hind wing, mediella strongly arched, areolet small and .... tially, trochanters, hind femora above except for 0.2 at the base ... and hind edge of gastral terga, red.
(XU-(QWRPRO ± ,661
'HVFULSWLRQRIWZRQHZVSHFLHVRI(QFOLVLV +\PHQRSWHUD ,FKQHXPRQLGDH DQGVXSSRUWIRUWKHVHFUHWRU\UROHRIW\ORLGVLQLFKQHXPRQLGPDOHV 6$17,$*2%25'(5$DQG(67()$1Ë$+(51È1'(=52'5Ë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¶V DQWHQQDH GXULQJ FRXUWVKLS WKHUHE\ HQKDQFLQJ IHPDOH UHFHSWLYLW\ 7KLV K\SRWKHVLV ZDV FRQILUPHG E\ %LQ HW DO
VWXG\LQJ WKH LFKQHXPRQLG 3LPSOD WXULRQHOODH / ,FKQHXPRQLGDH 3LPSOLQDH EXW QR RWKHU VXFK VWXGLHVDUHUHSRUWHG ,Q WKLV SDSHU WKH SUHVHQFH RI SRUHV LQ WKH PDOH W\ORLGV RI (QFOLVLV VSHFLHV DUH GHPRQVWUDWHG 7KHVH SRUHV DUH DVVRFLDWHG ZLWK LQWHUQDO FXWLFXODU GXFWV ZKLFK LQGLFDWH D VHFUHWRU\ UDWKHU WKDQ D VHQVRU\ IXQFWLRQ 7KLV GLVFRYHU\ LV IXUWKHU VXSSRUW IRU WKH K\SRWKHVLV RI ,VLGRUR HW DO FRQFHUQLQJWKHELRORJLFDOUROHRIW\ORLGVLQPDOHV RI,FKQHXPRQLGDH 0$7(5,$/$1'0(7+2'6 6SHFLPHQVZHUHFROOHFWHGLQ E\PHDQVRI0DODLVHWUDSV SODFHG DW WZR ORFDOLWLHV QHDU WKH 0HGLWHUUDQHDQ FRDVW RI 6SDLQ 7KH ORFDOLWLHV ZHUH )RQW 5RMD $OFRL $OLFDQWH ¶¶¶1 ¶¶¶: DQG %DUUDQFR GH 6DUDJXWLOOR 6HUUD 9DOHQFLD ¶¶¶1 ¶¶¶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p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
Fig 1. Seasonal activity of adults of E. dichroma sp. n. blade (only the antennomeres with tyloids to facilitate manipulation), treated again with warm potassium hydroxide for 10 minutes and then washed with distilled water. After dehydration in a graded ethanol series (40, 70, 90 and 100%), the antennae were critical-point-dried in a Electron Microscopy Sciences EMS 850 apparatus before gold coated for scanning. Terminology. Terms used for surface sculptures on sclerites were adapted from Eady (1968). RESULTS
Enclisis dichroma sp. n. (Figs 1, 3–6, 14, 18, 22–26) Female. Body length (without ovipositor) 4.7–8.6 mm. Head 0.6–0.9 mm long and 1.0–1.6 mm wide. Thorax 1.6–2.9 mm long, 0.9–1.5 mm wide (mesoscutum). Front wings 3.8–6.0 mm long. Petiole 0.5–0.9 mm long. Postpetiole 0.3–0.6 mm long and 0.4–0.6 mm wide. Ovipositor sheaths 2.0–3.5 mm long. Coloration. Reddish. Spots on the cheeks close to base of mandibles, mandibular teeth, vertex, occiput and postocciput partially or completely, upper half of temples, oral and occipital carina, basal flagellar segments above, hind edge of propleurum, sutures of meso and metanotum, tegula, sternum between coxae, all coxae partially, trochanters, hind femora above except for 0.2 at the base, and base of petiole black. Gastral segments, except for the central basal area of the second one, fore and middle femora above, tibiae partially and tarsi brown to dark brown. Hind margin of terga 2–3 and palpi usually light brown. Antennae with white marks on the 6–9 flagellar segments. Seventh gastral segment with a white membrane. Wings with transverse dark band. Description. Head (Fig. 14). Transverse, 1.7–1.8 times as wide as long, strongly constricted behind the eyes. Temple 0.3 times as long as eye (viewed from above). Vertex, frons and face with dense punctures on granulate background. Temples with small and very sparse setiferous punctures on a smooth and polished background. Malar space strongly granulate, about 1.1–1.2 times as wide as basal width of mandible. Clypeus moderately 402
Fig 2. Seasonal activity of adults of E. schwarzi sp. n.
