 View Figures 163–170 ( Fig. 167 View Figures 163–170 ).
Cymatodera oblita Horn. Reproductive organs: Female- Spermathecal capsule feebly incised, poorly sclerotized; spermathecal gland attached to middle of capsule; bursa copulatrix not saccular; bursal plate present ( Fig. 169 View Figures 163–170 ).
Cymatodera tricolor Skinner. Reproductive organs: Female- Spermathecal capsule bifid, poorly sclerotized; spermathecal gland attached to middle of capsule; bursa copulatrix not saccular; bursal plate present ( Fig. 170 View Figures 163–170 ).
Cymatodera serena Barr. Reproductive organs: Female- Spermathecal capsule feebly incised, poorly sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix not saccular; bursal plate as in Fig. 172 View Figures 171–184 ( Fig. 171 View Figures 171–184 ).
Cymatodera tutoides Barr. Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed and very long; testis comprising 1 follicle ( Fig. 173 View Figures 171–184 ).
Cymatodera undulata (Say) . Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed and very long; testis comprising 1 follicle ( Fig. 175 View Figures 171–184 ). Female- Spermathecal capsule slightly bifid, poorly sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix not saccular; bursal plate as in Fig. 176 View Figures 171–184 ( Fig. 174 View Figures 171–184 ).
Lecontella cancellata (Leconte) . Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed; testis comprising 1 follicle ( Fig. 178 View Figures 171–184 ). Female- Spermathecal capsule very narrow and elongated, poorly sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular; bursal plate as in Fig. 181 View Figures 171–184 ( Fig. 184 View Figures 171–184 ).
Lecontella gnara Wolcott. Reproductive organs: Female- Spermathecal capsule very narrow and elongated, poorly sclerotized; spermathecal gland particularly large, attached to base of capsule; bursa copulatrix saccular; bursal plate present ( Fig. 179 View Figures 171–184 ).
Monophylla terminata (Say) . Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed; testis comprising 1 follicle ( Fig. 183 View Figures 171–184 ). Female- Spermathecal capsule not distinguishable; spermathecal gland well defined; bursal plate as in Fig. 180 View Figures 171–184 ( Fig. 177 View Figures 171–184 ).
Monophylla californica (Fall) . Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed; testis comprising 1 follicle (Fig: 182). Female- Spermathecal capsule not distinguishable; spermathecal gland well defined; bursal plate present ( Fig. 185 View Figures 185–194 ).
Onychotillus vittatus Chapin. Alimentary canal: Stomodaeum long, with proventriculus not well defined externally; ventricular papillae well defined; 4 cryptonephridial malpighian tubules; proctodaeum well developed ( Fig. 186 View Figures 185–194 ). Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed; testis comprising 1 follicle ( Fig. 187 View Figures 185–194 ). Female- Spermathecal capsule bifid; spermathecal gland attached to base of capsule; bursal plate as in Fig. 189 View Figures 185–194 ( Fig. 188 View Figures 185–194 ).
Orthocladiscus dispar (Gorham) . Reproductive organs: Male- Two pairs of accessory glands, medial pair branched, with central longitudinal dark streak, lateral gland not branched; testis comprising 6 follicles ( Fig. 190 View Figures 185–194 ).
Pallenis Castelnau ( Madagascar) . Species. Reproductive organs: Female- Spermathecal capsule poorly sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular; bursal plate present ( Fig. 191 View Figures 185–194 ).
Tillus elongatus (Linnaeus) . Alimentary canal: Stomodaeum long, with proventriculus not well defined externally; ventricular papillae well defined; 6 cryptonephridial malpighian tubules; proctodaeum well developed ( Fig. 194 View Figures 185–194 ). Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed; testis comprising 1 follicle ( Fig. 192 View Figures 185–194 ). Female- Spermathecal capsule long and narrow; spermathecal gland attached to base of capsule; bursal plate present ( Fig. 193 View Figures 185–194 ).
Tillus notatus Klug. Reproductive organs: Female- Spermathecal capsule long and narrow; spermathecal gland attached to base of capsule; bursal plate present ( Fig. 195 View Figures 195–200 ).
Tillus Olivier (Africa) . Species a. Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed; testis comprising 1 follicle ( Fig. 196 View Figures 195–200 ).
Tillus Olivier (Africa) . Species b. Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed; testis comprising one follicle ( Fig. 197 View Figures 195–200 ).
Tilloidea unifasciata (Fabricius) . Reproductive organs: Male- Two pairs of accessory glands, medial pair not branched, with central longitudinal dark streak, lateral gland bilobed; testis comprising 1 follicle ( Fig. 198 View Figures 195–200 ). Female- Spermathecal capsule long, narrow and bifid; spermathecal gland attached to base of capsule; bursa copulatrix not saccular, bursal plate as in Fig. 200 View Figures 195–200 ( Fig. 199 View Figures 195–200 ).
