Studies on Neotropical Fauna and Environment

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Studies on Neotropical Fauna and Environment Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713817190 New record, geographic variation and redescription of Apedilum elachistus Townes (Diptera: Chironomidae: Chironominae) Mariano Donato ab; Analía C. Paggi b; John H. Epler a Laboratorio de Sistem tica y Biolog a Evolutiva (LASBE), La Plata, Argentina b Instituto de Limnolog a “Dr. Ra l A. Ringuelet” (ILPLA), La Plata, Argentina

Online Publication Date: 01 August 2008 To cite this Article: Donato, Mariano, Paggi, Analía C. and Epler, John H. (2008) 'New record, geographic variation and redescription of Apedilum elachistus Townes (Diptera: Chironomidae: Chironominae)', Studies on Neotropical Fauna and Environment, 43:2, 125 — 134 To link to this article: DOI: 10.1080/01650520701437707 URL: http://dx.doi.org/10.1080/01650520701437707

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Downloaded By: [Donato, Mariano] At: 16:33 9 July 2008 ‘r Rau ‘‘Dr. n umrzto fvrainadcovariation and variation of summarization and syst biological in of analysis measurements in sexes. both of mainly adults specimens, and southern between US northern differences from found material it We with samples. species, compared and this measured of was distribution known southernmost of Grosso southwards. its extending range specimens thus Mato (Argentina), province New in Aires Buenos (1995). species Nolte this elachistus of by (Uruguay) finding (Brazil) Indios los the de examined Piedra and specimens from (1988) male Epler two by were America South 1996). Reiss Guatemala, & (Spies in Brazil occurring widespread and is region, Nicaragua with and latter Neotropical Nearctic The the the Neotropics, in in limit. USA southern the has the in known it occurs its and in Nearctic as the Guatemala distribution in USA the restricted and occurs Canada former the in The in regions. distributed Neotropical both and Nearctic 1988), Epler, 1945; (Townes, subcinctum Rcie 0Jn 06 cetd7My2007) May 7 accepted 2006; June 30 (Received http://www.informaworld.com 10.1080/01650520701437707 DOI: online # 1744-5140 print/ISSN 0165-0521 ISSN [email protected] Email: author. *Corresponding genus The Introduction a Donato Mariano of redescription and variation geographic record, New ARTICLE ORIGINAL 125–134 2008, August 2, No. Environment 43, and Vol. Fauna Neotropical on Studies hrnmde Chironominae) Chironomidae: aoaoi eSistema de Laboratorio 08Tyo Francis & Taylor 2008 h rmr olo egahcvariation geographic of goal primary The the represents collected material new the As for data distributional only The boe,V,adlnt fploeeVfrfmls Consequently, females. for V palpomere of length and VR, abdomen, fdroeta ea,nme fpelrste iglnt,V,lnt ftoa,adlnt fadmnfor III, abdomen palpomere of of a length length setae, reflect and temporal thorax, of differences of number these thorax, length of VR, if BV length number length, setae, setae, determine length, IX wing acrostichal Component gonocoxite to wing setae, of of Principal prealar number order number a the of and in in males, number specimens variation performed setae, American North–South dorsocentral were show North of results Tree and The South Spanning differentiation. geographic between Minimum found and dissimilarities Analysis to Due species. elachistus Apedilum Keywords: female. and male, pupa, lA igee’ IPA,L lt,Argentina; Plata, La (ILPLA), Ringuelet’’ A. ´l aebe on ytefrtato in author first the by found been have Apedilum onsand Townes Apedilum ab ,Analı *, tc Biologı y ´tica 2 onsicue h species the includes Townes egho ou,BV notum, of length , hrnmde hrnmne egahclvrain etoia region Neotropical variation; geographical Chironominae; Chironomidae; ; ons14 srcre rmAgnia ersnigtesuhrms eodo this of record southernmost the representing Argentina, from recorded is 1945 Townes aC Paggi C. á mtc stedescription the is ematics .elachistus A. aEouia(AB) ue eL lt,L lt,Argentina; Plata, La Plata, La de Museo (LASBE), Evolutiva á .elachistus A. b n onH Epler H. John and 1 ubro oscnrlste egho apmr V eghof length IV, palpomere of length setae, dorsocentral of number , Townes A. A. in c rwodil,Foia(L,USA (FL), Florida Crawfordville, c ifrnito.Teeoe elo o statistical (H hypothesis for null the geographic look reject to a evidence we reflect Therefore, differences to differentiation. order such in the if method statistical determine a to of application the Due is 1969). of specimens ern Sokal bet found & of dissimilarities kind (Gabriel this location co and these characters, relates one of analysis observable terms more in made or are populations area these among the number in a localities at in of organisms studies comparable for of re populations basis variation The area. geographic an over distributed characterist the in patterns otenseies nteohrhn,tealter- the hand, (H other hypothesis the native