convex and polished, with sparse weak punctures, about 2.2–2.3 times as wide as long, its apical margin straight, with sharp lateral edges and a blunt angulation on midline. Lower mandibular tooth clearly longer than upper tooth. Antennae with 23–25 segments; first flagellar segment 3.6–3.7 times as long as wide (Fig. 18). Genal carina joining oral carina behind base of mandible at a distance of 0.6–0.8 times the basal width of the mandible. Hind ocelli separated from eye about 1.5–1.8 times their diameter. Space between hind ocelli 1.5–1.8 their diameter. Thorax (Fig. 3). Pronotum weakly strigose on the hind edge. Epomia present. Mesoscutum polished, weakly coriaceous in the centre; weak and very sparse punctures on the lateral lobes, more deep and dense in the centre but always smaller than their interspaces. Notauli deep, reaching about 0.5–0.6 the distance to scutellum. Scutellar groove with longitudinal carinae. Scutellum smooth and polished, finely punctate and without lateral carinae. Mesopleurum finely rugose on the upper side of sternaulus, except on and in front of the speculum; the background polished with very sparse setiferous punctation, sometimes weakly granulate in the rugose area. Speculum smooth, polished and completely hairless. Sternaulus sinuous, its hind end, weakly downward toward base of middle coxa, only strongly distinct on 2/3 the length of mesopleurum. Metapleurum conspicuously rugose; rest of propodeum weakly rugose or smooth, on a polished or finely granulate background. Both of its transverse carinae strong and complete but the apical transverse carina not forming a sublateral crest. Propodeal spiracle circular and joining pleural carina. Juxtacoxal carina strongly present at least at the base. Ramulus of fore wing absent. Radial cell, 2.8–3.3 times as long as its maximum width. Second recurrent vein weakly inclivous or almost vertical. Areolet small and pentagonal. Nervulus opposite basal vein. Nervelus vertical, strongly angular and intercepted a little below the middle. Hind femora 4.6–4.8 times as long as high.
)LJV± 0RUSKRORJ\RIWKHWKRUD[DQGJDVWHURI(GLFKURPD±WKRUD[ODWHUDOYLHZ ±ILUVWWRWKLUGJDVWUDOWHUJD ±RYL SRVLWRUWLS ±RYLSRVLWRUVHQVRU\UHFHSWRUV V