Epiphloeinae Kuwert : Iontoclerus sericeus (Klug) . Reproductive organs: Male- Two pairs of accessory glands, neither gland branched; testis multifollicular ( Fig. 201 View Figures 201–207 ). Female- Spermathecal capsule poorly sclerotized; spermathecal gland attached to distal third of capsule; bursa copulatrix saccular ( Fig. 202 View Figures 201–207 ).
Peloniinae Opitz : Chariessa dichroa (Leconte) . Reproductive organs: Male- Two pairs of accessory glands, neither gland branched; seminal vesicle bulbous; testis multifollicular ( Fig. 204 View Figures 201–207 ). Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 203 View Figures 201–207 ).
Chariessa pilosa (Forster) . Reproductive organs: Male- Two pairs of accessory glands, neither gland branched; testis multifollicular ( Fig. 205 View Figures 201–207 ). Female- Spermathecal capsule well sclerotized, wrinkled; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 206 View Figures 201–207 ).
Chariessa vestita (Chevrolat) . Reproductive organs: Male- Two pairs of accessory glands, glands particularly long and unbranched; testis multifollicular ( Fig. 207 View Figures 201–207 ).
Pelonium alcicorne Klug. Reproductive organs: Female- Spermathecal capsule well sclerotized, wrinkled; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 208 View Figures 208–215 ).
Pelonium amabile Spinola. Reproductive organs: Male- Two pairs of accessory glands, glands unbranched; testis multifollicular ( Fig. 209 View Figures 208–215 ).
Pelonium auripenne (Hope) . Reproductive organs: Male- Two pairs of accessory glands, glands unbranched; testis multifollicular ( Fig. 210 View Figures 208–215 ). Female- Spermathecal capsule well sclerotized, wrinkled; spermathecal gland attached to base of capsule; bursa copulatrix saccular, bursal plate present ( Fig. 211 View Figures 208–215 ).
Pelonium badeni Gorham. Reproductive organs: Male- Two pairs of accessory glands, glands unbranched; testis multifollicular ( Fig. 212 View Figures 208–215 ).
Pelonium fasciculatum Klug. Reproductive organs: Male- Two pairs of accessory glands, glands unbranched; testis multifollicular ( Fig. 213 View Figures 208–215 ). Female- Spermathecal capsule well sclerotized, wrinkled; spermathecal gland attached to base of capsule; bursa copulatrix saccular, bursal plate present ( Fig. 214 View Figures 208–215 ).
Pelonium disconotatum Pic. Reproductive organs: Male- Two pairs of accessory glands, medial pair particularly long, glands unbranched; testis multifollicular ( Fig. 215 View Figures 208–215 ).
Pelonium leucophaeum Klug. Reproductive organs: Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 216 View Figures 216–222 ).
Pelonium lituratum Kirby. Reproductive organs: Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 217 View Figures 216–222 ). Nervous organs: Ventral nerve chord and brain as in Fig. 219 View Figures 216–222 .
Pelonium nigroclavatum Chevrolat. Reproductive organs: Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 218 View Figures 216–222 ).
Pelonium peninsulare Schaeffer. Reproductive organs: Male- Two pairs of accessory glands, glands unbranched; testis multifollicular ( Fig. 220 View Figures 216–222 ). Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 221 View Figures 216–222 ).
Pelonium placidum Schenkling. Reproductive organs: Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 222 View Figures 216–222 ).
Pelonium posticum Klug. Reproductive organs: Male- Two pairs of accessory glands, glands unbranched; testis multifollicular ( Fig. 223 View Figures 223–230 ).
Pelonium quadriplagiatum Kuwert. Reproductive organs: Male- Two pairs of accessory glands, glands unbranched; testis multifollicular ( Fig. 224 View Figures 223–230 ). Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 225 View Figures 223–230 ).
Pelonium semirufum Gorham. Reproductive organs: Male- Two pairs of accessory glands, glands unbranched; testis multifollicular ( Fig. 226 View Figures 223–230 ). Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 227 View Figures 223–230 ).
Pelonium viridipenne Kirby. Reproductive organs: Male- Two pairs of accessory glands, glands unbranched; seminal vesicle saccular; testis multifollicular ( Figs. 229, 230 View Figures 223–230 ). Female- Spermathecal capsule well sclerotized, wrinkled; spermathecal gland attached to middle of capsule; bursa copulatrix saccular, but very small ( Fig. 228 View Figures 223–230 ).
