A preliminary investigation of some morphological characters in adult females of the simulium damnosum complex Togo

I J1r I I 2 3 4 5 6 ? 8 9 l0 1l t2 13 l4 l5 r6 t7 18 19 20 tl 22 23 24 25 A Prelinrinary Investigation of Some l'lorphologica'l Characters in Adul t Females of the Sinrul iunr dantnosum complex from Togo A.R. Soponis and B.V. Peterson Oota.O.! til Jl rt tO': l'0.'!'3lOc altl I2 3 4 5 .6 7 8 9 10 1l L2 l3 L4 15 l6 t7 l8 l9 20 2L 22 23 24 25 I NTROT)UCT I ON Tlre problent of variation wi thin the species Sirnul.iurn dantnosum Theobal.d has been vlell docurrrented in the I i terature (Crisp .l956, Lewis 1958, Mc Crae 1966, Disney 1970, Gouteux )975, Ganns and Vajinre 1975). The identification o sibl ing spec'ies and the status of the taxonontic invesilgations of this conrplex have been recently revier.red (Vajime and Dunbar .l975). To date-, eight sibling species occur in l.lest Africa, of rvhjeh.six new species have been described frorn larv.ae collected in tlest Africa (Vaiinte and Dunbar .l975). There have been fcw investigations on the varjation of morphological characteps in the adul ts of the S'imul iunr datnnosum complex (Lervis .l958, Lewis and Duke .|966, HSusernrann 1969, Lewis and Raybould .l974). In fact, studies of the variation of rnorphologica'l characters in the adults of Simuliidae are rare (Davies .l949, Davies .l957, Rubtzov .l974). A honrogeneous collection of adult fenrales of the Sinrulium dantnosum comp)ex from Togo provided the opportunity to study the varjation of some morphol ogi cal characters. Because the di f ferences i n geni tal i c characters can be difficult to detect initially, characters excluding those of the genitalia r.rere chosen for study. The data appear to indicate that there are at 'least tr,ro groups. vri thin the col lection of adul t fenrales fronr Togo. Taking the ecological and cytogenetical evidence into account, jt is possible that tvlt sibling species of the S. danrnosunr complex are represented by the tlo grouPs' and that these species can be separated from each other on the basis of a combination of six nrorpho'logical characters in the adult fenrales. Holever, these results are tentative, and further studies are needed before the identitr of the different species can be established with certainty. oot..6.:l l:r!l r3to', l'o.' r'! loc a?i ! I 2 3 4 5 6 7 8 9 10 u t2 l3 L4 15 16 L7 l8 19 20 2l 22 23 24 25 -2- SCOP[ (}t STUDY This 'investigation is a prel inrirrary search for morphological characters that nray be useful in the separation of the sibling species of the Simulium damnosunr cornplex. A total of 55 charactcrs based on the morphology of the adult female, exclucling tire genitalia, t,rere examinccl ancl measured or counted. These characters r/ere chosen frclm scattered descriptions of Sinruliidae and frorr studies of variation in Chi ronornidag,,-since no conrparable studics have been dolr in the Simuli idae. The characters rt,ere arralysed for grouping by plotting the frequency distrjbutions of selected characters. OOa. l,.t t:rJ,,trO'i l'C.')'rlOl ) alll I2 3 4 5 6 7 8 9 10 l1 L2 13 l4 l5 l6 t7 18 l9 20 IL 22 23 24 25 -3- I.'ATERIALS AIID I'ItIIIODS Ninety-seven adu'lt fernales of 5:Ltlt]_:_q1i