*DVWHU )LJV ± )LUVW JDVWUDO WHUJXP ZHDNO\ DQG ILQHO\ JUDQXODWH WKH KLQG VXEODWHUDO HGJH WHQGLQJ WR EH VPRRWK DQG SROLVKHG 0HGLDQ GRUVDO FDULQDH DEVHQW RU ZHDNO\ LQVLQXDWHG RQ SHWLROH EHFRPLQJ REVROHVFHQW RQ WKH KLQG KDOI RI SRVWSHWLROH 3RVWSHWLROH ± WLPHV DV ORQJ DV LWV PD[LPXP ZLGWK GRUVDOO\ PHDVXUHG WKH VSLUDFOHRQO\VOLJKWO\EHKLQGWKHPLGGOH6HFRQGDQGUHVW RI WHUJD FRULDFHRXV ZLWK PRGHUDWHO\ GHQVH VHWLIHURXV SXQFWDWLRQ KLQGFHQWUDOSDUWRIWKHIRUPHUZLWKWKLVSXQF WDWLRQ PXFK PRUH VSDUVH 6HFRQG JDVWUDO WHUJXP ± WLPHVDVORQJDVLWVPD[LPXPZLGWK GRUVDOO\PHDVXUHG 7K\ULGLDVPDOODQGURXQGHGZHDNO\FRQFDYH2YLSRVLWRU VOLJKWO\ XSFXUYHG ZLWK QRGXV DQG VWURQJ WHHWK RQ WKH DSH[ RI ORZHU YDOYH LWV WLS ± WLPHV WKH PD[LPXP KHLJKWDWQRGXV )LJ 8SSHUDQGORZHUYDOYHVZLWKWZR OLQHV RI FRQVSLFXRXV VHQVRU\ UHFHSWRUV )LJ V 2YL SRVLWRUVKHDWKV±WLPHVDVORQJDVWKHKLQGWLELDH 0DOH%RG\OHQJWK±PP+HDG±PPORQJ DQG±PPZLGH7KRUD[±PPORQJ± PP ZLGH PHVRVFXWXP )URQW ZLQJ ± PP ORQJ 3HWLROH ± PP ORQJ 3RVWSHWLROH ± PP ORQJ DQG±PPZLGH6HFRQGWHUJXP±PPORQJ &RORUDWLRQ %ODFN 7KRUD[ PRVWO\ H[FHSW VXWXUHV DQG D[LOODU\WURXJKRIPHVRQRWXPDQGPHWDQRWXP IURQWDQG
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± WLPHV DV ZLGH DV ORQJ 7HPSOH ± WLPHV DV ORQJ DV H\H 0DODUVSDFHDERXW ±WLPHVDVZLGHDVEDVDOZLGWKRI PDQGLEOH &O\SHXV ± WLPHV DV ZLGH DV ORQJ DSLFDO PDUJLQDVLQIHPDOH$QWHQQDHZLWK±VHJPHQWV ILUVW IODJHOODU VHJPHQW ± WLPHV DV ORQJ DV LWV PD[LPXP ZLGWK W\ORLGV RQ WKH WKWK IODJHOODU VHJPHQWV )LJ WKH\ KDYH VPDOO SRUHV RSHQLQJ DPRQJ MX[WDSRVHG GRUVDOFXWLFXODUSODWHV )LJV ZKLFKDUHDVVRFLDWHG LQWHUQDOO\ ZLWK FXWLFXODU GXFWV )LJ )ODJHOOXP ZLWK VWURQJO\ SURWUXGLQJ VHWDH DPRQJ VKRUW SXEHVFHQFH )LJ *HQDOFDULQDMRLQLQJRUDOFDULQDEHKLQGEDVHRIPDQ GLEOHDWDGLVWDQFHRI±WLPHVWKHEDVDOZLGWKRIWKH PDQGLEOH +LQG RFHOOL VHSDUDWHG IURP H\H E\ DERXW
)LJV± 0RUSKRORJ\RIWKHWKRUD[DQGJDVWHURI(VFKZDU]L±WKRUD[ODWHUDOYLHZ ±ILUVWWRWKLUGJDVWUDOWHUJD ±RYL SRVLWRUWLS ±RYLSRVLWRUVHQVRU\UHFHSWRUV V
