Revised phylogenetic analysis of Indian species of genus Macrophya Dahlbom (Hymenoptera: Symphyta; Tenthredinidae: Tenthredininae) M.S. Saini1 and L. Kaur2 DepartmentofZoology, Punjabi University, Patiala, India. 1email: [email protected]) ( 2email: [email protected]) ( Abstract Revisedphylogeneticanalysiswasperformedfor14speciesofthegenusMacrophya(Hymenoptera: Tenthredinidae) usingthe phylogenetic analysis package PAUP, based on 15ofthe morphological characters mostcommonlyusedforMacrophya species identification.Speciesdescriptionswere derived primarily from "Indian Sawflies Biodiversity" vol. II Saini (2007), Saini and Bharti (1996), Saini efa/. (1996), and Saini and Vasu (1997). The results are discussed in terms ofevolutionary trends or biological maxim that "nature prefer to modify the already existing structures so as to copewith new needs." Keywords: Phylogenetic analysis, Macrophya, Evolutionary trends. Introduction ThegenusMacrophya(Hymenoptera:Tenthredinidae) The purpose of present study is to trace the long iswidelydistributedgenuswithitsrepresentativesavailablein evolutionary history which modified generalizations into almostallmainregionsoftheglobe.Withregardtoitsaffinities, specializationsofextremeform tosuitthecircumstances in it shares most of its characters with Pachyprotasis Hartig. which subsequently insects dwelled. Parsimony analysis is EvenwithinMacrophyatherangeofcharactersissowidethat used to investigate phylogenetic relationships among timetotimemanyofitssubgenerawereproposed(Malaise, Macrophya species, using data based on morphological 1945)andbecauseofnodistinctboundariestheyallgotmerged charactersmostcommonlyusedforMacrophyaidentification. (Ross, 1937; Gibson, 1980). Today noneofits subgenus is With the help of literature contributed by Saini efa/. (1996) considered to be valid (Abe and Smith,1991). The genus and Saini and Vasu (1997) some different characterswere Macrophya was first described by Dahlbom (1835) as a usedtotracephylogenyofgenusMacrophya. subgenusofTenthredoLinnaeus,onthebasisofbodyshape, Materials and Methods lengthandformofantenna. Hedividedthissubgenusintotwo Speciesdescriptionswerederived primarilyfrom subsections"A"and"B".Hartig(1837)appliednamestothese "Indian Sawflies Biodiversity" vol. II Saini (2007), Saini twosubsectionsusing T.(Macrophya)forsubsection"B"and and Bharti (1996), Sainiera/. (1996), and SainiandVasu T.(M.)(Pachyprotasis)forsubsection"A". Bothofthesewere (1997)andthecharactersused intheanalysiswerethose laterrecognisedasvalidgenerabyWestwood(1840). givencomparablyforall, oralmostall, species. Tenthredo The genus is characterized by venation as in rufipodusSainiand Bharti, 1996wasalso included inthe Pachyprotasis, buttheanalcellmayhaveacrossvein.Malar analysisastheoutgroup. Revised phylogeneticanalysis space mostly shorter than the diameterof an ocellus. The wascarriedwith a computerprogram PAUPversion4.0, hindlegsarestronglybuilt,andthekneesreachingandmostly with morecharactersascomparedtoearlierstudy (Saini exceedingtheapexoftheabdomen(Saini,2007).Thelarval and Kaur, 2010), has been used to infer the phylogeny stagesfeedonvarietyofwildherbs,shrubsandtrees.Generally (Swofford, 2000). In total 15 morphological characters adultsfeedonpollens,flowernectarorleafjuiceexudingfrom were used in the phylogenetic analysis Thesewere: woundscausedbystrongmandibles. However, manyrobust LForewing: (0-without infuscated band, 1-with infuscated speciesindulgeinzoophagy(Cameron,1882;Rohwer,1913; band) Benson,1938; Malaise,1945;Naito,1988andGoulet,1996). 