lgryqsu11 y/ere selected from the followjlg localitjes in the Guinea savanna of Togo, C. Vaiinte collector: (1) R'iver l4ono at Tetenou near Tohoun, 25.'1.l.75, adults caught biting and adul ts reared f rorrt puPae. (2) Dayes Elavanyo village near Palj.rrle, 26.11.75, adults caugltt biting. (3) R'iver Anroutchou at Pa'linre/Bidou Road , ?g.l'l .75, adults reared frorn pupae. (4) River l,Jar.ra at Ghana/Togo border near Badclu, 29..l.l.75', adults caught bi ti ng. (5) River l.lono at Kpessi north of Anie, 30..l.l.75, adults caught biting. (6) River I'lono at Tchartrba near Lama-Kara, .|..l2.75, adults reared from pupae. (7) River I'lo at Sokode/Bassari Road, 2.12.75, adults reared front'pupae. (B) River l'1o at Bagan near Bassari, 4.12.75, adults reared from pupae. (9) River Kpelou by Sara-Kar.ra near Lama-Kara, 5.12.75, adults reared from pupa' Each specinren vras nunrt-rcred (to avoid bias) and exarnined in alcohol . Iach specimen r.las tlren assigned to one of tlre classes A, B, or C after Lervis and Duke (.l966): A, basal vring tufts whol'ly or partly dark; B, basal r'rin9 tufts pa1e, anterior coxae dark; C, basal vrirrg tufts pa1e, anterior coxae pale. Slides of each specimen r.rere prepared as follovrs: rvings and'legs of one side v/ere removed frorn the rest of the body and placed into solutions of distilled water, gl aci al acet'ic aci d, propanol , cedan'rood oi I , and mounted di rectly onto a sljde from cedarlood oil-balsam into balsarn. The rest of the body r'las placed into either hot .|01 KOH or col d 2.5-53 KOll until the specimen r',as cleared, then (rr)ra o.-r l.'rJl ,5JO 2l rt.'! 3lof Alil I2 3 4 5 6 7 8 9 10 l1 L2 r3 t4 l5 16 t7 l8 l9 20 2L 22 23 24 25 rjnsed in disti l led water. The l,t'ad r.ras dissected from tlre thorax, and measurentents urere takcn in glyceri'ne on a sl ide. The palpus, antennae, and nrouthparts vere d'issccted and placed in glycerine under a coversl ip., Tlte thorax and abdornen r./erc stored in glycerine r'r'ithin a rrricrovjal that r'tas attachr to the sl ide. l.leasurcrrrcnts ylere taken vri th the aid of an ocular micronteter. Al I nreasurenrcnts t'rere takerr by tlre senior author as fol lorvs: Head. I.lidth (Fig. l): nraxinrum distance, usual'ly at l.evel of anterior margin of c)ypeus. Interocular Distance (Fig. l): frons at narrolest point tlidth of Eye (Fig. I ): (1./idth of Head - Interocular Distance ) x I Frontal Ang'le (f ig. 2): angle formed by crossed 'lines touching anterior and posterior nrargins of nredial jndentations of eye (lr/ygodzinsky and Coscari n .l973, fi g. 2, after Knoz I 965 ) . Antenna (F jg. 3): t.otal 1en9th of segments I and. 2, plus flagellor,reres Palpus (fig. 4): total length of each segnrent Thorax Length (Fjg. 6): anterior edge of scutunr to posterior ed-oe of postnoturn along straight axis in lateral vievr. l,li ngs . Length (Fig. 5): distance from center of lrumeral crossvcin to tip (Lervis and Duke .l966). I,,idth (l'ig. 5): nraximum distance, usual iy at basal 1/3. Legs. Length (Fig. 7): Iength of each segment along nredian axis. O0 la.O., l.'r!l ,110 el O"r'rltr' A?il I,! J 4 5 (r 7 t{ 9 t0 ll t2 13 l4 l5 l6 t7 l8 l9 20 2l 2'2 2J 1'l 25 -5- Ierr,ltlr of Wlrrte liurtd ort llirid