On specimens. southern differe morphometric are n erdi h aoaoyi niiulcontainers (18–20 individual temperature net in room a laboratory at with the collected in were reared and Larvae swarms. of netting New methods and Materials specimens. ern be differences geographic one nsie ihCnd balsam. Canada with slides on mounted pdlmelachistus Apedilum .elachistus A. pdlmelachistus Apedilum .elachistus A. aeilwscletdb hand by collected was material 1 otltsta hr r no are there that postulates ) prsn odfeecsin differences to mparisons ne td.Comparisons study. under ensuhr n north- and southern ween csbtennrhr and northern between nces we otenadsouth- and northern tween t nteeitneof existence the on sts srdsrbda larva, as redescribed is c fognssta are that organisms of ics u ) h pcmn were specimens The C). ons(Diptera: Townes h i fti study this of aim the , b nttt eLimnologı de Instituto 0 )thatthere á Downloaded By: [Donato, Mariano] At: 16:33 9 July 2008 V,rtoo egho ut egho ) number R M); of on length to setae Cu of Ratio of length Venarum of wing; prealar of ratio of width (VR, number wing; of setae; length scutellar setae; the of dorsocentral flagellomeres); of number basal number by setae; setae; more acrostichal divided the of number of flagellomere length elongated combined of apical length length of ratio setae; of (AR, Ratio clypeal Antennal of I–V; of palpomeres number number abdomen; setae; of length temporal thorax; of length are: 1969). local dimensions Ross other & if (Gower detect apart account) far into help are taken actually are but to plot on together a close superimposition look in PCA) which for points of pairs as useful (i.e. distortions is (such It ordinations path them. best the unite for looking by to other each to relate data the high- of influence the variables. reducing relatively variance the and with variables variance has of small influence matrix the variable inflating each Correlation of by influence The the in equalizing of units. expressed advantage of not same matrix are symmetric variables the square the a since in on Correlation based performed is PCA study their and The this highlight to differences. data, as and way high-dimensional a similarities such in in data the Minimum patterns expressing of and cation (PCA) Principal (MST). Tree Analysis Spanning groups: between Components differences investigate indepen- analyzed was multivariate stage the dently. examined’’). developmental perform ‘‘Material see to Each details more from used (for nine were analysis and Argentina USA) from USA) 14 the (seven the USA), from larvae nine 16 the and and from Argentina from four and (five and Argentina pupae females Argentina from adult from eight (10 USA), (four males the from adult five and 15 Uruguay of total A analysis Multivariate and FS) MLP). M (Argentina, (USA, Plata & La A de Florida Museo Florida Tallahassee, at Florida Arthropods JE), University, of specimens (USA, Collection The following collection State the mean. to Epler’s correspond the institutions: study by this for followed examined given are ranges measurements All as (1987). Epler and (1980) egho eaassdvddb egho ii;LR tibia; of of length ratio by (LR, divided Ratio metatarsus of Leg length chaetica; sensilla of number 126 h hrcesue ntePAfrbt sexes both for PCA the in used characters The how see to useful method statistical another is MST identifi- for useful technique statistical a is PCA to order in applied were methodologies Two Sæther follows nomenclature morphological The .Dnt tal. et Donato M. + R 1 ubro ea nR on setae of number ; 4 1–3 + 5 ; eest rn,mdadhn e,respectively); leg, hind and mid front, Beinverha to refers i n eaassdvddb obndlnt of length combined BV by 2–5; divided tarsomeres metatarsus and bia hrcesnme fsteo R on setae of number characters apmr I,lnt fadmn egho palpo- BV of length, length wing abdomen, V, on mere of of length length setae, III, temporal representation palpomere of acros- number of VR, number setae, two significant including: tichal total 1), (Table first 1 Several the had Component the respectively). of 21.05%, specimens, characters 63.64% and male (42.58 for variation of accounted PCA components the In Results analyses. both and perform to X; used was tergite on IX. gonocoxite setae on setae on of of setae number number of females: number VIII; cerci; of of sternite analysis length notum; for following of length taken the were Additionally, measurements respectively). leg, hind S,cmie egho femur SV of metatarsus; of length length Schenkel-Schiene-Verha combined and (SV, respectively); leg, ea nR on setae ewe .6ad28 m h eut rmMST from results abdomen The of mm. 2.81 length and and and 2.26 thorax mm 1.39 of 1.51 between length and between 0.94, 0.93 and length between 0.6 between wing VR mm, specimens setae, 2.00 northern prealar setae, dorsocentral hand, 1–2 9–13 and setae, other 1.42 acrostichal and 10–12 the between 0.69 have abdomen On between of