Pelonium Spinola , species a-c ( Brazil). Reproductive organs: Male- Two pairs of accessory glands, glands unbranched; testis multifollicular ( Fig. 231 View Figures 231–241 ). Female- Spermathecal capsule well sclerotized; spermathecal gland attached to base of capsule; bursa copulatrix saccular ( Fig. 232 View Figures 231–241 ). Species b. Reproductive organs: Male- Two pairs of accessory glands, glands unbranched; seminal vesicle saccular; testis multifollicular ( Fig. 233 View Figures 231–241 ). Species c. Reproductive organs: Male- Two pairs of accessory glands, glands unbranched; seminal vesicle saccular; testis multifollicular ( Fig. 234 View Figures 231–241 ).
Tarsosteninae Jacquelin du Val : Tarsostenodes leucogramma Elston. Reproductive organs: Male- Two pairs of accessory glands, glands unbranched; testis comprising 6 follicles ( Fig. 235 View Figures 231–241 ).
Blackburniella hilaris (Westwood) . Reproductive organs: Female- Spermathecal capsule well sclerotized; spermathecal gland attached to apex of capsule; bursa copulatrix saccular ( Fig. 236 View Figures 231–241 ).
Parapylus sedlaceki (Kolibáč) . Reproductive organs: Female- Spermathecal capsule well sclerotized; spermathecal gland attached near base of capsule; bursa copulatrix saccular, bursal plate present ( Fig. 237 View Figures 231–241 ).
Tarsostennus carus (Newman) . Reproductive organs: Male- One pair of accessory glands, gland unbranched; seminal vesicle bulbous; testis comprising 6 follicles ( Fig. 239 View Figures 231–241 ). Female- Spermathecal capsule well sclerotized; spermathecal gland attached near middle of capsule; bursa copulatrix saccular ( Fig. 238 View Figures 231–241 ).
Tarsostenodes guttulus (White) . Reproductive organs: Female- Spermathecal capsule well sclerotized; spermathecal gland attached near base of capsule; bursa copulatrix saccular ( Fig. 240 View Figures 231–241 ).
Korynetinae Laporte : Dolichopsis haplocnemodes Gorham. Reproductive organs: Female- Spermathecal capsule well sclerotized; spermathecal gland attached to apex of capsule; bursa copulatrix saccular ( Fig. 241 View Figures 231–241 ).
Comparative morphology and taxonomic significance of variations
This discussion concerns general descriptions of the alimentary canal and mesodermal reproductive organs. It is not possible to make any comments about the central nervous system and brain, which has not been extensively studied in the Cleridae . Opitz illustrated the ventral nerve cord and brain of Aphelocerus coactus Opitz ( Opitz, 2005: 22) and Necrobia violacea Linnaeus ( Opitz, 2011b: 51) . This work also includes opinions about the taxonomic significance of some of the differences noted in the alimentary canal and mesodermal reproductive organs. The comparative and taxonomic discussion includes information gleaned from previously published works as noted in Opitz (2010a) .
General morphology- Alimentary canal. The gross morphology of the alimentary canal consists of a stomodaeum, ventriculus, cryptonephridial malpighian tubules, and proctodaeum ( Fig. 242 View Figures 242–246 ). The stomodaeum includes a distinct pharynx ( Fig. 243 View Figures 242–246 ), a well-developed esophagus ( Fig. 244 View Figures 242–246 ), and a usually muscular proventriculus ( Fig. 245 View Figures 242–246 ). The pharyngeal intima exhibits several folds that eventually transform into a distinct number of primary and secondary folds in the proventriculus. Intimal spines are numerous in the pharynx, scarce in the anterior region of the esophagus, long and stout and posterior region of the esophagus and anterior region of the stomodaeum. The stomodaeal valve ( Fig. 246 View Figures 242–246 ) consists of lobes that vary in size, shape, and number, but their position in the valve is constant among the taxa examined.
