Äîêóìåíò âçÿò èç êýøà ïîèñêîâîé ìàøèíû. Àäðåñ îðèãèíàëüíîãî äîêóìåíòà : http://zmmu.msu.ru/spec/pdf/mostovsky_mb_mikhailovskaya_mv_2003_153-165.pdf Äàòà èçìåíåíèÿ: Fri Oct 17 16:07:18 2008 Äàòà èíäåêñèðîâàíèÿ: Sun Apr 10 05:39:46 2016 Êîäèðîâêà:

(XU - (QWRPRO ± ,661

$UHYLHZRI3DODHDUFWLF *\PQRSKRUD 0DFTXDUW 'LSWHUD 3KRULGDH ZLWK GHVFULSWLRQ RI QHZ VSHFLHV
0,.+$,/ % 0267296., DQG 0$5,1$ 9 0,.+$,/296.$<$




3DOHRQWRORJLFDO ,QVWLWXWH 3URIVR\X]QD\D 6WUHHW 0RVFRZ 5XVVLD HPDLO SKRULGV#KRWPDLOFRP 0RXQWDLQ7DLJD 6WDWLRQ )DU (DVWHUQ %UDQFK RI 5XVVLDQ $FDGHP\ RI 6FLHQFHV *RUQRWD\R]KQRH 0DULWLPH 7HUULWRU\ 5XVVLD

.H\ ZRUGV 7D[RQRP\ 3KRULGDH *\PQRSKRUD 3DODHDUFWLF QHZ VSHFLHV QHZ V\QRQ\P\ OHFWRW\SHV GLVWULEXWLRQ $EVWUDFW 3DODHDUFWLF VSHFLHV RI WKH JHQXV *\PQRSKRUD DUH UHYLHZHG )RXU QHZ VSHFLHV * DPXUHQVLV VS Q * JRUQRVWDHYL VS Q * SHUSURSLQTXD VS Q DQG * YLFWRULD VS Q DUH GHVFULEHG IURP WKH (XURSHDQ 5XVVLD 0LGGOH $VLD DQG 5XVVLDQ )DU (DVW )HPDOHV RI WZR RWKHU VSHFLHV IURP WKH )DU (DVW DUH GHVFULEHG XQGHU FRGH OHWWHUV * ODFLQLDWD 0LFKDLORYVND\D LV V\QRQ\PLVHG XQGHU * SUHVFKHUZHEHUDH 'LVQH\ /HFWRW\SHV RI * DUFXDWD 0HLJHQ  DQG * YHUUXFDWD 6FKPLW]  DUH GHVLJQDWHG 7KH ODWWHU VSHFLHV LV UHGHVFULEHG 3DODHDUFWLF IHPDOHV RI * QLJULSHQQLV GHPRQVWUDWH ZLGH YDULDWLRQ RI WD[RQRPLFDOO\ LPSRUWDQW IHDWXUHV DQG PD\ LQ IDFW UHSUHVHQW D JURXS RI FORVHO\ UHODWHG VSHFLHV 2QH IHPDOH RI * QLJULSHQQLV ZKLFK KDV EHHQ FDXJKW DOLYH LV UHFRUGHG WR EH LQIHFWHG ZLWK IXQJL ,1752'8&7,21 DQG 6 UHVSHFWLYHO\ ZLWK ILJXUHV LQGLFDWLQJ WKH QXPEHU RI DQ DEGRPLQDO VHJPHQW :LQJ WHUPLQRORJ\ IROORZV 'LVQH\  0DWHULDO VWXGLHG LV NHSW LQ WKH IROORZLQJ FROOHFWLRQV =RR ORJLFDO 0XVHXP &DPEULGJH 8. ± &0= =RRORJLFDO ,QVWLWXWH 6W 3HWHUVEXUJ ± =,1 =RRORJLFDO 0XVHXP RI 0RVFRZ 8QLYHU VLW\ ± =008 ,QVWLWXWH RI %LRORJ\ DQG 3HGRORJ\ 9ODGLYRVWRN ± ,%3 0XVHXP 1DWLRQDO G¶+LVWRLUH 1DWXUHOOH 3DULV ± 01+1 /RV $QJHOHV &RXQW\ 0XVHXP 86$ ± /$&0 =RRORJLVFKHV )RUVFKXQJVLQVWLWXW XQG 0XVHXP $OH[DQGHU .RHQLJ %RQQ ± =)0. =RRORJLFDO 0XVHXP RI 8QLYHUVLW\ RI +HOVLQNL ± =08+ FROOHFWLRQ RI WKH ILUVW DXWKRU ± 0%0 FROOHFWLRQ RI WKH VHFRQG DXWKRU ± 090 7$;2120<

7KH JHQXV *\PQRSKRUD 0DFTXDUW FRPSULVHV PRUH WKDQ VSHFLHV UHFRUGHG IURP WKH +RODUFWLF DQG 2ULHQWDO UHJLRQV (XURSHDQ VSHFLHV DUH FRYHUHG E\ 'LVQH\  DQG $VLDQ VSHFLHV KDYH EHHQ WUHDWHG E\ %URZQ  /LX  DQG 0LFKDLORYVND\D  +RZHYHU WKLV JHQXV LV VWLOO SRRUO\ NQRZQ LQ WKH 3DODHDUFWLF 5HJLRQ ,Q WKLV SDSHU ZH DUH UHYLHZLQJ 3DODHDUFWLF VSHFLHV RI WKLV JHQXV ZLWK VSHFLDO UHIHUHQFH WR WKH WHUULWRU\ RI 5XVVLD DQG DOOLHG FRXQWULHV -DSDQHVH VSH FLHV DUH VDWLVIDFWRU\ NH\HG DQG LOOXVWUDWHG E\ %URZQ  DQG ZH GR QRW LQFOXGH WKHP XQOHVV WKH\ DUH UHFRUGHGIURPWKHPDLQODQG 5HSUHVHQWDWLYHV RI WKH JHQXV *\PQRSKRUD KDYH TXLWH GLVWLQFWLYH JHQHUDO DSSHDUDQFH DQG DUH HDVLO\ UHFRJQLVDEOH XQGHU ORZ PDJQLILFDWLRQ E\ WKH UHGXFHG IURQWDO WKRUDFLF DQG FRVWDO VHWDWLRQ WKH VOHQGHU KLQG OHJV ZLWK WKH WLELDH ODFNLQJ RI D ORQJLWXGLQDO KDLU SDOLVDGH DQG WKH ODUJH ZLQJV H[FHSW LQ WKH EUDFK\SWHURXV * ODSLGLFROD ZLWK D SHFXOLDU YHQDWLRQ 0DOH K\SRS\JLXP LV XVXDOO\ ZLWK GUDZQ LQWR WKH DEGRPHQ DQG FLUFXPYHUWHG FRXQWHU FORFNZLVH %LRORJLFDO GDWD RQ *\PQRSKRUD DUH VFDUFH DQG VXPPDULVHG UHFHQWO\ E\ %URZQ 
0$7(5,$/$1'0(7+2'6 )HPDOHV RI *\PQRSKRUD ZHUH VWXGLHG PRXQWHG RQ VOLGHV DV KDV EHHQ SURSRVHG E\ 'LVQH\  0DOHV ZHUH VRDNHG LQ FROG VROXWLRQ RI .2+ IRU VHYHUDO KRXUV WKHQ ULQVHG LQ ZDWHU DQG DOFRKRO DQG WUDQVIHUUHG WR D GURS RI JO\FHURO IRU IXUWKHU VWXG\ 3HUPDQHQWO\ WKH\ DUH VWRUHG LQ SLQQHG PLFURYLDOV 2FFDVLRQDOO\ PDOHV KDYH EHHQ PRXQWHG RQ VOLGHV VLPLODUO\ WR IHPDOHV 7KH GUDZLQJV ZHUH PDGH LQ WUDQVPLWWHG OLJKW XVLQJ D /HLFD '0/% PLFURVFRSH ZLWK GUDZLQJ DWWDFKPHQW 7HUPLQRORJ\ RI WKH VWUXFWXUHV RI WKH DHGHDJDO FRPSOH[ IRO ORZV %URZQ  DOWKRXJK D GLIIHUHQW RSLQLRQ KDV EHHQ H[SUHVVHG RQ WKH KRPRORJ\ RI VRPH VWUXFWXUHV 'LVQH\  $EEUHYLDWLRQV RI DEGRPLQDO WHUJLWHV DQG VWHUQLWHV DSSHDU DV 7

*\PQRSKRUD DPXUHQVLV VS Q )LJV ± 'LDJQRVLV 6WURQJ PHGLRODWHUDOV DEVHQW 1RWRSOHXUDO ULGJH SDOH EURZQ &RVWD ZLWKRXW DEUXSW WKLFNHQLQJ EHIRUH 5 3RVWHULRU PDUJLQ RI OHIW VLGH RI HSDQGULXP ZLWKRXW OREHV RU LQFLVLRQV /HIW VLGH RI K\SDQGULXP ZLWK VPDOO SURFHVV
(W\PRORJ\ )URP $PXU UHJLRQ

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


are remarkably longer in posterior half of T6. Posterior margin of left side of epandrium oblique, with bristles arranged in row on prominent ridge (Fig. 4). Right side of epandrium with knob-like posteroventral lobe bearing bristles and curly hairs (Fig. 5). Left side of hypandrium with small scoop-like process (Fig. 6), right side of hypandrium simple. Ring of basiphallus somewhat elongated (Fig. 7). Cerci moderately long. Hypoproct somewhat elongated.
Type material. Holotype, k, Russia, Amur Region, Zeya, 30.vii.1978 (A. Shatalkin) (ZMMU).