±WKHLUGLDPHWHU6SDFHEHWZHHQKLQGRFHOOL± WLPHVWKHLUGLDPHWHU2WKHUIHDWXUHVDVLQIHPDOH 7KRUD[5DPXOXVDEVHQWLQDIHZFDVHVVOLJKWO\SUHVHQW DWEDVH5DGLDOFHOO ±WLPHVDVORQJDVLWVPD[LPXP ZLGWK+LQGIHPRUD±WLPHVDVORQJDVKLJK2WKHU FKDUDFWHUVDVLQIHPDOH *DVWHU )LUVW JDVWUDO WHUJXP FRULDFHRXV WKH KLQG HGJH WHQGLQJWREHVPRRWKDQGSROLVKHG0HGLDQGRUVDOFDULQDH DEVHQW 5HVW RI JDVWUDO WHUJD VWURQJO\ FRULDFHRXV ZLWK GHQVH VHWLIHURXV SXQFWDWLRQ 6HFRQG JDVWUDO WHUJXP ± WLPHV DV ORQJ DV LWV PD[LPXP ZLGWK VHHQ IURP DERYH 7K\ULGLDHORQJDWHG 7\SH PDWHULDO +RORW\SH IHPDOH %DUUDQFR 6DUDJXWLOOR 6HUUD 9DOHQFLD 6SDLQ ,9 OHJ ) /XQD &(8$ 3DUDW\SHV VDPH ORFDOLW\ ,,, PDOH ,,, IHPDOH PDOHV ,,,,9 IHPDOH PDOHV ,9 IHPDOHVPDOHV IHPDOHPDOH01&1 IHPDOH PDOH 6& ,99 PDOH 9 PDOHDOOWKHPOHJ)/XQD (W\PRORJ\GL *UHHN WZR FKURPXV *UHHN FRORXUHG7KH VSHFLHVQDPHUHIHUVWRWKHEODFNJDVWHUDQGUHGKHDGDQGWKRUD[
+DELWDW%DUUDQFRGHO6DUDJXWLOOR 6HUUD9DOHQFLD LVD YDOOH\ LQ WKH 3RUWD&RHOL PRXQWDLQV QHDU WKH HDVWHUQ 6SDQLVK FRDVW 7KH PLQLPXP KHLJKW RI WKLV YDOOH\ DERYH VHDOHYHOLVPDQGLWVPD[LPXPP7KH0DODLVH
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±PP SHU \HDU ZLWK PRVW IDOOLQJ LQ VSULQJ DQG DXWXPQ $QQXDO PHDQ WHPSHUDWXUHV DUH XVXDOO\ DERYH &0LQLPXPWHPSHUDWXUHVXQGHU&DUHRFFDVLRQDOO\ UHFRUGHG IURP 'HFHPEHU WR )HEUXDU\ 'RFDYR HW DO %LRORJ\ 7KLV VSHFLHV KDV D FOHDU SHDN RI DFWLYLW\ LQ VSULQJ )LJ EXWLWLVXQNQRZQZKHWKHULWKDVDQRWKHU SHDN LQ DXWXPQ OLNH RWKHU UHODWHG VSHFLHV +RVWV DUH XQNQRZQ (QFOLVLVVFKZDU]LVSQ )LJV±±
Figs 19–22: Morphology of males of Enclisis species, heads in dorsal view. 19 – E. infernator; 20 – E. schwarzi; 21 – E. ornaticeps; 22 – E. dichroma.
Figs 11–18: Morphology of females of Enclisis species. 11–14 – heads, dorsal view. 11 – E. infernator; 12 – E. schwarzi; 13 – E. ornaticeps; 14 – E. dichroma. 15–18 – basal antennomeres. 15 – E. infernator; 16 – E. schwarzi; 17 – E. ornaticeps; 18 – E. dichroma.
Female. Body length (without ovipositor) 5.1–5.9 mm. Head 0.6–0.7 mm long and 1.1–1.3 mm wide. Thorax 1.8–2.1 mm long, 0.8–1.0 mm wide (mesoscutum). Front wings 4.0–4.5 mm long. Petiole 0.4–0.6 mm long. Postpetiole 0.4–0.6 mm long and wide. Ovipositor sheaths 1.8–2.0 mm long. Coloration. Black. Clypeus, mandibles, basal antennal segments ventrally, hind half of postpetiole, second and
Figs 23–26: Antennal morphology of males of E. dichroma. 23 – 11th-14th flagellar segments showing tyloids (ty); 24 – tyloid of 12th flagellomere showing cuticular pores (p); 25 – pores of tyloid (p); 26 –internal ultrastucture of tyloid showing numerous secretory cuticular ducts (cd).