The ventriculus ( Fig. 247 View Figures 247–252 ) is well developed and may be smooth, papillose, or with small transverse wrinkles. Between the ventriculus and the proctodaeum there are the malpighian tubules which attach to the posterior region of the ileum. In the proctodaeum, the pylorus is short and muscular and is marked anteriorly by a 6-lobed pyloric valve ( Fig. 248 View Figures 247–252 ). The anterior intestine is divided into a well-defined ileum and a distally broadened colon. The posterior intestine is composed of a well-developed rectal sac and an elongate rectum proper, which is twice as long in the female as in the male (a morphological development to enable the female to extend her lengthy ovipositor out of the abdomen). The proctodaeal intima shows numerous small folds in the pylorus and 6 larger folds in the colon ( Fig. 249 View Figures 247–252 ). The folds terminate in the region of the rectal valve. The rectal intima exhibits small folds anteriorly and long slender folds posteriorly ( Fig. 250 View Figures 247–252 ). Information about the alimentary canal of the Cleridae , as part of taxonomic revisions, was published recently by Ekis (1977b) and Opitz (2003, 2004, 2005, 2006, 2007, 2008, 2009a, 2009b, 2010b, 2011a, 2011b, 2012)
General morphology-Reproductive organs. When fully developed, the male reproductive organs consist of a pair of testes, vas deferens, seminal vesicle, usually two pairs of accessory glands, and the ejaculatory duct ( Fig. 252 View Figures 247–252 ). The number of testicular follicles per testis varies as does the number of accessory glands and shape of seminal vesicle. The term multifollicular is used to indicate that the testis has more than one follicle; this to distinguish from the singular follicle testis as found in most Tillinae . The female organs ( Fig. 251 View Figures 247–252 ) consist of a pair of ovarioles, lateral oviduct, medial oviduct, bursa copulatrix, spermathecal capsule, spermathecal gland, and the vagina. In some species the base of the bursa copulatrix shows a bursal plate ( Fig. 148 View Figures 147–162 ). There is substantial variation in the number of follicles that constitute the ovary, shape and composition of the spermathecal capsule, attachment site of the spermathecal gland to the capsule, length of the spermathecal gland, and in the presence or absence of a saccular bursa copulatrix.
Taxonomic implications-Alimentary canal. A very long stomodaeum occurs in Somatochaetus Menier and Ekis , Metaxina Broun , and Katachaetosoma Opitz , of Chaetosomatidae , and Thaneroclerus of Thaneroclerinae , and Ababa Casey of Isoclerinae, of the Cleridae . The elongated condition of the stomodaeum was considered a primitive characteristic by Crowson (1972: 340) and Opitz (2010a: 116) . All other Cleridae studied to date show a reduced foregut, presumably a manifestation of the loss of the crop ( Crowson, 1972: 340). The number of stomodaeal lobes that comprises the stomodaeal valve varies at subfamily level. Primitively there are 8 lobes in Thaneroclerinae and Tillinae whereas the derived condition, involving less than 8 folds, is found in the other subfamilies ( Opitz, 2010a: 98 ). The development of the ventricular papillae varies at subfamily level. They are primitively poorly developed in the Thaneroclerinae , Isoclerinae, Epiphloeinae , Enopliinae , Peloniinae, Tarsosteninae , and Neorthopleurinae. Their well-developed, derived, state is found in the Tillinae , Clerinae , Hydnocerinae , and Korynetinae . Presence of 6 malpighian tubules is the primitive condition found in most of Tillinae , Clerinae , Hydnocerinae , and Korynetinae , whereas in the Thaneroclerinae , Isoclerinae, Neorthopleurinae, Epiphloeinae , Peloniinae, Enopliinae , Tarsosteninae , there are 4 tubules, which reflects the derived condition for this character.
Taxonomic implications-Reproductive organs. Some characteristics of the male and female mesodermal reproductive organs are particularly taxonomically useful at the generic and species- group levels ( Opitz 2010a: 51 ). In nearly every species of Tillinae examined, the testis is comprising one follicle and the median accessory gland exhibits a central longitudinal dark streak ( Fig. 154 View Figures 147–162 ). A similar dark streak is found in the spermatophoral accessory glands of antophilic species of Clerinae ( Opitz, 2003) . The spermathecal capsule is bifid and the bursa copulatrix is not saccular in tilline females ( Fig. 158 View Figures 147–162 ). Also, at the base of the bursa there is a bursal plate whose configuration is significant at the species level ( Fig. 159 View Figures 147–162 ). In the Hydnocerinae , the males of Isohydnocera Chapin and Phyllobaenus Dejean are characterized by having only one pair of accessory glands ( Fig. 12 View Figures 1–12 ) that are extraordinarily long ( Fig. 12 View Figures 1–12 ), and in most species of Phyllobaenus Dejean the spermathecal capsule is round and heavily sclerotized ( Fig. 29 View Figures 28–35 ). A saccular bursa copulatrix in not present in all hydnocerine examined. In the Clerinae genus Enoclerus Gahan the mode of bifurcation of the medial accessory glands are taxonomically significant at species-group levels. In various species of subfamily Peloniinae the spermathecal capsule is transversally wrinkled ( Fig. 206 View Figures 201–207 ). A bursal plate is found in the Peloniinae genus Pelonium Spinola. The spermathecal gland is extraordinarily long and the bursal plate is present in the Korynetinae genus Parapylus Blackburn. Information about the mesodermal reproductive organs of the Cleridae , as part of taxonomic revisions, was published recently by Ekis (1977a, 1977b), Ekis (1978), Opitz (1998, 2003, 2004, 2005, 2006, 2007, 2008, 2009a, 2009b, 2009c, 2010a, 2010b, 2010c, 2011a, 2011b, and 2012), and Yang, et. al. (2011).
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