thorax mm. 2.12 length of and length between mm 0.84 VR 1.35, setae, mm, 1.47 and prealar and 1.28 1.12 2–4 between acrostichal setae, length 2–4 wing dorsocentral possess 16–24 specimens setae, abdomen. wing of southern length setae, and Thus, thorax prealar of length of VR, length, number of setae, number setae, dorsocentral and acrostichal northern of component number and are southern this specimens between in values The different scores former. with best the in with mainly characters but components, second of specimens component. first the the along mostly of groups observed the consisting is of dispersion B the Internal group specimens. comprising northern and A specimens group second southern plotted: and first were the when components formed the two were that of with groups shows major 1 number Figure association 1). (Table and strong component second showed setae setae clypeal scutellar of number chaetica, h rga TSS eso . Rhf 1998) (Rohlf, 2.0 version NT-SYS, program The h eaainbtensuhr n northern and southern between separation The lnse(V obndlnt ffemur of length combined (BV, ¨ltnisse 4 + 5 .elachistus A. en R LR AR, vein, 1–3 eest rn,mdadhind and mid front, to refers 1 1–3 n egho hrx The thorax. of length and cusi ohfrtand first both in occurs eest rn,mdand mid front, to refers 1 ubro sensilla of number , + R + 1 ii iie by divided tibia en,nme of number veins, ¨ltnis + ti- Downloaded By: [Donato, Mariano] At: 16:33 9 July 2008 ubro ea nR on setae of Number it fwn ,190,618 0,9842 2,4179 2,9626 VR wing of Width wing of Length ubro raa setae prealar of Number ubro ctla ea 0,4493 setae scutellar of Number ubro oscnrlsetae dorsocentral of Number ubro crsia setae achrostical of Number AR egho apmr 2,9708 V palpomere of Length egho apmr V2,6014 IV palpomere of Length LR egho apmr I 3,1164 III palpomere of Length SV SV BV BV LR chaetica sensilla of Number BV R on setae of Number SV LR ubro lpa ea ,5 1,5434 2,851 1,457 I palpomere of Length setae clypeal of Number egho apmr I2,5773 II palpomere of Length specimens. of PCA 1. Figure 0,8976 0,8904 2 Component 2,6821 3,0003 1 Component setae temporal of Number abdomen of Length thorax of Length analysis. Character PCA male’s from obtained loadings PCA 1. Table 3 2 1 3 2 1 3 2 1 pdlmelachistus Apedilum + 4 + R 5 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2,7188 2,9462 ,640,9007 3,3634 ,552,1913 1,5555 ,731,6422 1,8783 ,530,1914 0,5533 0,9371 1,6352 2,2834 2,9171 1,1484 1,0957 0,9883 0,1322 0,0602 0,8366 1,5661 0,3203 3,358 ,5 0,4004 3,354 ae.Mnmmsann rei ueipsd ( superimposed. is tree spanning Minimum males. 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 0,4339 0,8637 1,3343 0,3193 0,1589 0,1995 0,5333 0,4609 1,1497 1,2825 1,0735 2,2103 0,8643 2,2167 0,775 1,794 0,833 tde nNorpclFuaadEnvironment and Fauna Neotropical on Studies ftmoa ea,lnt fploeeII wing III, palpomere of length BV length, setae, number thorax, temporal of length of setae, IX num- 2) gonocoxite (Table characters of ber from 1 components (59.19 representation Component variation significant two had total respectively). the first 15.53%, of and the 74.73% for that accounted showed analysis rdetadgorpi itiuino specimens of H distribution therefore and geographic character and between gradient relationship geographic direct a is the This as evident. interpreted and becomes specimens the above of a distribution mentioned results, high- with loadings these measurements on est the Based between PCA. relation of direct those with agree hrcesA,nme fsuelrste LR setae, The scutellar V. of palpomere of number length AR, and characters of VR length IV, palpomere abdomen, of length setae, dorsocentral otyaogtescn opnn.I h aeof case the In component. second the along the mostly of the consisted second contains B 2). (Figure and group specimens A northern first and group specimens the plotted: southern when were formed components were major Two 2). associa- groups (Table strong component second showed the width with tion wing and I palpomere ubro eslacatc nlgI,LR II, leg on chaetica sensilla of number o eaeseies h rnia component principal the specimens, female For nenldseso ftegop a observed was groups the of dispersion Internal 2 ou egh BV length, notum , 0 sacpe o h C fmales. of PCA the for accepted is m otenseies ( specimens; Northern ) 1 ,BV 3 ubrof number , 1 eghof length , N southern ) 127 3 , Downloaded By: [Donato, Mariano] At: 16:33 9 July 2008 LR ubro ea ntrieX441 0,5275 4,4814 3,6108 IX gonocoxite on setae X of tergite Number on setae of Number it fwn ,321,4067 0,6143 3,3352 4,3124 R on setae of Number VR wing of Width wing of Length setae prealar of Number setae scutellar of Number setae dorsocentral of Number setae achrostical of Number egho ec ,3 0,8693 0,3166 3,839 4,2628 VIII sternite on setae of Number cerci of Length notum of Length R2,2519 0,5123 0,449 0,5975 4,1003 1,1567 1,4937 4,158 4,3596 3,5237 3,331 AR V palpomere of Length IV palpomere of Length III palpomere of Length II palpomere of Length I palpomere of Length SV BV LR 0,1179 III chaetica sensilla of Number ubro ea nR on setae of Number ubro lpa setae clypeal of Number setae temporal of Number SV BV BV SV LR 2,4496 II chaetica sensilla of Number n 175 and esecp hrce BV character have except speci- scores southern mens highest and northern with between values characters different the all females, 0,0637 4,4461 4,1299 2 Component 1 Component abdomen of Length thorax of Length PCA female’s from Character obtained loadings PCA analysis. 