405
)LJV± $QWHQQDOPRUSKRORJ\RIPDOHVRI(VFKZDU]L ±WKWK IODJHOODUVHJPHQWVVKRZLQJW\ORLGV W\ ±W\ORLGRI WKIODJHOORPHUHVKRZLQJFXWLFXODUSRUHV S ±SRUHVRIW\ORLG S ±LQWHUQDOXOWUDVWXFWXUHRIW\ORLGVKRZLQJQXPHURXVVHFUH WRU\FXWLFXODUGXFWV FG
WKLUG JDVWUDO VHJPHQWV UHGGLVK 3DOSL GLVWDO KDOI RI IURQW DQG PLGGOH IHPRUD WLELDH DQG WDUVL OLJKW EURZQ )ODJHO OXP WHJXOD KLQG WLELD DQG WDUVL EURZQ $QWHQQDH ZLWK D ZKLWH PDUN GRUVDOO\ RQ WKH WKWK IODJHOODU VHJPHQWV 6HYHQWK JDVWUDO VHJPHQW ZLWK D ZKLWH PHPEUDQH :LQJV ZLWKDWUDQVYHUVHGDUNEDQG 'HVFULSWLRQ+HDG )LJ 7UDQVYHUVH± WLPHV DV ZLGH DV ORQJ VWURQJO\ FRQVWULFWHG EHKLQG WKH H\HV 7HPSOH WLPHV DV ORQJ DV H\H YLHZHG IURP DERYH )DFHIURQVDQGYHUWH[ZLWKGHQVHSXQFWXUHVRQJUDQXODWH EDFNJURXQG 7HPSOHV ZLWK VPDOO DQG VSDUVH VHWLIHURXV SXQFWXUHV RQ D JUDQXODWH EDFNJURXQG RQ WKH XSSHU KDOI DQG RQ D VPRRWK DQG SROLVKHG EDFNJURXQG RQ WKH ORZHU KDOI &KHHNV VWURQJO\ JUDQXODWH ZLWK D YHU\ VKDOORZ DQG VSDUVH SXQFWDWLRQ DERXW ± WLPHV DV ZLGH DV EDVDO ZLGWK RIPDQGLEOH &O\SHXVDERXW ±WLPHV DVZLGH DV ORQJ PRGHUDWHO\ FRQYH[ SROLVKHG RU VOLJWKO\ JUDQXODWH ZLWK YHU\ VSDUVH GHHS SXQFWXUHV LWV DSLFDO PDUJLQVWUDLJKWZLWKVKDUSODWHUDOHGJHVDQGDEOXQWDQJX ODWLRQ RQ PLGOLQH /RZHU PDQGLEXODU WRRWK ORQJHU WKDQ XSSHU WRRWK $QWHQQDH ZLWK ± VHJPHQWV ILUVW IODJ HOODU VHJPHQW ± WLPHV DV ORQJ DV ZLGH )LJ *HQDOFDULQDMRLQLQJRUDOFDULQDEHKLQGEDVHRIPDQGLEOH
DWDGLVWDQFH±WLPHVWKHEDVDOZLGWKRIWKHPDQGL EOH +LQG RFHOOL VHSDUDWHG IURP H\H E\ DERXW ± WLPHV WKHLU GLDPHWHU VSDFH EHWZHHQ KLQG RFHOOL ± WKHLUGLDPHWHU 7KRUD[ )LJ 3URQRWXP ZLWK ZHDN ORQJLWXGLQDO ZULQNOHV(SRPLDSUHVHQW0HVRVFXWXPSROLVKHGVOLJWKO\ JUDQXODWH LQ WKH FHQWUH ZLWK ZHDN DQG YHU\ VSDUVH SXQF WXUHVWKHLULQWHUVSDFHVWZLFHWKHLUGLDPHWHU1RWDXOLGHHS DQG UHDFKLQJ DERXW ±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