Gymnophora arcuata (Meigen, 1830) (Figs 8, 9, 65) Diagnosis. Mediolaterals differ from adjacent hairs, but much weaker than bristles on vertex. Notopleural ridge pale. Costa abruptly thickened before R1, with clear spot. Wing length of specimens from European Russia 2.9­3.9 mm, that of West European specimens 2.4­3.8. Mean costal index 0.51, range 0.49­0.54. Mean costal sections ratio 6.38:1.40:1, range 5.30­7.44:1.06­1.69:1. Left side of epandrium with small anteroventral lobe (Fig. 8). Left side of hypandrium with long process, which is greatly swollen at mid-length (Fig. 9). Female with T4 and T5 entirely absent, sclerotised glands absent. Apex of female abdomen with paired ventral sclerites, which are somewhat elongate and bear numerous (mean 30, range 24­37) setae grouped mainly along outer margin of sclerite (Fig. 65). Cerci on average 6.3 times as long as broad, range 6­7.
Type material examined. Lectotype (here designated), l, rectangle printed label "Museum Paris coll. Meigen", small disc "2892 40" and "Meigen" on reverse side [handwritten], rectangle label "Phora arcuata l" [handwritten], rectangle label "arcuata Becker vidit 1898", and small green rectangle "98". Paralectotype, l, small disc "2892 40" and "Meigen" on reverse

Figs 1­3. Thorax and wing details of Gymnophora. 1 ­ left face of thorax of G. healeyae, showing dark notopleural ridge; 2 ­ wing of G. verrucata; 3 ­ wing base of G. victoria sp. n. Abbreviations: S ­ anterior spiracle, M ­ mesopleuron. Scale: 0.1 m in Figs 1, 3; 1 mm in Fig. 2. (Fig. 1 courtesy of Dr. R.H.L. Disney)

Figs 4­9. 4­7 ­ Details of hypopygium of G. amurensis sp. n.: 4 ­ left face of hypopygium; 5 ­ right face of epandrium; 6 ­ process of left side of hypandrium; 7 ­ ring of basiphallus; 8­9 ­ G. arcuata, left face of hypopygium and process of left side of hypandrium. Scale: 0.1 mm.

154


VLGH >KDQGZULWWHQ@ UHFWDQJOH ODEHO ¨3KRUD DUFXDWD l´ >KDQGZULW WHQ@ DQG VPDOO JUHHQ UHFWDQJOH ¨´ 2WKHU PDWHULDO H[DPLQHG (QJODQG l 6XUUH\ 6WHSSLQJ 6WRQHV YLL l /HLFHVWHUVKLUH 6ZLWKODQG :RRG YLL 3 &KDQGOHU k :LQGVRU )RUHVW YLLL 3 &KDQGOHU l ,VOH RI :LJKW :DOOHU¶V &RSVH YLL 3 &KDQGOHU k 1RUWK
.DPFKDWND NP 12 .R]\UHYVN YLL =ORELQ =,1 k $PXU 5HJLRQ =H\D YLL $ 6KDWDONLQ k 6RXWKHUQ 0DULWLPH 7HUULWRU\ NP 6( RI 8VVXULMVN YLL $ 2]HURY =008 k 0DULWLPH 7HUULWRU\ NP 6( RI 8VVXULMVN *RUQRWD\R]KQRH ±YL DQG ±YLL 0 0LFKDLORYVND\D 090 'LVWULEXWLRQ 5XVVLD (DVW 6LEHULD -DSDQ 1RUWK $PHULFD

*\PQRSKRUD JRUQRVWDHYL VS Q )LJV ±  'LDJQRVLV 0HGLRODWHUDOV ZHDN 1RWRSOHXUDO ULGJH GDUN &RVWD DEUXSWO\ WKLFNHQHG DQG ZLWK FOHDU VSRW 3RVWHULRU PDUJLQ RI OHIW VLGH RI HSDQGULXP ELOREHG +\SRSURFW ORQJHU WKDQ FHUFL ZLGHQHG GLVWDOO\ )HPDOH DEGRPHQ ZLWK 7 DEVHQW DQG 7 JUHDWO\ UHGXFHG 6FOHURWLVHG JODQGV FOHDUO\YLVLEOHYHQWUDODSLFDOVFOHULWHVSDLUHG
(W\PRORJ\ 7KLV VSHFLHV LV QDPHG LQ WKH PHPRU\ RI 'U *1 *RUQRVWDHY 0RVFRZ 6WDWH 8QLYHUVLW\

&RPPHQWV 7KH JUHDWO\ VZROOHQ SURFHVV RI WKH OHIW VLGH RI WKH K\SDQGULXP LV WKH PRVW GLVWLQFWLYH IHDWXUH RI WKH PDOHV RI WKLV VSHFLHV +RZHYHU ZH IRXQG WKDW UDUHO\ WKLV SURFHVV LV OHVV VZROOHQ DQG PD\ EH HDVLO\ FRQIXVHG ZLWK WKDW RI * SHUSURSLQTXD VS Q ,Q WKLV FDVH RQH VKRXOG UHIHU WR WKH YHQWUDO OREH RI WKH OHIW VLGH RI WKH HSDQGULXP ZKLFK LV PXFK VPDOOHU WKDQ LQ * SHUSURSLQTXD VS Q )HPDOHV RI DUFXDWD DUH YHU\ VLPLODU WR WKRVH RI * SHUSUR SLQTXD 0HLJHQ  GHVFULEHG * DUFXDWD EDVHG RQ WZR IHPDOHV EXW GLG QRW GHVLJQDWH WKH KRORW\SH 7R FODULI\ WKH LGHQWLW\ RI WKLV VSHFLHV ZH KDYH UHH[DPLQHG WKH V\QW\SHV DQG GHVLJQDWHG WKH OHFWRW\SH DQG SDUDW\SH 7KH W\SH ORFDOLW\ FDQQRW EH GHWHUPLQHG IURP WKH ODEHO GDWD DOWKRXJK 0DFTXDUW  ZKR HUHFWHG WKH JHQXV *\PQRSKRUD ZLWK * DUFXDWD DV WKH W\SH VSHFLHV QRWHG WKDW WKLV VSHFLHV LV GLVWULEXWHG LQ WKH QRUWKHUQ )UDQFH DQG QRUWKHUQ *HUPDQ\ *\PQRSKRUD IDVWLJLRUXP 6FKPLW] )LJV  'LDJQRVLV 6WURQJ PHGLRODWHUDOV DEVHQW 1RWRSOHXUDO ULGJH GDUN &RVWD ZLWKRXW DEUXSW WKLFNHQLQJ EHIRUH 5 /HIW VLGH RI HSDQGULXP ZLWK QHDUO\ VWUDLJKW RU MXVW VOLJKWO\ FRQFDYH SRVWHULRU PDUJLQ ZLWK DERXW WHQ EULVWOHV DQG ZLWK VPDOO URXQGHG HODERUDWLRQ RQ LQQHU VXUIDFH /HIW VLGH RI K\SDQGULXP ZLWKRXW SURFHVV 0DOH FHUFL VKRUW )HPDOH DEGRPHQ ZLWK 7 DEVHQW 7 DQG 7 UHGXFHG 7 ZDWFKJODVVVKDSHG 7 WULDQJXODU 6FOHURWLVHG JODQGV DEVHQW YHQWUDO DSLFDO VFOHULWH XQSDLUHG
0DWHULDO H[DPLQHG 5XVVLD k %XU\DWL\D 9LWLP 5 %DLVVD ±YLL 0RVWRYVNL 0%0 k 0DJDGDQ 5HJ 6LELW 7\HOODNK $ERULJHQ YLLL $QWRQRYD =008 k