S.rsitarsus: lengtlr alonq nredian axis. l'IidLh of lttrc ll.rs itarsUs (f ig. 7): iltdximunt distance, usual'ly distal 3/4. Courrts wcrc tu[:ert at 400 and 1000 rrragrrifications, usua]ly on the riglt of ;tcrit'cd structul't's. Most courtts are obvious; hoylever, a few deserve expla.atiolt. Slrirrcs ancJ setae on ve'in R of tlre wing, and setae on vein Rs of the win$ r,rere courtted frottt ct'c.rssvejn t'-tn to the poiqt-where the vein hits the Costa. uing tuf I setae were nlorc accurotely recorded wherr the sockets, not the setae, rvere countctl. 0f tltc nraxillary dentition, the lateral teeth include the apical tooLh, wlti le tlre rerna'ining teeth were recorded as medial . The frontal angle I'Jas,ncasurcd on a drawing rvith a protractor to the nearest degree. The dravrilrg was produced ft'otn a head ntoun.ted in g'ly,cerine, projected rvith the a.id of a lrlicroprojector'. Color classes A, B, and C were rechecked on the slidc- trroun ted s pec i ntelts . A'E' I.' i 4 5 6 7 I I 9 . l0 ll t2 l4 l5 16 t7 l8 t9 2tJ 2t 22 23 24 25 -6- RE5ULl S llte salltprle size, rdtlgt), tnedtl, and standard cleviation are listed for each of the 55 cltaracters in Apperrdix l. tlistograrns were plotted for only 33 of the 55 characters since ntatry charactet's showed'little or no variation (see Appen4ix r). Thc hist<lgrattts shotved three main patterns of frequency distributions: onc peak, two peaks and nrore than two peaks. Distributions of the one peak pattern were recorded for the follovling rrine characters: Iength of palpal segrrrents I and 3; width of wing; interocular distance; numl,er of teeth on the nraxi l1a (medianrlateral ); width of fore trasitarsus; length of hind tarsa'l segrnents 2-5; and'length of fope tarsal se!;rnen tS 2- 5 . Distributions of ttre two peak pattern were recorded for the folloning six characters (Fi9s. 8-13): length of ntid basitarsus, length of fore basitar-sus, lcrlgth of fore femur,.length of wing, length of palpal segnent 4, and number of spines on wing vein R. Variations of this two peak pattern were recorded for the lengrth of palpa'l segrnent 2 and'length of mid tarsal segments 2-5 (Fi9s. .l4, l5). Distributions of the more than two peak pattern were recorded for the following l6 characters: length of white band on hind basitarsus; total number of teeth on nraxilla; length of fore tibia; length of hind.basitarsus; length of hind fernur; number of setae on wing veins R and basal R; length of mid tibia; lerrgth of antenna; length of hind tibia; width of head; length of mid fenrur; frontal ang'le; 'length of palpal segnrent 5; nunrber of setae on wing vein Rs; and nunrtrer of wing tuf t setae. ,1 rr,.,l.\rf r.,leo ) -7- DISCU55ION Distributir.ltrs of tltc onr'[,ea[. pattern viere not of lruch intc.r.est since ortly otte grou;tiItg was presetrt. Tlre djstriLrutjons of color class A ref'lect tlte' onc peak pattet'rt in al I but two characters (length of palpal segment I , lerrgth of forc tarsa'l segtrrcnts 2-5). Color class B ref lects tlre one peak patterrt in four cltaracters (nunrber of median teeth on the maxilla, width of for'e basitarsus, lengtlr of palpal segrpelt l, and length of hind tarsal segments 2-5). Irr those charactcrs where aitf,". class