SOHWHEXWWKHDSLFDOWUDQVYHUVHFDULQDQRWIRUPLQJDVXEODW HUDOFUHVW3URSRGHDOVSLUDFOHFLUFXODUDQGMRLQLQJSOHXUDO FDULQD -X[WDFR[DO FDULQD VWURQJO\ SUHVHQW DW OHDVW DW WKH EDVH 5DPXOXV RI IRUH ZLQJ DEVHQW 5DGLDO FHOO ± WLPHV DV ORQJ DV LWV PD[LPXP ZLGWK 6HFRQG UHFXUUHQW YHLQ YHUWLFDO RU ZHDNO\ UHFOLYRXV $UHROHW SHQWDJRQDO 1HUYXOXVRSSRVLWHEDVDO YHLQ1HUYHOXVYHUWLFDOVWURQJO\ DQJXODU DQG LQWHUFHSWHG D OLWWOH EHORZ WKH PLGGOH +LQG IHPRUD±WLPHVDVORQJDVKLJK *DVWHU )LJV ± )LUVW JDVWUDO WHUJXP JUDQXODWH 0HGLDQGRUVDOFDULQDHGLVWLQFWRQSHWLROHEHFRPLQJREVR OHVFHQWRQWKHKLQGKDOIRISRVWSHWLROH3RVWSHWLROHDVORQJ DV LWV PD[LPXP ZLGWK GRUVDOO\ PHDVXUHG WKH VSLUDFOH RQO\VOLJKWO\EHKLQGWKHPLGGOH6HFRQGDQGUHVWRIWHUJD FRULDFHRXV ZLWK PRGHUDWHO\ GHQVH VHWLIHURXV SXQFWDWLRQ KLQGFHQWUDOSDUWRIWKHIRUPHUZLWKWKLVSXQFWDWLRQPRUH VSDUVH 6HFRQG JDVWUDO WHUJXP WLPHV DV ORQJ DV LWV PD[LPXP ZLGWK GRUVDOO\ PHDVXUHG 7K\ULGLD URXQGHG YHU\ VOLJKWO\ GHYHORSHG 2YLSRVLWRU ZHDNO\ XSFXUYHG ZLWK QRGXV DQG VWURQJ WHHWK RQ WKH DSH[ RI ORZHU YDOYH LWV WLS ± WLPHV WKH PD[LPXP KHLJKW DW QRGXV )LJ 8SSHUDQGORZHUYDOYHVZLWKWZROLQHVRIFRQVSLFXRXV VHQVRU\UHFHSWRUV )LJV 2YLSRVLWRUVKHDWKV± WLPHVDVORQJDVWKHKLQGWLELDH 0DOH%RG\OHQJWK±PP+HDG±PPORQJ DQG ±PPZLGH7KRUD[±PPORQJ± PP ZLGH PHVRVFXWXP )URQW ZLQJ ± PP ORQJ 3HWLROH ± PP ORQJ 3RVWSHWLROH ± PP ORQJ DQG±PPZLGH6HFRQGVHJPHQW±PPORQJ DQG±PPZLGH &RORUDWLRQ %ODFN 0DQGLEOHV FO\SHXV SDUWLDOO\ SUR[LPDOHGJHRIIHPRUDDQGVHFRQGWURFKDQWHUVWK\ULGLD DSH[RIVHFRQGWHUJXPKLQGKDOIRIWKLUGXVXDOO\WKHKLQG KDOI RI IRXUWK DQG KLQG HGJH RI ILIWK UHGGLVK 7HJXOD ZKLWH WR EURZQ 6RPHWLPHV WZR VSRWV RQ WKH IURQWDO RUELWVFROODURISURQRWXPPDQGLEXODUEDVHVYHQWUDOSDUW RIIURQWDQGPLGGOHWURFKDQWHUV\HOORZ3DOSLGLVWDOKDOI RI IURQW DQG PLGGOH IHPRUD WLELDH DQG IURQW WDUVL OLJKW EURZQ $QWHQQDH KLQG WLELDH PLGGOH DQG KLQG WDUVL EURZQ 'HVFULSWLRQ+HDG )LJ 7UDQVYHUVH± WLPHV DVZLGHDVORQJ9HUWH[SROLVKHGZLWKYHU\VOLJKWO\VHWLI HURXV SXQFWXUHV 7HPSOH ± WLPHV DV ORQJ DV H\H SROLVKHG DQG SXQFWXUHG OLNH YHUWH[ 0DODU VSDFH DERXW ± WLPHV DV ZLGH DV EDVDO ZLGWK RI PDQGLEOH &O\SHXV ± WLPHV DV ZLGH DV ORQJ DSLFDO PDUJLQ DV LQ IHPDOH $QWHQQDH ZLWK ± VHJPHQWV ILUVW IODJHOODU VHJPHQW ± WLPHV DV ORQJ DV ZLGH W\ORLGV RQ WKH WKWK IODJHOODU