'HVFULSWLRQ %RG\ HQWLUHO\ GDUN EURZQ ERG\ OHQJWK ± 7KLUG DQWHQQDO VHJPHQW EURZQ 3DOSL EURZQ ZLWK ± VWURQJ EULVWOHV ZKLFK DUH FOHDUO\ ORQJHU WKDQ PD[LPXP SDOS EUHDGWK 0HGLRODWHUDOV DW OHDVW WLPHV DV ORQJ DV DGMDFHQW KDLUV EXW UHPDUNDEO\ ZHDNHU WKDQ EULVWOHV RQ YHUWH[ 1RWRSOHXUDO ULGJH GDUN /HJV EURZQ :LQJ OHQJWK ± PP 0HDQ FRVWDO LQGH[ UDQJH ± 0HDQ FRVWDO VHFWLRQV UDWLR UDQJH ±± &RVWD ZLWK DEUXSW WKLFNHQLQJ EHIRUH 5 HQFORVLQJ FOHDU VSRW $[LOODU\ ULGJH ZLWK RQH RU WZR EULVWOHV +DOWHUHV EURZQ :LQJ PHPEUDQH JUH\ EURZQLVK 0DOH $EGRPLQDO WHUJLWHV ZLWK VFDWWHUHG VKRUW KDLUV YHQWHU EDUH H[FHSW VHJPHQW ZKLFK EHDUV ORQJLVK KDLUV DW UHDU 3RVWHULRU PDUJLQ RI OHIW VLGH RI HSDQGULXP ZLWK WZR VPDOO DQG EURDG OREHV XSSHU OREH WULDQJXODU ORZHU RQH URXQGHG )LJ  5LJKW VLGH RI HSDQGULXP QDUURZ ZLWK VHYHUDO UDWKHU VWURQJ EULVWOHV LWV SRVWHULRU PDUJLQ ZLWK GLVWLQFW QRWFK )LJ  %DVLSKDOOXV ZHOO VFOHURWLVHG URXJKO\ URXQGHG )LJ 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 6 GHYHORSHG DV ORQJ QDUURZ VFOHURWLVHG VWULSH 7 EOXQW WULDQJXODU ZLWK VFDWWHUHG VHWDH )LJ  $EGRPLQDO JODQGV VWURQJO\ VFOHURWLVHG &HUFL WLPHV DV ORQJ DV EURDG UDQJH ± $SLFDO VFOHULWHV URXQGHG SDLUHG
7\SH PDWHULDO +RORW\SH k .LUJL]L\D &KDWNDOVNLM 0RXQWDLQ 5LGJH 6DU\&KLOHN 9 / =LPLQD 3DUDW\SHV k l VDPH GDWD k VDPH GDWD H[FHSW =KHORNKRYWVHY l VDPH GDWD H[FHSW $UNLW 9, / =LPLQD $OO W\SHV LQ =008




Figs 10­18. 10 ­ G. fastigiorum, left face of epandrium; 11­13 ­ G. gornostaevi sp. n.: 11 ­ left face of epandrium; 12 ­ right face of epandrium; 13 ­ aedeagal complex; 14­15 ­ G. gotoi, left face of hypopygium and aedeagal complex; 16­17 ­ G. healeyae, left face of hypopygium and process of left side of hypandrium; 18 ­ G. integralis, left face of epandrium. Scale: 0.1 mm. (Figs 14, 15 courtesy of Dr. B.V. Brown, Fig. 16 courtesy of Dr R.H.L. Disney)

Gymnophora gotoi Brown, 1988 (Figs 14, 15, 38, 51) Diagnosis. Strong mediolaterals absent. Notopleural ridge dark. Costa gradually thickened and without a clear spot. Left side of epandrium with notably drawn posterior margin and thin and narrow ventral process. Ring of basiphallus massive, with small aperture. Left side of hypandrium with small broad process. Male cerci somewhat elongate. Female T7 wedge-shaped, with massive proximal portion, T8 elongate with somewhat tapered posterior end and several hairs. Sclerotized glands absent. Abdominal venter with paired apical sclerites.
Material examined. Russia: 3l, Maritime Territory, 18 km SE Ussurijsk, Gornotaezhnoe, 17­22.vii.1996 and 28­31.vii.1996 (M. Michailovskaya) (MVM). Distribution. Russian Far East, Japan.

Gymnophora healeyae Disney, 1980 (Figs 1, 16, 17, 39, 52) Diagnosis. Mediolaterals developed but clearly shorter than bristles on vertex, especially in females. Notopleural ridge dark. Costa abruptly thickened before R1, but without clear spot. Left side of epandrium with small posteroventral lobe and several bristles along posterior margin. Left side of hypandrium with prominence. Female abdomen with T3, but T4 and T5 reduced, abdominal glands poorly sclerotised, abdominal venter with paired apical sclerites.
Type material examined. Holotype, 1k, England, Bear Park, 10.viii.1978 (Coulson). Paratypes, 3l, Rowantree Scar, Yorkshire, 23.ix.1976 (Coulson), 1l, Salt Lake Quarry, North Yorkshire, 1.vii.1977 (Disney), 1l Waskerley Beck, 30.vii.1976 (Coulson). The types housed in CMZ.

156


Figs 19­26. 19­20 ­ G. multipinnacula, left face of epandrium and details of aedeagal complex; 21 ­ G. nigripennis, left face of hypopygium; 22­24 ­ G. pararcuata: 22­23 ­ let and right faces of epandrium; 24 ­ process of left side of hypandrium; 25­26 ­ G. perpropinqua sp. n., left face of hypopygium and process of left side of hypandrium. Scale: 0.1 mm. (Fig. 21 courtesy of Dr. B.V. Brown) Other material examined. England: 1k, 2l, Dun Fell 1700', 2.viii.1978 (Coulson), 1k, Newby Head, 11.viii.1978 (Coulson) (CMZ). Distribution. England.

Gymnophora integralis Schmitz, 1920 (Figs 18, 40, 41, 53) Diagnosis. Mediolaterals at most just little longer than other hairs on frons in both sexes. Notopleural ridge dark. Costa with abrupt thickening, enclosing clear spot. Posterior margin of left side of epandrium notably convex and fringed with bristles and hairs. Left side of hypandrium with large parallel-sided process. Hypoproct longer than cerci. Female abdomen with all tergites fully developed, a pair of sclerotised glands that discharge into a common chamber, whose opening to the exterior is enlarged and transversely elongated. Abdominal venter with paired apical sclerites.
Material examined. Romania: 1l, Banat, env. de-Baine Herculane, 6.x.1968 (L. Matile) (MNHN); Russia: 1l, Belgorod Region, 20­30.iv.1987 (V. Kolyada) (MBM), 32k, 15l, Maritime Territory, 18 km SE of Ussurijsk, Gornotayozhnoe, 30.iv­3.vi.1996 (M. Michailovskaya) (MVM). Additionally, one slide-mounted male from Schmitz' collection (ZFMK) has been studied and figured; this slide has no data labels. Distribution. Europe, Russian Far East, Japan. This may turn to be transpalaearctic species.

Diagnosis. Mediolaterals distinctive but remarkably smaller than bristles on vertex. Notopleural ridge dark. Costa with abrupt thickening before R1 and encloses clear spot. Left side of epandrium with irregular posterior margin and bristles concentrated in the lower part. Left side of hypandrium without process. Inner left arm of aedeagus with five to six peaks, which are clearly visible on prepared hypopygium. Female abdomen with T3 reduced, T4 and T5 absent, abdominal glands sclerotised and clearly visible. Abdominal venter with paired apical sclerites.
Material examined. Russia: 1k, Amur Region, Zeya, 5.vii.1978 (Belov), 1k, Maritime Territory, 40 km SE of Ussurijsk, 26.viii.1983 (A. Ozerov) (ZMMU), 11k 17l, Maritime Territory, 18 km SE of Ussurijsk, Gornotayozhnoe, 25.vii. and 18­31.viii.1996 (M. Michailovskaya) (MVM). Most of specimens were caught in copula. Distribution. Russian Far East, Japan.

Gymnophora nigripennis Schmitz, 1926 (Figs 21, 43, 55, 56, 66, 67) Diagnosis. Mediolaterals at most just slightly differ from adjacent hairs. Notopleural ridge dark. Costa gradually thickened before R1. Posterior margin of left side of epandrium oblique and has small elaboration. Left side of hypandrium without process. Male cerci short. Females with abdominal T4 absent. T3 and T5 developed but obviously less than T2 or T6, sometimes reduced to stripe or small patches or developed as semicircular sclerites. T7 narrow wedge-shaped, T8 elongate triangular to pentagonal. Abdominal venter with single apical sclerite,

Gymnophora multipinnacula Brown, 1987 (Figs 19, 20, 42, 54) Gymnophora verrucata Michailovskaya, 1997: 3 non Schmitz, 1927 (misident.).