2. Table 128 pcmn of specimens eghbten199 n 204.6 and 189.95 between length n egho apmr ewe 5.6and 151.56 between V palpomere of 199.55 1.32 and length 1.19 between and VR mm, 2.86 101 and palpo- 1.92 and between of 81 between length IV setae, dorsocentral mere 15–18 2.38, and .8ad19 m BV mm, 1.97 101.04 and and 1.68 80.83 III palpomere of between length setae, temporal 7–10 and mm, 1.03 1.16 between thorax of length setae, IX gonocoxite 4tmoa ea,lnt fploeeIIbten53 BV between III 58 palpomere 23– of and mm, 0.76 length and setae, 0.65 temporal between 24 thorax of length setae, elachistus A. 3 2 1 3 2 1 3 2 1 2 ewe .7ad45,ntmlnt ewe 162 between length notum 4.54, and 4.17 between m m .I otat otenfml pcmn of specimens female northern contrast, In m. .Dnt tal. et Donato M. ,wn eghbten12 n .2mm, 1.42 and 1.26 between length wing m, m ,BV m, r hrceie ytogncxt IX gonocoxite two by characterized are .elachistus A. 1 + 4 + R 5 ewe .5ad27,26–32 2.74, and 2.65 between 1 2 ewe .4ad38,notum 3.89, and 3.64 between r hrceie y14–21 by characterised are m 3 ,wn eghbetween length wing m, h otenfemale southern The . m 2 2 2 2 2 2 2 2 2 2 ,lnt fabdomen of length m, m 2 2 ,170,1914 4,1167 0,2521 0,1845 0,585 4,0717 4,2312 0,1577 ,390,0835 3,8214 4,4329 ,821,2018 1,1376 3,6892 3,5758 0,4606 0,134 4,2738 4,2582 0,0007 1,5073 2,7058 3,7285 2,6966 1,0229 2,9235 2 ,BV m, ,7 1,9286 0,875 ,0 0,5931 4,203 2,097 ,10,1989 0,81 1 ewe 2.16 between 2 2 2 2 2 2 2 2 2 0,2304 0,8894 0,9757 0,1132 1,7934 1,1019 1,5259 0,1391 0,8245 2 1,24 .6m,V ewe .5ad14 n eghof length 128 and and 1.43 120 and between V 1.35 palpomere between VR between mm, 1.46 90 IV palpomere and of 70 length setae, dorsocentral eaePAas cet h rpsdH proposed the accepts relationship also direct PCA a female is female of there distribution geographic and gradient specimens, character between male case the in the of As specimens. and the above of distribution mentioned geographic loadings results highest These measure- with the PCA. new ments between from relationship these direct results a from the demonstrate obtained with agrees MST for data mean The the specimens replacing character. northern by each the for with controlled values northern was missing the it of length for and and entries northern thorax abdomen, of missing length a characters to on due specimen with is specimens. result related southern remaining This the more southern with one than was specimen showing PCA, that of specimen results the with agree ‘ln’ Hxe 98 sue nodrt account to term order in the used variations. is Usually geographic 1938) microevolutionists. of (Huxley sources ‘‘cline’’ important for spatial most of the study patterns among the are reasons variation geographic these as manifested For are variation. and species indi- a within of may morphological) viduals events (e.g. phenotype these variations or Hence, produce genotype accom- 1977). small in be (Endler change may a frequencies large frequencies a that by gene panied is in change populations Mendelian spatial An and within passive. diploidy and relationships of active consequence both cap- species, important migratory affecting the different geographical of factor the abilities of is Another interbreeding degree them. sexual certain between a be isolation will this with and disrupted, of correlated be will members gender certain the opposite with the Among other. interbreeding each sexual from latter, distance occur- greater those between at between ring frequent than more individuals be neighbouring will conditions, ecological reproduction indivi- equal Given sexual among species. effective a interactions of the duals sexual regulate of that of one is facts establishment This important survive. to most ‘‘size’’ ecologi- the certain an a needs of space population cal a within individual An Discussion larval at level. character differentiation pupal geographic and a is there of Therefore, evidence groupings. no any show not did mens specimens. northern south- and ern between differences/similarities the summarizes h C efre o aveadppespeci- pupae and larvae for performed PCA The .elachistus A. m ,lnt fadmnbten12and 1.2 between abdomen of length m, pcmn n hrfr,the therefore, and specimens m .MTddnot did MST m. 0 .Table3 Downloaded By: [Donato, Mariano] At: 16:33 9 July 2008 specimens. iue2 C of PCA 2. Figure nCiooia (Sa Chironomidae in takes corre- finally