VHJPHQWV )LJ WKH\ KDYH VPDOO SRUHV RSHQLQJ DPRQJ MX[WDSRVHG GRUVDO FXWLFXODU SODWHV )LJV WKHVH SRUHV DUH FRQQHFWHG LQWHUQDOO\ WR FXWLFXODU FDQDOV )LJ )ODJHOOXP ZLWK VWURQJO\ SUR WUXGLQJ VHWDH DPRQJ VKRUW SXEHVFHQFH )LJ *HQDO FDULQD MRLQLQJ RUDO FDULQD EHKLQG EDVH RI PDQGLEOH DW D GLVWDQFH RI WKH EDVDO ZLGWK RI PDQGLEOH +LQG RFHOOL VHSDUDWHG IURP H\H E\ D GLVWDQFH RI DERXW ± WKHLU GLDPHWHU 6SDFH EHWZHHQ KLQG RFHOOL DERXW ± WLPHV WKHLUGLDPHWHU2WKHUFKDUDFWHUVDVLQIHPDOH
7KRUD[ 5DPXOXV DEVHQW 5DGLDO FHOO ± WLPHV DV ORQJDVLWV PD[LPXPZLGWK+LQG IHPRUD±WLPHV DV ORQJDVKLJK2WKHUIHDWXUHVDVLQIHPDOH *DVWHU )LUVW JDVWUDO WHUJXPJUDQXODWH ZLWK GHQVH VHWLI HURXVSXQFWDWLRQWKHKLQGHGJHWHQGLQJWREHVPRRWKDQG SROLVKHG 0HGLDQ GRUVDO FDULQDH DEVHQW 5HVW RI JDVWUDO WHUJD VWURQJO\ FRULDFHRXV ZLWK GHQVH VHWLIHURXV SXQFWD WLRQ 6HFRQG JDVWUDO WHUJXP ± WLPHV DV ORQJ DV LWV PD[LPXPZLGWK VHHQIURPDERYH 7K\ULGLDHORQJDWHG 7\SH PDWHULDO +RORW\SH IHPDOH )RQW 5RMD $OLFDQWH 6SDLQ 9 OHJ)/XQD &(8$ 3DUDW\SHV ,9 9 PDOH 9 IHPDOH PDOHV PDOH 01&1 PDOH 6& 99, PDOH 9,,, ,; PDOHV ,;; IHPDOHPDOH DOOWKHPOHJ )/XQD (W\PRORJ\7KLVVSHFLHVLVGHGLFDWHGWR'U0DUWLQ6FKZDU] /LQ]$XVWULD IRUKLVYDOXDEOHZRUNRQWD[RQRP\RI&U\SWLQDH
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± PP SHU \HDU LQ WKLV DUHD LV LUUHJXODUO\ GLVWULEXWHG WKURXJKRXW WKH \HDU ZLWK PD[LPXPVLQVSULQJDQGDXWXPQ$QQXDOPHDQWHPSHUD WXUHVDUHIURPWR&LQWKHORZODQGVDQGXQGHU& LQWKH KLJKODQGV 0LQLPXP WHPSHUDWXUHV XQGHU & XVX DOO\RFFXUIURP2FWREHUWR0D\ %RURQDWHWDO %LRORJ\7KLVVSHFLHVVHHPVWREHELYROWLQHZLWKSHDNV RIIOLJKWDFWLYLW\LQVSULQJDQGDXWXPQ )LJ +RVWVDUH XQNQRZQ ',6&866,21
(XURSHDQVSHFLHVZHUHUHYLVHGNH\HGDQGLOOXVWUDWHGE\ 6FKZDU] 7KHQHZVSHFLHVDUHFORVHO\UHODWHGWR( RUQDWLFHSV 7KRPVRQ DQG ( LQIHUQDWRU $XEHUW LQKDYLQJDSROLVKHGPHVRVFXWXPZLWKVSDUVHSXQF WXUHV VPDOOHU WKDQ WKH LQWHUVSDFHV SROLVKHG DQG UXJRVH PHVRSOHXUXP ZLWK VSDUVH VHWLIHURXV SXQFWXUHV EODFN WKRUD[ RU LI UHGGLVK WKHQ WKH VFXWHOOXP LV DOVR UHGGLVK KLQGWDUVLZLWKRXWDZKLWHULQJDQGILUVWIODJHOODUVHJPHQW OHVVWKDQILYHWLPHVDVORQJDVZLGH$QWHQQDHRIIHPDOH DQGPDOHZLWK±DQG±VHJPHQWVUHVSHFWLYHO\ 7KHQHZWD[DFDQEHGLVWLQJXLVKHGDVIROORZV