157


Figs mollis, below R.H.L.

27­35. 27­28 ­ G. prescherweberae, left face of hypopygium and right side of epandrium from below; 29­30 ­ G. quartoleft face of epandrium and bifurcate process; 31­33 ­ G. verrucata, left and right faces of epandrium and hypandrium from respectively; 34­35 ­ G. victoria sp. n., left face of epandrium and distiphallus. Scale: 0.1 mm. (Fig. 27 courtesy of Dr. Disney)

bearing fan of long setae and short hairs between them. Sclerotised glands absent. Comments. The recognition of this species is probably the most troublesome in the genus. Females of this species demonstrate wide variation of the shape of their tergite 8 (Figs 55, 56) and tergites 3 and 5. The female from the northern Maritime Territory seems to be the closest to European specimens, as evidenced by the shape of its abdominal tergites, the shape of the internal sclerotised ring, which is not bent, and the straight hairs between the setae on the apical sclerite of the abdomen. Other specimens from the Far East and China collected by Licent show other states of these characters. Their T5 developed as a semicircular sclerite (except for the specimen from Kamchatka, whose T5 is greatly reduced), the internal sclerotised ring is more or less bent, and the hairs on the apical sclerite is not straight but somewhat curly (Fig. 67). Chinese specimen collected by Przewalsky has the internal sclerotised ring being not bent and straight hairs on the apical sclerite, besides the semicircular T5. However, at the present state of knowledge all these differences seem to be insufficient to put these females as a separate species now, especially without a confident association with males. Brown (1988) reported some other unidentified females of G. nigripennis-subgroup from Japan, and female of G. nepalensis is still unrecognised 158

(Brown, 1988). On balance, we decided to keep these doubtful specimens under the name of G. nigripennis, but the reader should bear in mind that they may in fact belong to other species. The female from the Kirov Region has been found to be filled with fungal hyphae that penetrate the whole body including the head and legs. This should be of particular interest, since this specimen was caught alive.
Type material examined. Holotype, l, "Museum Paris coll. Meigen" [printed], small disc "2875 40" [handwritten], "Phora nigripennis" [handwritten], "86" [pale bown rectangular label]. The holotype is housed in MNHN. Other material examined. Russia: 1l, "Gouv. Petrograd, Tobzhitsa, 30.vi.1934" (Shtakelberg) (ZIN), 1l, Vologda Region, Kich-Gorodetskij Distr., Sud. Gora village, 22.vii.1999 (Mostovski), 1l, Kirov Region, Yug R., Bolsheromanovo, 4­6.viii.1994 (Mostovski) (MBM), 1l, Amur Region, Zeya, 26.vii.1979 (A. Shatalkin) (ZMMU), 1k, 1l, Kamchatka, Kamchatka R., Kamaki settlement, 9.vii.1909 (A. Derzhavin) (ZIN); China: 1l, "montes Gan-ssu, China occid., Przewalsky, viii.[18]71" and "77442", 1l, "Mae eull ting, 1.vii.[19]33, Licent", 1l, "Mahochan, 15.vii.[19]18, Kansou O., Licent" (MNHN). Distribution. Europe, Far East, China.

Gymnophora pararcuata Brown, 1988 (Figs 22­24, 44, 57)


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
7\SH PDWHULDO H[DPLQHG 3DUDW\SH k -DSDQ )XNXRND 0W +LNRVDQ [ . 7DNHQR /$&0 2WKHU PDWHULDO H[DPLQHG 5XVVLD l $PXU 5HJLRQ =H\D L[ $ 6KDWDONLQ =008 k l 0DULWLPH 7HUULWRU\ NP 6( RI 8VVXULMVN *RUQRWD\R]KQRH ±YLL 0 0LFKDLORYVND\D 090 l 0DULWLPH 7HUULWRU\ /D]R 5HVHUYH L[ 0%0 'LVWULEXWLRQ 5XVVLDQ )DU (DVW -DSDQ

ZHGJHVKDSHG )LJ  6 IDLQW RU DEVHQW 7 ZDWFKJODVVVKDSHG )LJ  $SLFDO VFOHULWHV URXQGHG ZLWK VHWDH UDQJH ± ZKLFK DUH PRUH GLVSHUVHG DQG QRW FRQILQHG WR RXWHU PDUJLQ RI VFOHULWH )LJ  &HUFL RQ DYHUDJH WLPHV DV ORQJ DV EURDG UDQJH ± 6FOHURWLVHGDEGRPLQDOJODQGVDEVHQW
7\SH PDWHULDO +RORW\SH k 8NUDLQH &ULPHD %DNKFKLVDUDM 'LVWU .D\DV']KLOJD YL 0RVWRYVNL =008 3DUD W\SHV 8NUDLQH k l &ULPHD $OXVKWD &KHO¶PHQFKL 5 Y 0RVWRYVNL k l &ULPHD 1LNLWD %RWDQ *DUGHQ Y 0RVWRYVNL 0%0 k VDPH GDWD DV KRORW\SH &0= VDPH GDWD DV KRORW\SH H[FHSW YLL l VDPH GDWD DV KRORW\SH H[FHSW YLL k l 6: &ULPHD 6HO¶EXNKUD 0W YL .RO\DGD k VDPH GDWD H[FHSW YL .RO\DGD =008 k VDPH GDWD .RO\DGD /$&0 k VDPH GDWD .RO\DGD =)0. k VDPH GDWD .RO\DGD =,1 l &ULPHD %DNKFKLVDUDM 'LVWU QU 3URNKODG QRH .L]LO&KLJLU 0RVWRYVNL =008 k &ULPHD %DNKFKLVDUDM 'LVWU 3URNKODGQRH ZLQGRZ YL 0RV WRYVNL k l &ULPHD %DNKFKLVDUDM 'LVWU .HUWPHO¶VNLM RDN IRUHVW YL 0RVWRYVNL k &ULPHD %DNKFKLVDUDM 'LVWU $NVX'HUH YL 0RVWRYVNL k l &ULPHD %DNKFKLVDUDM 'LVWU QU 3URNKODGQRH YL 0RVWRYVNL 0%0 l VDPH GDWD &0= l VDPH GDWD /$&0 l VDPH GDWD =)0. l VDPH GDWD =,1 5XVVLD k 0RVFRZ 5HJ QU 6WXSLQR YLL .RO\DGD k l .LURY 5HJ -XJ 5 %ROVKHURPDQRYR±YLLL 9 .RO\DGD k l VDPH GDWD H[FHSW ±YLLL 0RVWRYVNL 0%0

*\PQRSKRUD SHUSURSLQTXD VS Q )LJV 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± VHWDH WKDW DUH QRW GHQVHO\ DUUDQJHG DORQJ RXWHU PDUJLQ 6FOHURWLVHGDEGRPLQDOJODQGVDEVHQW
(W\PRORJ\ )URP /DWLQ SHUSURSLQTXD YHU\ VLPLODU

'HVFULSWLRQ %RG\ OHQJWK ± PP 7KLUG DQWHQQDO VHJPHQW EURZQ 3DOSL EURZQ QRW EURDGHQHG ZLWK VHYHUDO VHWDH ZKLFK DUH FOHDUO\ ORQJHU WKDQ PD[LPXP SDOSXV EUHDGWK 0HGLRODWHUDOV DW OHDVW WLPHV VKRUWHU WKDQ EULVWOHV RQ YHUWH[ 1RWRSOHXUDO ULGJH SDOH /HJV \HOORZLVK EURZQ :LQJ OHQJWK ± PP 0HDQ FRVWDO LQGH[ UDQJH ± 0HDQ FRVWDO VHFWLRQV UDWLR UDQJH ±± &RVWD ZLWK DEUXSW WKLFNHQLQJ EHIRUH 5 ZKLFK HQFORVHV FOHDU VSRW $[LOODU\ ULGJH ZLWK WZR WR WKUHHEULVWOHV+DOWHUHV\HOORZ 0DOH $EGRPLQDO YHQWHU EDUH ZLWK H[FHSWLRQ RI ODVW VHJPHQW ZKLFK EHDUV H[WUHPHO\ VKRUW VFDWWHUHG KDLUV DW UHDU 7HUJLWHV ZLWK GLVSHUVHG VKRUW KDLUV +\SRS\JLXP VLPLODU WR WKDW RI * DUFXDWD +RZHYHU HSDQGULXP ZLWK VWURQJO\ GHYHORSHG YHQWUDO OREH )LJ  DQG K\SDQGULDO SURFHVV RQO\ VOLJKWO\ VZROOHQ DW PLGOHQJWK )LJ  &HUFL PRGHUDWHO\ ORQJ +\SRSURFW VKRUW )HPDOH $OO DEGRPLQDO VHJPHQWV ZLWK VFDWWHUHG KDLUV ZKLFK DUH ORQJHU RQ ± VHJPHQWV HVSHFLDOO\ DORQJ KLQG PDUJLQ 9HQWHU ZLWK ORQJLVK KDLUV DORQJ KLQG PDUJLQ RI VHJPHQWV DQG 7 WUDSH]RLGDO 7 DQG 7 DEVHQW 7