which speciation cline which at broken place. stage the the the to called without sponds he third that theory Recently a stage including a 2002). speciation parapatric of presented (Salomon assumption (2002) take rejected Salomon to nowadays theory speciation cline classical is for and speciation necessary parapatric place, are barriers that new generate speciation. would parapatric by groups species and isolated process eventually evolutionary as an interpreted the of be stages could these successive variation two inter- that clinal of proposed hybrid forms (1938) a two Huxley species with zone. sub-areas polytypic gradation definite (subspecies), cline in groups breeding geographical stepped geographical definite the in comprise along and polytypic of to smoothly types gradient, refers two varying cline are smooth species a There variations: connected. clinal are when groups continuous discontin- or be the groups isolated may involves cline it if A etc.) uous axis. behavioural, geographical character a some ecological, along of genetic, values measured (phenotypic, the of variation e tde aeadesdgorpi variation geographic addressed have studies Few suggested genetics population in studies the Since gradual the as cline a defined (1938) Huxley pdlmelachistus Apedilum wdl17;Lneeg1980; Lindeberg 1979; ¨wedal eae.Mnmmsann rei ueipsd ( superimposed. is tree spanning Minimum females. tde nNorpclFuaadEnvironment and Fauna Neotropical on Studies Clfri)hv o R(.308) high (0.83–0.85), AR R on low setae of a number have (California) male adult of forms elachistus apparent two described has genus the For 1980). Savage n atws xsi ot mrc n another and Americas. America the North in axis in north–south axis way, east–west this one In between specimens. fit elachistus close southern length a and and show females VR northern for abdomen, V palpomere of of length IV, palpomere BV ftoa,nme ftmoa ea,lnt of length setae, BV temporal length, wing of III, number palpomere length setae, thorax, IX abdomen gonocoxite of of of of wing number length and number setae, and males, thorax for prealar of setae, length of VR, acrostichal length, number setae, of char- dorsocentral the number in Variation acters of variation. females geographic and present males from characters sensill (7–11). metatarsus palmate of number h i metata mid palma the of number low and 19), Foia aeahg R(.912) o number R low on (1.19–1.21), setae AR of high a have (Florida) 1 h nlsspromdhr hw htsome that shows here performed analysis The ubro oscnrlste eghof length setae, dorsocentral of number , hw w opooia aito axes, variation morphological two shows nteUA etr populations Western USA. the in + R 1 ss(–) Eas (4–5). rsus 81)adR and (8–12) m otenseies ( specimens; Northern ) + R 1 heiao h mid the on chaetica a 2–5 n R and (23–25) Apedilum esnil heiaon chaetica sensilla te 2 4 egho notum, of length , + enpopulations tern 5 02,adhigh and (0–2), pe (1988) Epler , .elachistus A. N southern ) 4 + 5 (17– 129 A. A. Downloaded By: [Donato, Mariano] At: 16:33 9 July 2008 Male metatarsus al .Cmaaiecatsoigrne en tnaddvainadnme fseiesmaue ftecharacters the of of measured specimens specimens northern of and number southern and of deviation Analysis standard Components mean, Principal range, the showing in chart used Comparative 3. Table 130 Female m nesohriestated). otherwise unless m raa ea 024415 5 1 5 1 0.03 4 3 5 1.31 7 1.35 1 4 3 5 9 5 5 0.05 11 1.28 20 0.20 0.14 0.43 5 2 1.062 5 1.38 12 6 4 24 10 0.49 4 3 4 1.23 2 4 1.48 5 0.10 10 7.14 12.50 25.32 5.42 7.07 9 0.79 0.35 10 14 16 0.83 26 72 0 1.13 35 146 32 0.94 100 1 10 10 10 16 2 173 10 32 90 45 10 105 0.08 40 7 0.60 0.07 1 0.18 0.52 1 0.99 5 2 123 12 1.78 5 5 20 63 4 30 90 11 0.60 28 9 0.27 0.06 1.08 0.33 7 2.00 10 8 1.89 9 0.80 9 9 2.69 1 8 21.08 4 13 0.37 0.88 9.34 1 2.13 8.38 13.84 1.51 0.84 6 2.97 193 5.48 1 123 98 15 R 50 9 2 1.43 48 9 0.69 VR 217 2.12 mm) (in width Wing mm) 141 (in length 114 Wing 17 71 setae 53 10 Prealar 10 159 10 11 setae Scutellar 0.22 0.16 106 setae 0.36 Dorsocentral 86 2.52 setae 13 1.11 43 Acrostichal 3.60 40 AR 8 length V Palpomere 2.81 1.39 length 4.14 IV Palpomere length III Palpomere length II Palpomere 2.22 0.93 length I 3.19 Palpomere setae Clypeal setae Temporal mm) (in length Abdomen mm) (in length Thorax mm) (in length Total ea R Setae eslacatc nmid on chaetica Sensilla SV SV SV BV BV BV LR LR LR R raa ea 4 4 0 4 0.03 4 1 4 1.39 7.25 4 2 1.43 4 4 4 3 4 0.03 8 9 0 1.35 0.07 0.51 28.5 4 4 3 4 4 0.45 4 4 1.36 4 11 0.54 7 3 32 8 0.49 3 8 2 1.43 4 3 1 0.07 3 0.48 4 1.25 6 81 23 0.41 120 55 26 27 1.26 24 1 24 1.32 3 90 2 128 4 25 58 4 4 2 33 24 3 3 28 0.08 7 3 1.19 0.11 0.27 0.15 1 0.6 113 3 70 2 0.51 19 0.19 3 53 3 3 1.81 23 2.93 23 0.13 22 9 3 1.31 0.06 0.68 9 0.17 17 0.68 3 3 1.33 3.11 1.97 3 3 0.71 4 2.04 1.43 3 2 29 0.53 1.46 10 3 18 0.43 9 0.76 1 2.17 2.62 1.68 4 10 12 186 1 13.23 1.09 14 3 111 R 90 16 Setae 43 1.2 0.65 4.04 8 40 15 1.85 8 