doesn't reflect the overall paLtern,'it is most probably due to the snrall sanrple size. Lewis (.l958) measured the vring width.of 723 adult females of Sipulium _dgUlqtqll frorn northern Nigeria. He concluded that the frequency distribution (fig. I) "approximates to a normal curve". Lewis measured the rvidth along the 'line passing through the crossvein r-m of the wing, which was different froln this study. In this study the distribution of wing width was,a one peak pa tte rn 0istributions of the 2 peak pattern were of the nrost interest because tlre sante pattern was shared by six characters, the pattern was not complex, and color classes A and B reflected the overall pattern for nrost characters (Figs. B-13). The distribution of color class B would most probably reflect the overall pattern nrore closely if the sanrple size were larger. In order to detennine whether or not the same lndividuals occurred within the corresponding lrroups (peaks) of the six different characters, Table I was cbrrrpiled. After two characters (nreasurements) were conrpared, 87 out of the original 97 characters agrecd in grouping. After five characters (measurerrrents) were compared, 46 out of 97 specimens agreed in grouping. I 2 3 4 5 6 ? 8 I 10 l1 t2 13 l4 l5 l(r L7 l8 l9 20 2L 22 23 24 25 U, ra r., 1,, tt l' r,, -'l lrJ, 'l,,d ArB, -8- I ,| 3 .l 5 6 7 6 9 10 It t2 r3 I{ l5 l6 t? l6 r9 20 2r 22 2J 24 25 ) llur.rt-,vcr, rJlron a diffc,rerrt cltaracter (courtt) was cornpared to tlre five nrcosurerncnt orrly 30 of 97 specitrrcns agreed in tht grouping. In r.rrder to clctcrrrrinc whethcr or rrot color class affected grouping, the t1ata in Table I was arranged according to color class. Since alI color classes iv(.:te co,lsisterrtly represcnted in tlre ttvo groups, there appears to be no irrdicati, tlrat color class affects grouping. To detcnnine whether or not rearing-or lrabitat affected grouping, Table II was conrplied, based on ?2 reared and 65 nonreared specirnens, of which 54 were collectcd at or near the Mono River. There is orrly one reared specirnen from the l'lono River in the left group of the histogram, but reared specirnens from I'lorro consistently appear in the right group. Both reared arrd nonreared, front nrixed lrab'itats and from Mono, were consistently represented after the six characters were conrpared. Therefore, there appelrs to be no indication thdt rearing or habitat affects grouping. To determine whether or not collection site affected grouping, Table III vras conrpiled. A)though only tvto habitats are well represelrted (11 , 4) , af ter six clraracters were compared the different collection sites wcre represented jn ttre two different groups. Tlrerefore, there appears to be no indication tlrat ct-rl lection si te affects grouping. Based on the cytogenetic work of Vajime and Dunbar (1975), five species of the S.d. conrp)ex occur in the Guinea savanna of West Afrjca. 