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 *\PQRSKRUD SUHVFKHUZHEHUDH 'LVQH\ )LJV  *\PQRSKRUD ODFLQLDWD 0LFKDLORYVND\D V\Q Q 'LDJQRVLV 0HGLRODWHUDOV QRWDEO\ VWURQJ DOWKRXJK WKH\ DUH REYLRXVO\ ZHDNHU WKDQ WKRVH RQ YHUWH[ 1RWRSOHXUDO ULGJH GDUN &RVWD ZLWKRXW DEUXSW WKLFNHQLQJ EHIRUH 5 3RVWHULRU PDUJLQ RI OHIW VLGH RI HSDQGULXP ZLWKRXW DEUXSW H[FDYDWLRQ RU HODERUDWLRQ SURFHVV RI ULJKW VLGH RI HSDQ


Figs 36­49. Female tergite 7 of Gymnophora spp. 36 ­ G. fastigiorum; 37 ­ G. gornostaevi sp. n.; 38 ­ G. gotoi; 39 ­ G. healeyae; 40 ­ G. integralis; 42 ­ G. multipinnacula; 43 ­ G. nigripennis; 44 ­ G. pararcuata; 45 ­ G. perpropinqua sp. n.; 46 ­ G. prescherweberae; 47 ­ G. victoria sp. n.; 48 ­ G. sp. A; 49 ­ G. sp. B; 41 ­ G. integralis, opening of abdominal glands between T5 and T6. Scale: 0.1 mm. (Figs 36, 38 courtesy of Dr B.V.Brown, Fig. 39 courtesy of Dr . R.H.L. Disney)

drium with notch near its tip when viewed from below, and with bristles arranged in row. Left side of hypandrium without process. Cerci short. Female abdomen with T3­5 reduced, T7 approx. as long as broad, with numerous setae, T8 narrow wedge-shaped, with concave sides. Abdominal glands weakly sclerotised. Abdominal venter with paired apical sclerites, which are small and somewhat rounded.
Type material examined. Holotype, k, Thuringia, Appfelstadter Ried, grassland, 26.ix.1985 (J. Weipert) (CMZ). Paratypes. l, same data as holotype, k, l, same data as holotype except 1.vii.1985 (CMZ). Other material examined. Russia: 1l, M. Khekhtsir, 25 km S of Khabarovsk, 31.viii.1979 (Nartchuk) (ZIN), 6k, Maritime Prov., 18 km SE Ussurijsk, Gornotaezhnoe, 12­12.vii and 24­26.vii.1996 (M. Michailovskaya) (IBP). Distribution. Europe, Russian Far East.

of both left and right sides of the epandrium and the pattern of setation support the newly proposed synonymy. Gymnophora quartomollis Schmitz, 1920 (Figs 29, 30, 60) Diagnosis. In males, mediolaterals clearly differ from adjacent hairs, but at least 2 times weaker than bristles on vertex, mediolaterals absent in females. Notopleural ridge dark. Costa gradually thickened before R1. Left side of epandrium with excavated posterior margin, with three to four bristles, and with small triangle elaboration on inner surface. Left side of hypandrium without process. Female abdomen with T4 absent and T3 reduced, T7 long, narrow, triangular, T8 pentagonal. Paired sclerotised glands absent, apical sclerite single.
Material examined. Ukraine: 1k, 1l, Crimea, Bakhchisaraj Distr., Sel'bukhra Mt., 5­6.vi.1995 (Kolyada), 1l, same data except 15.vi.1995, 2l, Crimea, Bakhchisaraj Distr., Nauchnyj settlement, Kayas-Dzhilga, 23.vi.1995 (Mostovski) (ZMMU), 1k, Crimea, Bakhchisaraj Distr., Aksu-Dere, 25.vi.1995 (Mos-

Comments. This species was described nearly simultaneously from Germany and Russian Far East. The shape

160


WRYVNL k l &ULPHD %DNKFKLVDUDM 'LVWU .L]LO&KLJLU YL 0RVWRYVNL 0%0 5XVVLD k /HQLQJUDG 5HJ YLF /XJD 7ROPDWVFKHYR YLL 6WDFNHOEHUJ l /HQLQ JUDG 5HJ /XJD YLLL 6WDFNHOEHUJ l /HQLQJUDG 5HJ
KDV UHDSSHDUHG LQ +HOVLQNL DQG LV GHVLJQDWHG KHUH DV WKH OHFWRW\SH DQG UHGHVFULEHG 8QIRUWXQDWHO\ ZH FDQQRW ILQG DQ\ WUDFHV RI WKH IHPDOH WKH RWKHU V\QW\SH VSHFLPHQ WR GHVLJQDWHLWDVWKHSDUDOHFWRW\SH *\PQRSKRUD YLFWRULD VS QRY )LJV  'LDJQRVLV 5DWKHU VWURQJ PHGLRODWHUDOV SUHVHQW LQ PDOHV EXW DEVHQW IURP IHPDOHV 1RWRSOHXUDO ULGJH GDUN &RVWD ZLWKRXW DEUXSW WKLFNHQLQJ /HIW VLGH RI K\SDQGULXP ZLWK YHQWUDO OREH 'LVWLSKDOOXV LQGHQWHG )HPDOH 7 ZLGH ZHGJHVKDSHG 7 FUHVFHQW $EGRPLQDO YHQWHU ZLWK SDLUHG VFOHULWHV
(W\PRORJ\ )URP /DWLQ YLFWRULD ¨YLFWRU\´ WR FRPPHPRUDWH WKH \HDU RI WKH YLFWRU\ LQ WKH :RUOG :DU ,, ZKHQ RQH RI WKH SDUDW\SHV ZDV FDXJKW

*\PQRSKRUD YHUUXFDWD 6FKPLW] )LJV ± 'LDJQRVLV 6WURQJ PHGLRODWHUDOV DEVHQW 1RWRSOHXUDO ULGJH SDOH &RVWD ZLWK DEUXSW VZHOOLQJ DQG ZLWK FOHDU VSRW /HIW VLGH RI HSDQGULXP ZLWK REOLTXH SRVWHULRU PDUJLQ DQG SDWFK RI VKRUW EULVWOHV /HIW VLGH RI K\SDQGULXP ZLWK HORQ JDWH SURFHVV 5HGHVFULSWLRQ %RG\ OHQJWK PP 3DOSL GDUN ZLWK SDOHU VWHP 0HGLRODWHUDOV LQGLVWLQFW IURP RWKHU KDLUV 1RWRSOHXUDO ULGJH SDOH /HJV \HOORZLVK :LQJ OHQJWK PP )LJ  &RVWDO LQGH[ &RVWDO VHFWLRQV UDWLR &RVWD ZLWK DEUXSW WKLFNHQLQJ EHIRUH 5 WLS WKDW HQFORVHV FOHDU VSRW :LQJ PHPEUDQH VKDGRZHG WLQJHG DORQJ IRUH PDUJLQ DQG WR OHVV GHJUHH DORQJ ILUVW DQG VHFRQG WKLQ YHLQV +DOWHUHV SDOH \HOORZLVK /HIW VLGH RI HSDQGULXP ZLWK SRVWHURGRUVDO EXQFK RI EULVWOHV ZKLFK DUH QRW ORQJHU WKDQ ORQJHVW EULVWOHV RQ FHUFL 3RVWHULRU PDUJLQ RI OHIW VLGH RI HSDQGULXP REOLTXH ZLWKRXW LQFLVLRQV RU OREHV )LJ  5LJKW VLGH RI HSDQGULXP ZLWK GHHS SRVWHULRU QRWFK DQG ODWHURYHQWUDO OREH EHDULQJ IDVFLFOH RI ORQJ DQG VKRUWHU EULVWOHV )LJ  /HIW K\SDQGULDO OREH ZLWK PRGHUDWHO\ ORQJ SURFHVV RQO\ VOLJKWO\ QDUURZHG DSLFDOO\ )LJ  $HGHDJXV HQODJHG FRPSOH[ %DVLSKDOOXV PDVVLYH ZLWK VPDOO DSHUWXUH $QWHULRU OREH RI GLVWLSKDOOXV EURDG DQG FXUYHG UHVHPEOLQJ WKDW RI * SDUDUFXDWD LQ UHODWLYH SRVL WLRQ WR EDVLSKDOOXV EXW PRUH PDVVLYH &HUFL VRPHZKDW HORQJDWH +\SRSURFW VKRUW
7\SH PDWHULDO H[DPLQHG /HFWRW\SH KHUH GHVLJQDWHG k 5XVVLD ZLWK ODEHOV ¨8VVXUL 6SDVVNDMD ,; < :XRUHQ WDXV´ ¨*\PQRSKRUD YHUUXFDWD 6FKPLW] k +RORW\SH´ >6FKPLW]¶ KDQGZULWLQJ@ DQG ¨0XV =RRO + IRUV 6SHF W\S 1R *\PQRSKRUD YHUUXFDWD 6FKPLW]´ 7KH OHFWRW\SH LV KRXVHG LQ WKH =08+ 2WKHU PDWHULDO H[DPLQHG 5XVVLD k $PXU 5HJLRQ NP : RI 6YRERGQ\M .OLPRXWV\ RDN IRUHVW L[ =LQRYLHY =,1 'LVWULEXWLRQ 5XVVLDQ )DU (DVW