10 98 VR 205 mm) (in width 10 Wing mm) 119 0.71 (in length 101 Wing 17 56 127 0.62 4 4 2.55 setae 113 56 4 Prealar 152 10 1.44 setae Scutellar 0.39 0.06 101 setae 131 0.39 Dorsocentral 3.23 81 2.42 setae 1.11 16 32 Acrostichal 3.53 88 2.70 30 AR 7 length V Palpomere 123 2.86 1.23 1.16 length 3.96 IV 10 Palpomere 1.04 length III Palpomere length 10 9.39 II Palpomere 1.92 1.03 length I 107.85 3.05 Palpomere 8.46 setae Clypeal 126.55 121.24 setae Temporal mm) (in length 133.87 Abdomen mm) (in 95.90 length Thorax mm) (in length Total 106.09 HV HR length Gonostylus length Gonocoxite + 4 + R 3 2 1 3 2 1 3 2 1 5 1 ea 981 5 10 8 19 0 4 1 3 4 2 setae ea 01 431 51 5 8 15 25 8 10 3 14 19 10 setae 4 .Dnt tal. et Donato M. + + R 5 1 Minimum value 82 1431 91 4 4 4 2 15 20 19 22 10 18 3 3 4 1 21 21 25 22 18 21 .529 .601 020 .326 .04 5 3 0.40 0.19 4 0.09 2.67 5 3.73 3 0.18 1.85 2.93 0.24 4 0.15 2.79 3.89 5 1.95 3.86 3 0.01 2.29 3.01 0.04 2.08 0.10 0.74 4.16 3.45 2.40 1.77 0.61 1.16 0.76 2.63 10 0.66 3.67 10 1.27 2.12 8 0.18 0.23 2.76 0.73 10 0.04 3.66 0.57 10 1.07 0.14 8 1.80 2.93 0.19 2.54 10 4.03 0.07 3.43 10 1.86 0.07 8 2.05 2.81 2.35 0.02 0.77 3.79 3.21 0.03 0.59 2.15 1.74 1.19 0.94 2.26 0.63 3.11 1.22 1.96 0.69 0.56 1.13 476110411735 ot mrcnseiesNrhAeia specimens American North specimens American South Maximum au enSD Mean value n Minimum value pdlmelachistus Apedilum Maximum au enSD Mean value maueet r in are (measurements n Downloaded By: [Donato, Mariano] At: 16:33 9 July 2008 tarsus metatarsus al .Continued. 3. Table hsseiswt h salsmn fanwentity; new a of of areas establishment extensive the however, with species this .3 ppladlra descriptions). larval and 1970, Beck, (pupal & 30 p. Beck (placement); 173 p. 1965, Sublette description). elachistus Apedilum Taxonomy an Therefore, below. given imagos. is female larva, diagnosis the species and of emended male redescription and a for pupa, call America South study. this in found variation geographic osdrtospeetu rmdsrbn new These a findings 1983). Future describing variation. clinal of (Endler of from type any us zones or species prevent transition considerations sepa- by distributions state character rated character- particular areas geographic of by a mosaic ized a addition, of consists In usually known. pattern this partially of biology is the and species samples, any by represented not 5 examined Material imma- and female and stages). male ture adult of of (redescription (misdetermination 143 88, 64, subcinctum 47, p. 1962, unsArs .B ont 34 Gonnet, Buenos Argentina, (MLP), exuviae B. pupal and larval M. 58 Aires, Buenos „„ LR hind mid on chaetica Sensilla mid on chaetica Sensilla SV SV SV BV BV BV LR LR u .elachistus A. 01 hs hrce ifrne ol edt h pi of split the to lead could differences character These nec elachista Paralauterborniella elachistus Apedilum new The pdlmelachistus Apedilum 3 2 1 3 2 1 3 2 1 n 3 and 9 34 aaatronel elachista Paralauterborniella 0 W,1May2004,leg.M.Donato;2 RR ). pdlmelachistus Apedilum ihppleuie(L) Argentina, (MLP), exuviae pupal with ilcaiytetxnmcsau fthe of status taxonomic the clarify will Townes ons pe 98 .109 p. 1988, Epler Townes: ons14,p 3(adult 33 p. 1945, Townes Minimum .elachistus A. value 72 07 72 024 4 2 3 20 21 22 25 17 18 4 4 4 2 20.75 23 24 25 17 20 .830 .400 .331 .600 4 4 4 0.03 0.02 4 0.06 3.06 4 3.92 0.07 1.86 4 3.11 0.15 4 2.92 3.94 0.04 4 1.94 4.35 0.01 4 2.69 2.99 0.00 3.03 0.69 4.54 0.04 3.89 0.56 2.74 1.8 1.14 0.69 2.85 3 0.57 4.17 3 1.17 2.65 3 0.07 0.68 3 0.13 2.94 0.56 0.16 3 3.71 1.09 0.04 3 1.84 3.02 3 0.14 2.72 3.86 3 0.11 3.80 2.03 0.02 3 2.29 2.77 2.88 0.02 0.71 3.89 3.62 0.05 0.60 2.38 1.73 1.19 0.72 2.69 0.63 3.64 1.25 2.16 0.68 0.58 1.15 Twe) ulte& Sublette (Townes): aafo southern from data ot mrcnseiesNrhAeia specimens American North specimens American South Twe) Darby (Townes): Maximum au enSD Mean value ag were range ’ u 52 „„ 9 32.8 with 0 A. S, tde nNorpclFuaadEnvironment and Fauna Neotropical on Studies Male Buchanan. A. F. leg. 1967, January 25 16 Grosso, U. Mato leg. Brazil, 1992, (JE), 57 September larvae spring, six a Nolte; near pools rock S. lrd,S.Jh’ o,S.Jh’ ie;1 river; John’s St. Co., 4 and John’s St. Florida, USA., D 1 13 light, Wibmer; at Indios, los de 1 Nolte; .0m og boe .328,23 mlong. mm 2.31 1.43–2.81, abdomen 0.69–1.39, long, Thorax mm. mm 3.29 1.00 2.12–4.14, length Total 34 Gonnet, B. M. 58 Aires, Buenos aiona meilC. itrWildlife 1 leg. Tomasello; 1979, December P. December R. 4 (residence), Wister 26 1 Beach Epler; Palm Niland, Co., H. J. Co., near leg. 1981, Area Imperial Management California, 19 shoulder, April1980,leg.J.H.Epler;1 road on swarming lighthouse, at NWR, 15 Grosso, Mato Brazil, (JE), pupae idad o nis tlgt 4Dcme 93 leg. 1983, 1 December Wibmer; J. 24 G. light, at Indios, los de Piedra a 04 e.M oao 1 Donato; M. leg. 2004, May unsArs .B ont 34 Gonnet, Argentina, B. (MLP), M. larvae six Aires, Donato; Buenos M. leg. 58 ie,M .Gne,34 