0f these, -1. :-qrbrc-!rq, S. sirbanurn, S. danrrrosuqq, and !. gu.Jg"nlg have been collected in areas near Togo. Larvae identifjed as S. soubr"ense and.S. sirbanuni by Vojirrte were collected by him on the same,day and at the santc site as that given for'locality I (p.3). Specimens from locality l, both reared and nonreared, A!Ui -9- I <rr.., l.r-.1)r.esc,rLr:d i, both groulr.s of the 2 peak <Jistr.ibution (Tables I, IV)) d f ter s i x char''rcters wet'e corrrpared. Both of these species dre sympatri c3 r.rlrer.cvur. Uley occur (Vajirrre and Dunbar lgll, p. 1Zl). It nray be possitrle that .r thc two groups represent. these two species. l.lowever, further studies areItut'ijed to cUrrf itrlt tlte prel inrinary resul ts indicated here.6 Becauso adults of s-' 9ci!!9:tr! are known to fly <Iistances frol.5 to over7 20 knr' f ronr thei r breecling si tes (Dut<e 1gl4),. the possibi r i ty that nonrearedc flies foutttl at locality ) *o.e not bred there cannot be overrooked. However,9 redrc'd spc'citttens from locality I are represented in both groups of the twolo pt,ak distribution (Table III). ll Distribtrtiorts of the more than two peak pattern show as many as rz peaks12 (rrunrber of wing tuft setae). Distributions of the three peak patte* were13 r'ecorded for six characters: total number of teeth on rnaxir.ra, rength of hindl4 fetrlur' length of fore tibia, length of hind basitarsus, length of white band15 on hind basitarsus' and number of setae on wing vein R. Distributions of the16 four peak pattern were recorded for three characters: Iength of nrid tibia,17 'length of antenna, and nuntber of setae on wing vein basar R. gistribution of18 tlre five peak pattern was recorded for the length of the hind tibia.19 Distribution patterns with more than five peaks would indicate that20 thc following characters are too variabre to be useful in species separation:2L hcad width' frontal angle, length of pa'lpal segnrent 5, rnid femur length ,22 ,urnber of setae on wing vein Rs and nunrber of vring tuft setae.23 Although this study is based on a snral I samp.le size, the data appear to24 i,dfcate that two groups of sinrulium lgirug;gg are represented in the adult25 feinales fr,m Togo' These groups can be separated from each other by a con.ination AlcS -I ,, .] .l 5 6 7 8 9 t0 l1 t2 l3 l4 l5 l6 t7 l8 l9 20 2r 22 23 24 25 clf Llre lr.r'l lov;irr(l f ort: tr.t:, i I d i':, u5 , 4 , atrtl lrurrdrsr o f the two 9r'otrlri rrgs s i te . [,Jlre tlrc r or !. potrtrrcnse_ dnd 0rrly when 'lar'ger available carr the Sitntrl i trrrr d.arrrn_<.rsurn -.l0- chaructcrs ( rable IV): length of mi<j bas i tarsus , 'lengilr of 'lcngtlr of fore fcrrrur, length of wing, length of palpa'r segnrcnt spincs on wing voin R. There is no evidence to indicate that are affected by color c]ass, rearing, lrabitat, or collection not thr: two groups jn tlris study correspond to the species l. :jflggU can be establislred only after further study. sanrples of rearei, nraterial from identi f ied cytotype larvae are taxononrist confidently define the different species of the corrrplex in all stages of the I ife cycle. a?al I., .3 .l 5 b I 9 IU ll L2 l3 1.1 ,5 l6 t7 l8 l9 20 2L 22 2:l 24 25 coNtL US I0,lS A tot,r I uf 55 rtlut'pho'logi ca1 cltaracters in 97 adul t fenrales of the Simul-iurn {t,rrirlt)1,.tllrr I ll . corllp lex I rottt Togo were s tudierJ. Frequency di s tribut icl's of 33 cllrrl'ctCtet's wct'e p lotted, re5ul tirrg in three. nrain .patterns: one peak, two peaks I arr(l ntoro Llran tvlo pcaks. Jhc slrecies -S. l-il,ltlrylrl Vaiirtre and.Ounbar and S. soubrense_ Vajime and Duntrar nllry corrC:,1rclrrd to ilre two groups iir the two peak pa.ttern. If so, the aclult f't-'ntales of tlrcse ttvo spec'ies can be separated froln each other by a conrbination ol the follorvirtg charactcrs: length of mid basitarsus, length of fore basitarsus, lcrrgtli of fot'e fetttur, length of vring, length of paipal segrnent 4, and number of spitres <.rtl ttirrg vein R. There was no evidence to indicate that color c'lass, re,.rrirrg, lrabitatr or col'lection site affected the grouping. Several cltaracters appear to be too variable to be taxononrically useful irr specics sr:pdrdtion:. ltead width, frontal angle, length of palpal segrnent 5, 'lerrgth of rrtid fernur )ength, nulnber of setae on wing vein Rs, and number of wing tuft setae. A total of l0 characters v/ere founcl to havc three, four, or five peak distributions. Furthcr studies are needed to determine whether or not these peaks rept'esent species. Reared nraterial from identified cytotype laryae is t:srential to the separation of the species of the !innq'liunr damnosum complex in al'l life stages. A2llJ I') 3 4 5 6 7 8 9 l0 ll L2 l3 l4 l5 l6 L7 l8 l9 20 2t a1 2J 24 25 -12- LI IIRAIUI(t. CITLI) ('r'ts1,, G. 191,t,. 5ttrurl i_rrrtr and onchocerciasis ilr the ilortlrer-ri Territories of [lre co]tl coa:,t. lhc'Brit. Enrp. soc. for Ure Blind, ll.K. Lewis and Ct-r. l-trt., L()ndon D.tvics, D.l'1. .l949. Vari.rtiorr in taxonoruic clraracters of sorne Simuliiclae (UipLcru). Qtt. Itit_. 8t (t ): t8-Zt. D.tvics, L. .l957. A study of the black:fly, Sinrulium ornatr{m llg. (Diptera), rvith Particu'lar reference to its activity on grazing catfle. Bull. e1t. Ief. -4.!: 407.424. Uisrrey, R.ll.L. 1970. A note on variation within Sinrul ium Aar,lglm in the forcst zortc of l.lest Crtmsr"oofl. Ann. trop. l'led. P;rasit. gL (1): 'lZg-130. Dttlie, 8.0.t. 1974. llre ecology of onchocerciasis in relationship to the ecology of r,,an. -Ir1 Proceedings of the First JnternationaJ Congress of Ecology. Stt'ucture, Functioning, and Manageluent of Ecosysterns. The llague, Ttre Netherlands, Sept. 8-14, 1914, pp.323-329. Girt'ttts, R. arrd Vaj'ime, C.G. 1975. 0n the eco'logy and <listribution of the species of the !jgg]_ig1 damnosqm complex in different bioclinratic zones of Liberia and Guine.a. Tropelmed. parasit. 26 (3): 375-380. GoutcuX, J. -P. 1975. Larves de simulies (Diptera, Simuliidae) du r,.ivu reattacltab I es au corrrpl exe "damnosunr". Descri pti on de quatre types ' distinr:ts rrror"phologiquement. Cah. 0.R.s.T.0.lrt., ser. Ent. nrr6d. et !erer.i_L_q] . _lJ (a): ?37-243 llliuscrrnanfl, W. 1969. 0n the biology of Sinruljurrr danrnosunr Tlreobald, 1903, the nraitt vector of onchocerciasis in the Mahcnge Mountains, Ulanga, 'l'anzania. Acta trgp.-26 ('l): Z9-Og. tt '' ttt ., | (r.. r.'llrld A.'UT I? J .1 l{ortlrtrrrr li igi: r ia. &n, !-1gg. !t&_lg_tq:j!. 52: 216-231 . 5 [.urvis, t).J. arrd l)uku, 8.0.1.. 1966. 0rrchocerca-!_Ug]jgti conrplexes. I l. V<rri.rl" ion in l,lest African fernale Simu'lium d_amnosurn. |Ur. !I9l),_J.1gd.. Ptrasi_t. 60-: 3Jt:346: 8 letvis, D.J. arrd llaybould, J.N. 1974. Sonre Tanzanian Sinruliidae (Diptera). ftt-tgp'llgi:_t1' ggQ. l'1a9. Il0: 4l-50. l0 I'lcCt'.rc, ir..l,J.R. .l966. The I!tg]_l-um damnosqm species complex. E. A-fr. -Vir!g- I I !les- -lns-t..