'HVFULSWLRQ %RG\ OHQJWK ± PP 7KLUG DQWHQQDO VHJPHQW EURZQ 3DOSL EURZQ VOLJKWO\ EURDGHQHG ZLWK VHYHUDO VKRUW DSLFDO VHWDH ZKLFK DUH DW PRVW OLWWOH ORQJHU WKDQ PD[LPXP SDOSXV EUHDGWK 1RWRSOHXUDO ULGJH GDUN /HJV \HOORZLVK EURZQ :LQJ OHQJWK ± PP 0HDQ FRVWDO LQGH[ UDQJH ± 0HDQ FRVWDO VHFWLRQV UDWLR UDQJH ±± &RVWD JUDGXDOO\ DQG RQO\ VOLJKWO\ WKLFNHQHG EHIRUH 5 WLS )LJ  $[LOODU\ ULGJH ZLWK WZR UDUHO\ WKUHH RU IRXU ILQH EULVWOHV +DOWHUHV JUH\ EURZQ 0DOH $EGRPLQDO WHUJLWHV ZLWK VFDUFH VKRUW KDLUV DEGRPLQDO YHQWHU EDUH ZLWK H[FHSWLRQ RI ODVW VHJPHQW ZKLFK EHDUV VKRUW KDLUV DW UDUH /HIW VLGH RI HSDQGULXP ZLWK VWUDLJKW SRVWHULRU PDUJLQ ZLWK EURDG WULDQJOH YHQWUDO SURFHVV EHDULQJ SRRUO\ GLVWLQFW WXEHUFOH )LJ  5LJKW VLGH RI HSDQGULXP VLPLODU WR * SDOPXOD VHH %URZQ  +\SDQGULXP QHDUO\ V\PPHWULFDO IURP EHORZ 'LV WLSKDOOXV ORQJ ZHOO VFOHURWLVHG IRRWOLNH ZLWK REWXVH GHQWV DORQJ LWV SRVWHULRU PDUJLQ )LJ  &HUFL DQG K\SRSURFW HORQJDWH )HPDOH $OO DEGRPLQDO WHUJLWHV ZHOO GHYHORSHG ZLWK VFDWWHUHG ILQH KDLUV 7 ODUJH ZHGJHVKDSHG EURDGHQHG LQ DQWHULRU SRUWLRQ ZLWK IHZ ORQJLVK VHWDH )LJ  6 WULDQJXODU RU ZDWFKJODVVVKDSHG VRPHWLPHV GHYHORSHG DV EURDG PRGHUDWHO\ VFOHURWLVHG VWULSH 7 VPDOO FUHVFHQW ZLWK YHU\ IHZ VHWDH )LJ  9HQWHU RI DEGRPLQDO VHJ PHQW ZLWK WZR VPDOO KDLU\ VFOHULWHV $EGRPLQDO JODQGV ZHOO VFOHURWLVHG SDLUHG &HUFL WLPHV DV ORQG DV EURDG UDQJH ± RIWHQ VOLJKWO\ SRLQWHG $SLFDO VFOHULWHV URXQGHG UHFWDQJXODU
7\SH PDWHULDO +RORW\SH k .LUJL]L\D %DWNHQ 'LVWULFW 0DG\JHQ Y 3DQILORY 3ULW\NLQD =008 3DUDW\SHV 7XUNPHQL\D k l .RSHW'DJ )LU\X]D JRUJH LY =KHORNKRYWVHY l %DGNK\] $NDU&KHVKP\ LY 9 .RYDOHY =008 8]EHNLVWDQ k l 7DVKNHQW DU\N %RVVX LY 'REU]KDQVNLM .LUJL]L\D l ¨)UXQ]H 3LVKSHN 6HPLU Y´ / =LPLQ 7DMLNLVWDQ k .RQ GDUD JRUJH 9DU]RED YLOODJH P DOW Y *XVVDN RYVNLM k 6WDOLQDEDG *XO¶ELVWD 5 YDOOH\ LY 6WDFN HOEHUJ k l 6WDOLQDEDG IRRWKLOOV LY l VDPH GDWD LY k VDPH GDWD Y *XVVDNRYVNLM =,1 'LVWULEXWLRQ 0LGGOH $VLD

&RPPHQWV 7KH RULJLQDO GHVFULSWLRQ RI WKLV VSHFLHV LV UDWKHU YDJXH DQG LQVXIILFLHQW WR UHFRJQLVH LW DPRQJ RWKHU VSHFLHV 6FKPLW]  QRWHG WKDW KH EDVHG LW RQ WZR VSHFLPHQV D PDOH DQG D IHPDOH EXW GLG QRW GHVLJQDWH WKH KRORW\SH FOHDUO\ )RU D ORQJ WLPH WKH IHPDOH ZDV FRQVLG HUHG DV WKH KRORW\SH DQG WKH PDOH ZDV WKRXJKW WR EH ORVW %URZQ 0LFKDLORYVND\D  0HDQWLPH WKH PDOH ZLWK WKH RULJLQDO ODEHOV DQG 6FKPLW]¶V KDQGZULWLQJ




Figs 50­63. Female tergite 8 of Gymnophora spp. 50 ­ G. gornostaevi sp. n.; 51 ­ G. gotoi; 52 ­ G. healyae; 53 ­ G. integralis; 54 ­ G. multipinnacula; 55­56 ­ G. nigripennis; 57 ­ G. pararcuata; 58 ­ G. perpropinqua sp. n.; 59 ­ G. prescherweberae; 60 ­ G. quartomollis; 61 ­ G. victoria sp. n.; 62 ­ G. sp. A; 63 ­ G. sp. B. Scale: 0.1 mm. (Fig. 51 courtesy of Dr. B.V. Brown, Fig. 52 courtesy of Dr R.H.L. Disney)

Gymnophora sp. A (Figs 48, 62, 70, 71) Description Female. Third antennal segment and palpi brown. Palpi just slightly broadened, hairy, with 3 apical and ventroapical bristles, which are 1.5 times as long as maximum palpus breadth, and with numerous shorter setae disposed medially, laterally, and ventrally. Mediolaterals just slightly longer than adjacent hairs. Notopleural ridge dark. Legs yellow brown. Wing length 3.79 mm. Costal index 0.53. Costal sections ratio 9.41:1.47:1. Costa gradually thickened before R1, with unusually long clear spot before R1, it is narrowed abruptly distad of this spot. Axillary ridge with 3 bristles. Halteres yellow. Abdominal T2 broad trapezoidal, T3­6 absent. Segment 5 with few short hairs in middle. Segment 6 with numerous hairs on dorsal face, which are longer along posterior margin; only short hairs developed in row along posterior margin on ventral face. T7 wedge-shaped, with irregular sides (Fig. 48), 162

segment 7 hairy, with hairs being remarkably longer along posterior margin. S7 not developed. T8 generally rectangular, with strongly excavated anterior margin and slightly concave lateral and posterior margins, with numerous setae, which are somewhat longer on posterolateral corners (Fig. 62). Apical sclerites unusually wide and elongate, with short curly hairs arranged along posterior margin (Fig. 70). Cerci abruptly narrowed at midlength (Fig. 71). Abdominal glands small and poorly sclerotised although clearly visible; gland's opening between segments 5 and 6 distinct and bordered anteriorly with shagreened area.
Material examined. Russia: 1l, Maritime Territory, Lazo Reserve, 10.vii.1993 (Belokobyl'sky) (ZIN).