Gonnet, B. M. Aires, ( exuviae pupal and larval ont 34 Gonnet, 1 Donato; retn,Beo ie,M .Gonnet, B. M. (MLP), Aires, larvae five Donato; Buenos 34 M. leg. Argentina, 2004, May ie,M .Gne,34 Gonnet, B. M. Aires, u u u u ,30mal okpools, rock asl, m 300 W, 01 01 52 RR n 9 9 9 ( 34 34 32.8 n 5 „ J) S,Foia aul o,S.Marks St. Co., Wakulla Florida, USA, (JE), 0 0 ,1 a 04 e.M oao three Donato; M. leg. 2004, May 16 W, ,1 a 04 e.M oao 1 Donato; M. leg. 2004, May 12 W, 5ecp hnohriestated) otherwise when except 15 „ J) rga,Clna ot 1 Piedra 21, Route Colonia, Uruguay, (JE), 0 Minimum „ ,58 S, u MP,Agnia unsArs .B. M. Aires, Buenos Argentina, (MLP), 52 value J) rga,Clna ot 21, Route Colonia, Uruguay, (JE), 9 „ 32.8 u ihlra n ua xva (JE), exuviae pupal and larval with 01 „ 0 9 ,58 S, 34 J) S,SokIln BLT, Island Stock USA, (JE), Maximum 0 au enSD Mean value ,1My20,lg M. leg. 2004, May 1 W, u „ u 52 L) retn,Buenos Argentina, MLP), u 52 01 F) S,Pl Beach Palm USA, (FS), cme 93 e.G J. G. leg. 1983, ecember 9 9 32.8 9 32.8 6My19,lg U. leg. 1992, May 26 34 „ 0 MP,Argentina, (MLP), 0 ,1 a 2004, May 16 W, 0 ,58 S, ,58 S, u „ 57 u u 9 01 J) USA, (JE), u 52 ,57 S, 01 u 52 9 9 34 9 34 9 u 32.8 32.8 0 R 45 ,10 W, 0 ,1 W, with 9 n 131 W, u 0 0 S, S, S, „ Downloaded By: [Donato, Mariano] At: 16:33 9 July 2008 019 4() 1–0,13(6). 153 113–205, (7); lengths 94 Palpomere 70–119, (7). setae 20 (in 16–25, with clypeus Head males. 1.2–2.86, in as length Coloration Abdomen (7). length mm (7). Thorax 1.95 (7). mm 0.93 mm 2.88 0.65–1.16, 1.85–3.96, length Total Female (10). 2.55 (10) 0–3,17(10) 127 106–134, traoee5–3 5(10) 65 50–83, sternapodeme ,5(0 ea,ltrsent Xwt – ea (9). setae 1–2 with 64 IX 48–83, laterosternite setae, Phallapodeme (10) 5 6, Hypopygium and 4. Table Lengths in metatarsus. as legs hind of proportion on 0–1 metatarsus, Legs iue3 aewn of wing Male 3. Figure lengths Palpomere setae. 15 (in 12–17, with clypeus (13); Head 3. Figure in as grey spotted pale whitish with membrane dark Wing mostly whitish. abdomen legs brown; and thorax head, Coloration: 132 .506 .9m.V .–.5 .6 R 0.96. 0.6–1.35, VR mm. 0.49 0.35–0.6, Wing (14). setae 7 6–9, with Scutellum 2. 1–4, prealars Thorax (10). 179 123–217, (13); 116 90–141, (13); 5 4ste R setae; 14 25, m m m amt eslacatc:41,7o mid on 7 4–11, chaetica: sensilla Palmate . ) 25,3 7;2–6 4() 311 0(7); 70 53–101, (7); 34 23–56, (7); 31 22–56, m): ) 85,4 1) 07,4 1) 314 90 63–114, (13); 46 30–71, (12); 42 28–53, m): R04–.8 .8 eprlste72,16; 7–24, setae Temporal 0.48. 0.41–0.68, AR . R08–.3 .2 eprlste83,15 8–32, setae Temporal 1.02. 0.83–1.23, AR . og R10–.,14 1) V1.23–3.23, HV (10); 1.44 1.04–2.7, HR long. m Figure cotcas21,5 oscnrl –4 14; 9–24, dorsocentrals 5; 2–12, Acrostichals . ( .Dnt tal. et Donato M. n 5 xetwe tews stated) otherwise when except 8 ihu nlpit egmI ih4– with IX Tergum point. anal Without . ( 3 4 ) + iglnt .32 .4m;width mm; 1.64 1.13–2, length Wing . 5 ih01,5()setae. (9) 5 0–19, with m og ootls8–2,108 88–121, Gonostylus long. m pdlmelachistus Apedilum m og(0;transverse (10); long m m og Gonocoxite long. m . + R 1 ih8– with .806,05 7 m R11–.3 .3() R (7). 1.33 1.19–1.43, VR mm. (7) 0.54 0.48–0.68, setae. Wing 7 4–9, with Scutellum (7). 3 2–4, prealars 23; Thorax 016 98 80–116, adbelnt 0–4,121 103–149, length Mandible umre eeain h pce a lofudi a in found also from was species larvae the of vegetation; collection submerged the mentioned (1988) Epler Biology 4–10, 9; 6–13, 1.05. 17; 0.89–1.26, 12–25, AR 20; 6. 15–25, 18; (in 5.02 13–23, segments 73; 3.9–7.15, 96, antennal PSR of (10); Length 18.59 (10). 11.3–25.2, IPD (12); 83 103, Head stated) orhisa larva instar Fourth 1.65–3.20, DR (12). (8). setae 35 26 26–52, 17–32, II 2.23. T with of lobe of margin Anal row posterior Transverse on (8). hooklets abdo- mm 2.77 (8); (8); 1.68–3.2, length mm mm 1.01 3.3 men 0.66–1.24, 1.98–4.25, length length cephalothorax Total hyaline. Color: Pupa setae 1–2 with IX (5). gonocoxite one setae; each 7 6–9, with VIII ih1–2 1ste() R (7). setae 21 18–22, with 2,94 121, Abdomen 4. Table in shown are legs of Ta Legs 1 72,2,o idTa hind on 20, 17–24, ; amt eslacatc:1–5 2 nmiddle on 22, 18–25, chaetica: sensilla Palmate . iglnt .619,15 7 m width mm; (7) 1.55 1.26–1.97, length Wing . otetmlnt 1–5,134 113–157, length Postmentum . ( n cotcas61,9 oscnrl 15–32, dorsocentrals 9; 6–11, Acrostichals . m m 5 ;lnt 25,41 32–55, length m; og egmXwt –1 ea 7.S (7). setae 9 2–21, with X Tergum long. m ou 6–0,183 162–205, Notum . 4ecp hnohriestated) otherwise when except 14 m 1) etoetlpaewdh69– width plate Ventromental (12). m ( n 5 6ecp hnotherwise when except 16 4 + 1 5 egh n proportion and Lengths . m ih1–5 8ste(7). setae 18 10–25, with .VR15–.1 2.01 1.57–2.41, VPR m. m m .Mnu width Mentum m. og ec 73– cerci long; m m )(4:60– (14): m) m (15). m + R 1 Downloaded By: [Donato, Mariano] At: 16:33 9 July 2008

Table 4. Lengths (mm; ranges followed by the mean) and proportion of legs of Apedilum elachistus.