-.I!gti-L- -l g : 38- 39 . t2 liubtzot,, i.A. lg74 , The variabi I i ty of taxononric characters in black f I ics 13 (Diptera, Sirrruliidae); research tasks ancl methods. Ent. Re.v., tJash. r4 !_l_ (l ) : )s-24 l5 V;rjinc, C.G. and Dunbar, R.tJ. 1975. Chromosonral identification of eight species 6 7 l6 t7 l8 20 2L 22 23 24 25 l.tru/,.,. l{,u5. 'lo ttlt'trLif rra'Liorr of Czec}rr.r:lovakiorr [-rlackflie,, (Uiptcl..r, 5irir,rl i rd,r,.-'), [..q]_r9 j:]1.. _s.j.- !{_q_!.-_tl1r-iy. Purtiy-idnae.Brurl-qrsl-: q.: r-54. lcr;i:, [)..1. l95tt. Ubscrvdttons on Sirrruliurrr danrnqsunr Theobalcl at Lol,.oja in of tlre suhgenus Edvrardse,l lum near and inc'luding Sinrul ium (EOwqf{:S-l-ly-fff) .crgg)-.f:llnt Tlreobald (Diptcra: Sinruliidae). Ifggenlg!-. Parasit. -?!. (l): lil-138. Anrcrican black flies of the tribe [,rosimuliini (Sirnuliinae, Simu]iidae). 0_ul I . Aru.__Uus-_trgt._l!_i_st. l5l (2) : 129-200. 19 tJyqodz'insky, P. and Coscar6n, S. 1973. A review of the Mesoarrrerican and South 'l /r',116 ) Arit . 14 - llr lrrl,ullr,r.,I .1,.' t,',r',1r (u,r[,J.tlrg (, rtrJtd!tct.:, Ar(u!g.l dC((,tdlng ttt trrlur ( l,rt:r,) A. ll. llrrl C. Iul,lr I Cl,Jr u, tr r (l) tlrr,lttr of li1,l Ualt.,t,rr.,u. (2) trrrgth ul lort 8a.i I talrrr: (3) lrrrgth of fore ferrlrr ( 4 ) teng th of l', i,rg (5) Lerrgth.of Pallrat gegnrel1g 4 (6) t{unrtrr.r of Sptnes on Vcln R of lli ng Con4r.rr rson of Atr<.rve Srx Ch.rrdct&t's. (r) x (2) (r) x (2) x (3) (r)r(?)x(3)r(4) (r)r(z)r(3)x(4)x(5) Clasr A B c .A B c A I c A I c A b c A B c A I c A I c A I c A I c A B c Left Pcol l4 4 4 .19 I 5 l9 l{ I 30 l0 5 l2 5 4 ?t l0 4 l4 4 4 ll I 3 l0 4 3 3 2 2 I I I Rlght Pt'ak 50 23 ? 45 l9 I .t5 t3 2 t{ 17 I 50 22 2 3l n 2 45 l9 I t0 t2 I 30 t? I 2l n I t, 9 t Total llunDer of 5pec larn5 9l 9t 9l 9t 95 9? 8t 7l 60 a5 (l) r (2) x (l) r (4) x (s) x (0) 30 - 15 - l.tl,le I I . t)is t_r.rbuL iorr ul spe cirrrerrs, corrparirrg 6 t'c(u'irrit arrtl lr"rl-l iL,rt. L is le.ft peak, R cha rac ters , a rrangcd accortl i rrg i s ri ght peak of tr i s togrdrn. t <.r Cltot'actc,r' i (f) x (2) (1)x(2)x(3) (r)x(2)x(3)x(4) (l) x (2) x (3) x (4) x (5) (r)x(?.)x(3)x(4) Mi xe.d llab i ta t Reared L5 Rt6 L3 R ls L3 R l2 L2 R t2 .. Jllonreared 17 49 l5 38 14. 3l 5 27 2 17 l,lono Ri ver Reared 1 l3 '0 12 0 l0 0 l0 0 I Norr rea red I 28 7 25 7 l8 4 t7 2 l2 x(s) x(6) L I R t0 * s atrrc as ilt Table I Iol,lt.' ll I [ti:,trll,l/t.r(.r,r to t,-tt llet t iuri Ir i s Lo,lr.arrr. tt[ 5[).,L ilttcn5 t :iLt, (seu p. - 16 - cornl)Jri rrg 6 ctraracters, 3). L is left peak; R a rralrged accord i ng is riglrt peak 0f Col lt't.tit,rt Sitr: 1. r R 2. t. R 3. r- R 4. t. R 5. L R 6. L R 7. L R 8. L R 9. L R * san,e rrs ilr '[.tb]e l. 5 35 2 I 0 I 6 l2 2 2 0 0 I 0 0 I 2 I (r )x(2)x(3)x(4) -I.-:-- 5 26 2 I 0 I 5 12 ? 2 0 0 I 0 0 0 ? I (l )x(2)x(3) x(4)x(5) 3 26 I I 0 1 0I I 'l 0 0 I 0 0 0 I I ( I )x( 2)x(3)x(4) .____{J!X6) I l9 0 I 0 I 0 4 I I 0 0 0 0 0 q I I (DIJA(U.iolr(/l 7 3u 2 5 0 I 7 l6 2 ? 0 1 I 0 0 I 3 I _U_ l.rtrl,' lV. li.utrlr',, ol t.lr,rrtrr tcl'5 for Llrr, St-.pr.rr.