Gymnophora sp. B (Figs 49, 63, 64) Description Female. Third antennal segment and palpi brown. Palpi not broadened, hairy, with 3 apical and ventro-apical bris-


Figs 64­71. Details of female abdomen of Gymnophora spp. 64 ­ G. sp. B, sternite 7; 65­70 ­ apical sclerites: 65 ­ G. arcuata; 66­67 ­ G. nigripennis; 68 ­ G. perpropinqua sp. n.; 69 ­ G. prescherweberae; 70 ­ G. sp. A; 71 ­ left cercus of G. sp. A. Scale: 0.1 mm.

tles, which are at least 2 times as long as maximum palpus breadth, and with numerous shorter setae medially, laterally and ventrally. Mediolaterals similar to other hairs on frons. Notopleural ridge dark. Legs yellowish brow. Wing length 3.36 mm. Costal index 0.48. Costal sections ratio 6.02:1.56:1. Costa gradually thickened before R1. Axillary ridge with 3 bristles. Halteres yellow. Abdominal T2 broad trapezoidal, T3 developed as small rounded sclerite, T4 absent, T5 normally developed, wider than longer, T6 approximately as wide as long, with scattered short hairs in posterior half, T7 broad, parallel-sided (Fig. 49), S7 developed as broad plate (Fig. 64). Segment 7 with scattered hairs, those along posterior margin being 3­4 times as long as others. T8 triangular, with numerous setae in posterior half (Fig. 63). Apical sclerite single, with fan of long setae and numerous shorter hairs, which are arranged mainly along posterior margin. Cerci narrow, 4 times as long as wide, slightly tapered apically. Sclerotised abdominal glands absent.
Material examined. Russia: 1l, Sakhalin Is., nr YuzhnoSakhalinsk, 19.vii.1955 (Violovich) (ZIN). KEY TO THE SPECIES OF GYMNOPHORA 1. ­ 2. ­ 3. ­ 4. ­ Males and females brachypterous . . G. lapidicola (Bezzi) Males and females macropterous . . . . . . . . . . . . . . . . . . . 2 Males . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Females . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Costa abruptly thickened before R 1 (Fig. 2) . . . . . . . . . . . 4 Costa gradually thickened before R 1 (Fig. 3) . . . . . . . . . 11 Left hypandrial process absent . . . . . . . . . . . . . . . . . . . . . 5 Left hypandrial process present, but sometimes may be poorly developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Right side of epandrium with distinct notch (Fig. 12), hypoproct evidently longer than cerci and widened distally (Fig. 11) . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. gornostaevi sp. n. ­ Right side of epandrium without such notch, hypoproct clearly shorter and not widened (Fig. 19) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. multipinnacula Brown

6. Left hypandrial process strongly developed (e.g. Figs 9, 26, 33) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 ­ Left hypandrial process small and sometimes developed only as tubercle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 7. Left hypandrial process swollen at mid-length, notopleural ridge pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 ­ Left hypandrial process not so, notopleural ridge pale or dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ............................................. 9 8. Left hypandrial process greatly swollen (Fig. 9), left side of epandrium with small ventral lobe (Fig. 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. arcuata (Meigen) ­ Left hypandrial process only slightly swollen (Fig. 26), left side of epandrium with larger ventral lobe (Fig. 25) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. perpropinqua sp. n. 9. Left hypandrial process rather broad and parallel-sided, left side of epandrium broader, with bristles and hairs along posterior margin, hypoproct longer than cerci (Fig. 18) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. integralis Schmitz ­ Left hypandrial process narrower and evenly tapered (Fig. 33), left side of epandrium remarkably narrower, with bristles arranged in posterodorsal fascicle (Fig. 31), hypoproct shorter than cerci (Fig. 32) . . . . . . . . G. verrucata Schmitz 10. Notopleural ridge dark (Fig. 1), left side of epandrium with posterolateral elaboration, bristles despersed along posterior margin (Fig. 16), left hypandrial process developed as tubercle at most (Fig. 17) . . . . . . . . . . . G. healeyae Disney ­ Notopleural ridge pale, left side of epandrium oblique, with distinct prominence bearing bristles arranged in bunch (Fig. 22), left hypandrial process more developed (Fig. 24) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. pararcuata Brown 11. Gradual thickening of costa contains elongate clear spot before R1 . . . . . . . . . . . . . . . . . . . . presumed male of sp. A ­ Costa without any clear spot . . . . . . . . . . . . . . . . . . . . . 12 12. Left hypandrial process entirely absent, cerci evidently shor t . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 ­ Left hypandrial process present, cerci longer . . . . . . . . . 13 13. Left side of epandrium elaborated posteriorly, with narrow ventral process, and with dispersed bristles (Fig. 14), basiphallus as Fig. 15 . . . . . . . . . . . . . . . . . . . . G. gotoi Brown

163


± ± ±

±



±

± ± ± ±

± ±

/HIW VLGH RI HSDQGULXP REOLTXH ZLWKRXW DQ\ YHQWUDO SURFHVV DQG ZLWK EULVWOHV DUUDQJHG LQ URZ MXVW EHORZ FHUFL )LJ  EDVLSKDOOXV DV )LJ * DPXUHQVLV VS Q /HIW VLGH RI HSDQGULXP ZLWK VPDOO SRVWHURYHQWUDO HODERUDWLRQ RULHQWHG HLWKHU LQZDUGO\ RU RXWZDUGO\ )LJV  /HIW VLGH RI HSDQGULXP ZLWKRXW VXFK HODERUDWLRQ 3RVWHURYHQWUDO HODERUDWLRQ RI OHIW VLGH RI HSDQGULXP RULHQWHG RXWZDUGO\ SRVWHULRU PDUJLQ RI OHIW VLGH RI HSDQGULXP FOHDUO\ REOLTXH )LJ  * QLJULSHQQLV 6FKPLW] 3RVWHURYHQWUDO HODERUDWLRQ RI OHIW VLGH RI HSDQGULXP RULHQWHG LQZDUGO\ SRVWHULRU PDUJLQ RI OHIW VLGH RI HSDQGULXP OHVV REOLTXH RU QHDUO\ VWUDLJKW 3RVWHULRU PDUJLQ RI OHIW VLGH RI HSDQGULXP FRQFDYH RQO\ WKUHH WR IRXU EULVWOHV DSDUW IURP WLQ\ KDLUV RQ OHIW IDFH RI HSDQGULXP SRVWHURYHQWUDO HODERUDWLRQ VRPHZKDW WULDQJXODU DQG DWWDFKHG IDUWKHU IURP SRVWHULRU PDUJLQ )LJ  ELIXU FDWH SURFHVV )LJ  FOHDUO\ YLVLEOH ZKHQ K\SRS\JLXP YLHZHG IURP OHIW PHGLRODWHUDO EULVWOHV RQ IURQV ZHOO GLIIHU IURP DGMDFHQW KDLUV DOWKRXJK WKH\ DW OHDVW WLPHV DV ZHDN DV WKRVH RQ YHUWH[ * TXDUWRPROOLV 6FKPLW] 3RVWHULRU PDUJLQ RI OHIW VLGH RI HSDQGULXP QHDUO\ VWUDLJKW RU MXVW VOLJKWO\ FRQFDYH OHIW IDFH RI HSDQGULXP ZLWK PRUH QXPHURXV EULVWOHV DSDUW IURP WLQ\ KDLUV SRVWHURYHQWUDO HODERUDWLRQ URXQGHG DQG DWWDFKHG FORVHU WR SRVWHULRU PDUJLQ )LJ  ELIXUFDWH SURFHVV KDUGO\ YLVLEOH LI DHGHDJDO FRP SOH[ QRW UHPRYHG PHGLRODWHUDOV DV ZHDN DV DGMDFHQW KDLUV RQ IURQV * IDVWLJLRUXP 6FKPLW] 3RVWHULRU PDUJLQ RI OHIW VLGH RI HSDQGULXP PRUH RU OHVV FRQ YH[ ZLWKRXW DEUXSW H[FDYDWLRQ RU HODERUDWLRQ )LJ  SURFHVV RI ULJKW VLGH RI HSDQGULXP ZLWK PRUH RU OHVV URXQGHG WLS DQG QRWFK QHDU LWV WLS ZKHQ YLHZHG IURP EHORZ DQG ZLWK EULVWOHV DUUDQJHG LQ VRUW RI URZ )LJ  * SUHVFKHUZHEHUDH 'LVQH\ 3RVWHULRU PDUJLQ RI OHIW VLGH RI HSDQGULXP QHDUO\ VWUDLJKW ZLWK WULDQJOH YHQWUDO OREH DQG VPDOO WXEHUFOH MXVW EHORZ FHUFL )LJ  SURFHVV RI ULJKW VLGH RI HSDQGULXP QRWDEO\ GUDZQ ZLWK QRWFK VKLIWHG IDU PHGLDOO\ IURP LWV WLS ZKHQ YLHZHG IURP EHORZ DQG ZLWK GLVSHUVHG EULVWOHV GLVWLSKDOOXV IRRW VKDSHG KHDYLO\ VFOHURWLVHG DQG LQGHQWHG DW SRVWHULRU PDUJLQ )LJ  * YLFWRULD VS Q 7LS RI DEGRPHQ ZLWK VLQJOH YHQWUDO VFOHULWH 7LS RI DEGRPHQ ZLWK SDLUHG YHQWUDO VFOHULWHV &RVWD ZLWK DEUXSW WKLFNHQLQJ EHIRUH 5 )LJ  &RVWD JUDGXDOO\ WKLFNHQHG EHIRUH 5 )LJ  &RVWDO WKLFNHQLQJ ZLWK FOHDU VSRW )LJ  &RVWDO WKLFNHQLQJ ZLWKRXW FOHDU VSRW RU LI SUHVHQW LV REVFXUH 7 DQG 7 DV )LJV * KHDOH\DH 'LVQH\ 1RWRSOHXUDO ULGJH GDUN )LJ  DEGRPHQ ZLWK VFOHURWLVHG SDLUHG JODQGV 1RWRSOHXUDO ULGJH SDOH DEGRPHQ ZLWKRXW VFOHURWLVHG JODQGV 7± QRUPDOO\ GHYHORSHG LQWHUQDO DEGRPLQDO JODQGV GLV FKDUJH LQWR D FRPPRQ FKDPEHU ZKRVH RSHQLQJ WR WKH H[WH ULRU LV HQODUJHG DQG WUDQVYHUVHO\ HORQJDWHG )LJ  7 ZLWK EULVWOHV DUUDQJHG LQWR WZR JURXSV )LJ  7 FUHVFHQW )LJ  * LQWHJUDOLV 6FKPLW] 7 DOZD\V DEVHQW 7 DQG PD\ EH UHGXFHG RU DEVHQW RSHQLQJ RI LQWHUQDO DEGRPLQDO JODQGV LV PXFK VPDOOHU DQG OHVV FRQVSLFLRXV 7 LV V\PPHWULFDO 7 PRUH DEUXSWO\ QDUURZLQJ UHDUZDUGV )LJ  7 HORQ JDWH URXQGHG WULDQJXODU )LJ  7 QHDUO\ QRUPDOO\ GHYHORSHG * JRUQRVWDHYL VS Q 7 PRUH JUDGXDOO\ QDUURZLQJ )LJ  7 PDLQO\ UHFWDQ JXODU )LJ  7 UHGXFHG * PXOWLSLQQDFXOD %URZQ