Fe Ti Ta1 Ta2 Ta3 Ta4 Ta5 LR BV SV

Male P1 500–803, 689 415–681, 596 590–824, 726 280–427, 369 220–346, 288 135–254, 195 90–173, 121 1.07–1.27, 1.18 1.96–2.4, 2.12 1.74–1.95, 1.82 (14) (11) (11) (11) (11) (11) (11) (n511) (n511) P2 560–895, 751 440–742, 611 290–427, 370 130–224, 185 90–163, 136 50–112, 88 40–102, 80 0.56–0.66, 0.6 3.11–4.16, 3.58 3.21–4.03, 3.68 P3 580–946, 786 500–844, 703 380–620, 541 190–356, 297 170–305, 262 90–153, 131 60–122, 98 0.69–0.94, 0.77 2.26–3.01, 2.61 2.08–2.93, 2.73 (14) (14) (14) (14) (14) (14) (14) (14) (n514) (n514) Female P1 460–864, 597 415–651, 515 470–814, 600 195–376, 271 140–275, 196 90–183, 131 65–132, 95 1.09–1.25, 1.16 2.16–2.74, 2.52 1.73–2.03, 1.85 P2 520–915, 680 420–732, 553 240–427, 324 105–203, 147 70–147, 104 40–102, 66 45–97, 69 0.56–0.63, 0.58 3.64–4.54, 4.11 3.62–3.94, 3.83 P3 560–915, 709 510–864, 656 350–590, 455 185–315, 244 160–285, 216 70–142, 103 60–112, 84 0.68–0.72, 0.69 2.69–2.99, 2.83 2.88–3.11, 3.01

Male, n515; female, n57, except when otherwise stated. tde nNorpclFuaadEnvironment and Fauna Neotropical on Studies pe H 98 eosdrto ftegenus the of reconsideration A 1988. JH. Epler esma L atn mrmR erqe lviaAL, Oliveira Henriques R, Amorim Martins JL, Nessimian and European some in variation Qualitative 1980. B. Lindeberg Nature. principle. single taxonomic auxiliary and an Clines: trees 1938. spanning to JS. Huxley Minimum approach 1969. statistical JS. new Ross A JC, Gower 1969. RR. Sokal KR, Gabriel pe H 97 eiino h Nearctic the of Revision 1987. JH. Epler of study the in hypotheses causal Testing clines. 1983. and JA. speciation, Endler variation, Geographic 1977. JA. some Endler of stages with immature fields, rice California The with associated Midges 1962. 1970. RE. Darby EC. Beck WM, Beck References manu- and 7646-04. this CONICET Grant by of Society supported Geographic version National was for earlier work Crisci This an V. script. Jorge on thank comments to his wishes author first The biofilm-covered Acknowledgments with pool artificial was material an substratum. new The in 2003). rhithral found al. a et in (Nessimian vegetation zone submerged and 1995) (Nolte southern elachistus Likewise, (USA). A. Florida in pond brackish aaeHS 90 igcaatrvraini h Nearctic the in variation character Wing for 1980. used HMS. be can Savage that theory cline revised A 2002. M. Salomon termino- morphology chironomid of Glossary 1980. OA. Sæther nume pc, NT-SYS days: 1998. seven FJ. than Rohlf less in imago to egg From 1995. U. Nolte 93(itr:Ciooia) vlMng.9:1–139. Monogr. Evol Chironomidae). (Diptera: 1913 i eJnio eatmnodsge dos Levantamento do Janeiro. Estado de do (Diptera) Rio Chironomidae 2003. AM. Sanseverino 1978(1/2):115–121. Biol. Carol of Univ Acta Chironomidae). populations Nearctic 10:54–64. C. 42:219. St R J Stat Appl analysis. cluster linkage 18:259–278. Zool. Syst analysis. variation geographic 14:105–116. Suppl. Spixiana Chironomidae). (Diptera, 1945 nttt eisG(clgclsine) ubr1). number Study sciences); Advanced (Ecological (NATO G series Institute. Numerical 424–443, Institute Study p. editor. Advanced Springer-Verlag. NATO J, Berlin: a of Felsenstein proceedings In: taxonomy: variation. geographical 262. p. Press. University Princeton (NJ): Chironomidae). Princeton (Diptera: ecology their 32:1–206. to Hilgardia. reference special 33:29–42. Sci. Acad Fla Jl Q (Chironomidae). Chironomini hrnmde:apicplcmoet nlss e Am Mem analysis. 1983:299–308. components Soc. principal Entomol a Chironomidae): of species 29:509–517. Biogeogr. parapatric J without speciation. processes evolutionary of analyses quantified 14:1– Suppl. Scand Entomol 51. Chironomidae). (Diptera: logy Exeter (NY): Setauket 2.0. Version Software. system. analysis variate Melbourne: East ecosystems. 177–184. p. Publications. to CSIRO genes Chironomids—from elachistus Apedilum ocorre ni.Pb vlMsNcRoJ 98:1–16. J. Rio Nac Mus Avul Publ ˆncia. rhcais(Orthocladius) Orthocladius eefudi peea okpools rock ephemeral in found were Ciooia) n rntnP,editor. PS, Cranston In: (Chironomidae). ayassgracilentus Tanytarsus ia aooyadmulti- and taxonomy rical a e up(Diptera: Wulp der van nrseha e ˆneros Dicrotendipes Apedilum btt de ´bitats (Diptera, Townes, Kieffer, 133 Downloaded By: [Donato, Mariano] At: 16:33 9 July 2008 pe ,RisF 96 aao n ilorpyo Neotropical of bibliography and Catalog 1996. F. Reiss M, Spies Sa 134 wdlL 1979. L. ¨wedal n eia hrnmde(net,Dpea.Siin Suppl. Spixiana Diptera). 22:61–119. (Insecta, Chironomidae Mexican and the in insignilobus variation geographical and Chironomidae) .Dnt tal. et Donato M. grgt.EtmlSadSpl 10:133–138. Suppl. Scand Entomol aggregate. irpetabrundini Micropsectra .s.(Diptera: sp. n. Micropsectra onsH.14.TeNaci pce fTnieii mMidl Am Tendipedini. of species Nearctic The 1945. HK. Townes Stone In: Chironomidae. Family 1965. MS. Sublette JE, Sublette a.34:1–206. Nat. Dep S U Mexico. 276:142–181. of north Handb, America Agric of Agric A Diptera editors. the JR, Coulson of RH, catalogue Foote WW, Wirth CW, Sabrosky A,