)HPDOHV
7KRUD[ DQG KHDG EODFN $W PRVW D VSRW RQ IURQWDO RUELW FO\SHXV DQG PDQGLEOHV SDUWLDOO\ UHGGLVK 6SDFH EHWZHHQ KLQG RFHOOL ± WKHLU GLDPHWHU +LQG RFHOOL VHSDUDWHG IURPH\HE\DGLVWDQFHRIDERXW±WLPHVWKHLUGLDPHWHU )LJV ± +HDG DQG WKRUD[ PRVWO\ UHGGLVK 6FXWHOOXP DOZD\V UHG 6SDFH EHWZHHQ KLQG RFHOOL ± WKHLU GLDPHWHU +LQG RFHOOL VHSDUDWHG IURP H\H E\ D GLVWDQFH RI DERXW ± WLPHVWKHLUGLDPHWHU )LJV *DVWHU EODFN VRPHWLPHV SRVWSHWLROH UHGGLVK +LQG IHPXU WLPHVDVORQJDVGHHS)LUVWIODJHOODUVHJPHQW WLPHVLWVPD[LPXPZLGWK )LJ (LQIHUQDWRU $XEHUW ± 3RVWSHWLROH VHFRQG DQG WKLUG JDVWUDO WHUJD UHG +LQG IHPXU WLPHVDVORQJDVGHHS)LUVWIODJHOODUVHJPHQW± WLPHVLWVPD[LPXPZLGWK )LJ (VFKZDU]LVSQ *DVWHU UHGGLVK VRPHWLPHV EDVH RI ILUVW WHUJXP DQG IURP ILIWK WR DSH[ IXVFRXV )LUVW IODJHOODU VHJPHQW ± WLPHV LWV PD[LPXP ZLGWK )LJ +LQG IHPXU ± WLPHV DV ORQJDVGHHS (RUQDWLFHSV 7KRPVRQ ± *DVWHU EODFN WR GDUN EURZQ VRPHWLPHV SRVWSHWLROH DQG IURQWDQGRUKLQGHGJHRIVHFRQGWHUJXPUHGGLVK)LUVWIODJ HOODU VHJPHQW ± WLPHV LWV PD[LPXP ZLGWK )LJ +LQGIHPXU±WLPHVDVORQJDVGHHS (GLFKURPDVSQ
0DOHV
+LQG RFHOOL VHSDUDWHG IURP H\H E\ D GLVWDQFH RI DERXW ± WLPHV WKHLU GLDPHWHU 6SDFH EHWZHHQ WKHP ± WKHLU GLDPHWHU )LJV +HDG EODFN DW PRVW FO\SHXV DQG PDQGLEOHV UHGGLVK VRPHWLPHV D \HOORZ VSRW RQ IDFLDO DQGRUIURQWDORUELWV&R[DHDQGWURFKDQWHUVPRVWO\EODFN ± +LQG RFHOOL VHSDUDWHG IURP H\H E\ D GLVWDQFH RI DERXW ± WLPHV WKHLU GLDPHWHU 6SDFH EHWZHHQ WKHP ± WKHLU GLDPHWHU )LJV +HDG ZLWK PDQGLEOHV LQQHU RUELWVIDFHFO\SHXVFKHHNVDQGPRVWRIWKHWHPSOHZKLWH )URQW FR[DH YHQWUDOO\ DQG IURQW DQG PLGGOH WURFKDQWHUV ZKLWH *DVWHU EODFN DW PRVW D ILQH OLQH RQ WKH KLQG HGJH RI WHUJD \HOORZ+LQGIHPXU±WLPHVDVORQJDVGHHS (LQIHUQDWRU $XEHUW ± +LQGKDOIRIWKLUGWHUJXPRIJDVWHUDQGXVXDOO\DZLGHEDQG RQWKHKLQGHGJHRIVHFRQGDQGPRVWRIIRXUWKUHGGLVK+LQG IHPXU±WLPHVDVORQJDVGHHS (VFKZDU]LVSQ 7KRUD[ EODFN 6FXWHOOXP DQG SRVWVFXWHOOXP ZLWK \HOORZ PDUNV