<rtir4r of tlre tnt-r g).o1ps Of fcrlrolt:s tlf tlrt' S lllrlrl itrlll rl(lirrlro''ttttr cotttltlt:x 1r'orrr lclgo. fleasurenrerr t,s are givc^ .i n rrrcr.ons. llr":"rttt1rl'-'size Uf tlru lirst f-ive r:har'<rrters is 46; f,r the last ctrardcter, 30' Clto r'.rt-l tl' Lentytrr ol l'1td [JtrsiLar..sus lc'rrgth ol f ore 0.rsitar.sus Lc.rrgth r.rf I-or.c ['ernur Ierrgih t,f' li'ing Lr.ngLlr uf I)alpal ScArrrr,rrt 4 lluttrlrt-'t' o f Sp irres orr [,J i rrg Vein R Le&*P.si!- 268-299 371-412 525-s87 1918-20t8 lt2-138 6-7 ntsht pgd 309- 360 433-494 618-690 2163-2326 143-17 4 l0- 16 - tB - Fronta I Fig. 3 Fig. 5 I'lax imurn uri dth fore basi tarsus - Maximum width of wing teooti llecli arr Lenlr th of Fi g. 19 - f tg. ti (e, l(, a '1.; r 4d, CLur l'? (a- o lt,t t-or8 J)[:2b 4 Length in units; microns - x 10.3 Fig, 8. Frequency distribution of the length of the nrid basitarsus in adult females of Simul iutn danrrrr)tum, . ari'anged according to color claises'-A]U); ti(O), drrd C(H). # ?.n- ht Lerrg th i n un i ts ; nri crons - x .l0. 3 Ir'equency distribution of the )ength of the fore trasi tr'lrsusin aclult f enrales of Sirnul iunr danrnosunl, arranged according to color c'tasses A(A) ,--BTtr); Ena-C(gI. a <: 4t tr c, C! (+- -c)() ,l o!) E: :, ,< 20 Iu l6 t4 12 l0 B 6 4 ? M fig. 9. -20- TC c vl ,l'(U ao._EO CJ \-;rl- o +J €g ,o< Eut 'F o., ahL i./, :, G'Erce (I,u .q- OLF (lJoL-x ooq-(, I (uo -c +, +J (rlq-c O'r .IJ -cL{-, or,)L)E'(J ct (o E(u0J -c c,+Jc ,oq-L o! (o ac ,oo(oL (J E= Otff, tt ''J c@= Lr) g -c+)\ (r) oc a)J co .F E:I+, flfl,l .o01 .e ClLEI +, rul(/l1ol €et5l5'-l(-) -lc :rlq., El =.Fl . Cr Vtl'- flu-Elr- O L) c) cn t-L ro <r s^t suautJods jo JaqunN C)) .- cD (o \, c\l o F7-@ (rt L -21 .,, lt, 1(:l sr 14 I1- ilr;'tvt l0 -l- -1, I ' 8t C, I{r 6 -l- :2 4 -{- ,L rg. ll 55545352st dltwt&48 49 50 Fi g. {1 il J I -t- I '1- :t l + t t_ tn !lt: (J CJ c:- r u1 t t+- o !- o) -og :I 1? 2Lt l8 l6 l4 t? l0 B o 4 2 + 5756 Leng th i n un'i ts ; mi crons - x 40 . Bl Frcrluency distribution of the length of the uing in adul t fenrales of S'inrul iurn damno-slm, arranged accordirrgto co'l or cl asses A-(A)l-BlOI7anO-t(a) . ,L1 _n5a Lengtlr in units; nricrons - x 5.ll Frerlucrrcy distributiorr of the length of parpal segnrerrtin fernales of Sinrul iurrr danrnosum, arranged accclrding to color cl asses n(A)-,-ttln)l ano-T(Al . ['i g. 12 . C c, t OJ LT' o l- o, .ar el: l3 '? t, '?:t ,?,| i)'.) IrJ l6 l4 12 10 I () 4 ? Nulrber of spines lig. 13. Frequency distrjbution of the numLrer of spines on r.ring . vein R in adult femalcs of Sjmu]ium damnosun:, .irrcngcd according to color ctasses A-[EI;B(E);ai'?-C(lf) . ?4 l-2?I I20 -l- ? lB -lcrlt. 16 I'Jl H 14 + ,* l? +c)l ,'l0 l-O, Iit 8 -l- -ll .lo +-' I4 _l- 2) t_ 27 30 Length in uniLs; nricrons - x .l0.3 I-ig- 14. Frequency distribution of ilre length of the nridtarsal segrrrents z-5 in adul t f ernaies of simul iurndar,nosunr, arranged according to coror ctassei-'ria),F(o)-;o c(E). - tt c(u F: o €r C-L vl 9-o L.(uA r.' l) 37 30 ?.8 26 24 22 20 }B 16 t4 1? 10 8 6 4 2 -23- Length in units; nricrons l817l6 l9 - x 5'.ll ,>i Fig. 15. F retprency segnrent 2 arranoed and C (a) . distribution of the length of palpal in adul t females of Sinrul ium damrrosunr, accordi ns to color ct ailE-A-(Z)]-B1OI; ,l 14 15