7 LV PRUH RU OHVV QRUPDOO\ GHYHORSHG RU MXVW VOLJKWO\ UHGXFHG 7 ZLWK DQWHULRU PDUJLQ EHLQJ DW PRVW MXVW VOLJKWO\ FRQFDYH )LJ  7 V\PPHWULFDO DQG JHQHUDOO\ UHFWDQJXODU )LJ  FOHDU VSRW RQ FRVWD PRUH URXQGHG ± 7 JUHDWO\ UHGXFHG RU HQWLUHO\ DEVHQW DQWHULRU PDUJLQ RI 7 QRWDEO\ FRQFDYH )LJ  7 ZLWK DV\PPHWULFDO DQWHULRU SRUWLRQ )LJ  FOHDU VSRW RQ FRVWD VRPHZKDW HORQJDWHG * SDUDUFXDWD %URZQ 9HQWUDO DSLFDO VFOHULWHV RI DEGRPHQ PRUH URXQGHG ZLWK OHVV QXPEHU RI VHWDH ± ZKLFK DUH PRVWO\ QRW FRQILQHG WR WKH RXWHU PDUJLQ )LJ  FHUFL FD WLPHV DV ORQJ DV EURDG * SHUSURSLQTXD VS Q ± 9HQWUDO DSLFDO VFOHULWHV RI DEGRPHQ PRUH HORQJDWHG ZLWK VHWDH EHLQJ PRUH DEXQGDQW ± DQG PRVWO\ FRQILQHG WR WKH RXWHU PDUJLQ )LJ  FHUFL XVXDOO\ PRUH WKDQ WLPHV DV ORQJ DV EURDG * DUFXDWD 0HLJHQ 7± HQWLUHO\ DEVHQW RQO\ ZLWK KDLUV DQG VHWDH RQ VHJPHQWV DQG FRVWD ZLWK JUHDWO\ HORQJDWH FOHDU VSRW YHQWUDO DSL FDO VFOHULWHV EURDG ZLWK VKRUW EHQW VHWDH )LJ  FHUFL DEUXSWO\ QDUURZHG DW PLGOHQJWK )LJ  7 DQG DV )LJV * VS$ ± $W OHDVW ZHOO GHILQHG 7 LV DOZD\V SUHVHQW FRVWD ZLWKRXW DQ\ FOHDU VSRW 7± QRUPDOO\ GHYHORSHG 7 EURDG )LJ  7 URXJKO\ FUHVFHQW )LJ  * YLFWRULD VS QRY ± 7± DOZD\V VRPHZKDW UHGXFHG 7 QDUURZHU 7 V\P PHWULFDO 7 LV URXJKO\ DV ORQJ DV EURDG ZLWK QXPHURXV VHWDH )LJ  7 DV )LJ YHQWUDO DSLFDO SODWH DV )LJ * SUHVFKHUZHEHUDH 'LVQH\ ± 7 DSSUR[LPDWHO\ WLPHV DV ORQJ DV EURDG ZLWK IHZHU VHWDH )LJ  7 DV )LJ * JRWRL %URZQ 7 URXJKO\ WULDQJXODU )LJV  RU RWKHUZLVH )LJ  EXW QHYHU EURDG SHQWDJRQDO ± 7 EURDG SHQWDJRQDO )LJ  * TXDUWRPROOLV 6FKPLW] 6 GHYHORSHG DV QDUURZ VFOHURWLVHG VWULSH GHYRLG RI KDLUV RU VHWDH ± 6 EURDG ZLWK VHYHUDO VHWDH )LJ  7 DQG 7 DV )LJV * VS % 7 ZDWFKJODVVVKDSHG )LJ  7 EURDG WULDQJXODU VLPLODU WR )LJ * IDVWLJLRUXP 6FKPLW] ± 7 ZHGJHVKDSHG )LJ  7 RWKHUZLVH RU HORQJDWHG LI WUL DQJXODU )LJV  * QLJULSHQQLV 6FKPLW] $&.12:/('*(0(176 :H H[SUHVV RXU VLQFHUH WKDQNV WR (3 1DUWFKXN 93K =DLWVHY =,1 $/ 2]HURY =008 %9 %URZQ /$&0 3 9LONPDD =08+ 0 %D\ODF - &KDU ERQQHO 01+1 % 6LQFODLU =)0. IRU SURYLGLQJ W\SHV DQG FRPSDUDWLYH PDWHULDO LQ WKHLU FDUH DQG WR 9 .RO\DGD 0RVFRZ ZKR SDVVHG KLV FROOHFWLRQ RI SKRULGV IRU VWXG\ 7KH VHQLRU DXWKRU LV JUDWHIXO WR $



SK\ORJHQ\ DQG JHRJUDSKLF KLVWRU\ 6\VW (QWRPRO ± %52:1 %9 7HVWLQJ SK\ORJHQHWLF K\SRWKHVHV QHZ VSH FLHV RI *\PQRSKRUD 0DFTXDUW 'LSWHUD 3KRULGDH IURP -DSDQ DQG 1HSDO ZLWK DQ XSGDWHG ZRUOG FODVVLILFDWLRQ &DQ - =RRO ± %52:1 %9 1HZ VSHFLHV DQG UHFRUGV RI *\PQRSKRUD 0DFTXDUW 'LSWHUD 3KRULGDH IURP 6RXWKHDVW $VLD &RQWULEX WLRQV LQ 6FLHQFH 1DW +LVW 0XV /RV $QJHOHV &RXQW\ ± ',61(< 5+/ $ QHZ VSHFLHV RI *\PQRSKRUD 0DFTXDUW 'LSWHUD 3KRULGDH IURP
* $ 7D[RQRPLF 6WXG\ RI &KLQHVH SKRULG IOLHV 'LS WHUD 3KRULGDH 1RUWKZHVWHUQ $JULFXOWXUDO 8QLY ,8

/

5HFHLYHG 2FWREHU UHYLVHG -DQXDU\ DFFHSWHG 2FWREHU