2.Bodycolour: (0-notmetallicblue, 1-metallicblue) ) 54 Halteres, Vol.3, 2011 3.Clypeus: (0-not triangularly incised up to 1/4 of its 10. Circumocellar furrow: (0-fine or clear, 1-distinct, length, 1- triangularly incised up to 1/4 of its length) 2-indistinct, 3-sharp) 4. Frontal area: (0-not below level of eyes, 1-below 11. Postocellar area: (0-flat, 1-subconvex, 2-slightly level ofeyes) raised, 3-fine 5. Average length of body of female: (0-10 mm, 1- 12. Mesoscutellum: (0-slightly or roundly raised or above 10mm, 2-below 10mm) elevated, 1-subconvex, 2-prismatic, 3- flat) 6. Subapical teeth of tarsal claw: (0-smaller than 13. Mesepisternum: (0-roundly raised, 1-obtusely apical teeth, 1-longerorequal to apical teeth) raised, 2-depressed notobtusely raised) 7. Labrum: (0-with rounded anterior margin, 1-with 14. Antenna length: ( 0-2 times or more than 2 times truncate or subtruncate anterior margin, 2-with slightly of headwidth;1-less than 2 times of head width) emarginateanteriormargin) 15. Hind femora: (0-hardly reaching apexofabdomen, 8. Median fovea: (0-absent, 1-distinct, 2-indistinct) 1-exceeding apex ofabdomen) 9. Supraantennaltubercles: (0-insignificant, indistinct or not raised, 1-raised orsignificant) Table-1: Presence orabsence data forfifteen characters for 14species ofthe genus Macrophyaas used in the phylogenetic analysis; T. rufipodus Saini and Bharti, 1996 is included as an outgroup. 111111 Cr\cifioc \1/ 1 2 3 4 5 6 7 8 9 0 1 2 3 4 M. andreasi Saini and Vasu, 1997 0 0 0 0 2 1 0 1 0 3 1 0 - 1 1 M. brancuccii Muche, 1983 0 0 0 0 1 1 1 0 0 0 0 0 1 1 1 M. formosana Rohwer, 1916 0 0 0 0 1 1 2 1 3 1 0 0 1 M. gopeshwan Saini etal., 1986 0 0 0 1 2 1 1 1 0 0 2 1 0 M. khasiana Saini etal., 1996 0 0 0 1 0 1 1 1 0 1 0 1 1 M. maculicornis Cameron, 1899 0 0 0 1 1 0 1 1 0 0 0 0 0 M. manganensis Saini efal., 1996 0 0 1 0 0 2 2 1 3 0 0 1 1 M. naga Saini and Vasu, 1997 1 0 0 0 2 1 2 0 1 1 1 0 M. planata (Mocsary, 1909) 0 0 0 1 1 0 2 1 2 1 2 1 0 M. pompilina Malaise, 1945 1 1 0 1 0 0 2 1 3 1 1 0 0 M. pseudoplanata Saini etal, 1996 0 0 0 1 1 0 1 0 1 0 3 1 0 M. regia Forsius, 1930 0 1 0 1 2 1 0 0 3 2 0 0 1 M. rufipodus Saini etal., 1996 0 0 0 0 2 1 2 1 1 0 1 0 0 M verticalis Konow, 1998 0 0 0 1 2 0 1 0 1 0 0 0 1 1 T. rufipodus Saini and Bharti,1996 0 0 0 0 2 0 0 1 1 1 3 0 2 1 0 RevisedphylogeneticanalysisofIndianspeciesofgenusMacrophya Dahlbom... 55 Discussion The outgroup taxon T. rufipodus got separated apex ofabdomen (Character 15, state 0). The derived from all species of genus Macrophya on the basis of consensus tree identified two groupings to be present postocellar area fine (Character 11, state 3), in the cladogram. In the first group M. khasiana got mesepisternum depressed not obtusely raised isolated from M. verticalis, M. brancuccii, M. regia, M. (Character 13, state2) and hindfemora hardly reaching gopeshwari, M. planata, M. maculicornis and Results +++ T. rufipodus C>2 5>2 1>5 M khasiana 6 9 4+ M verticalis 2 4 5'! I I 11 Mbrancuccu 2>; M regia 5 7 44 M gopeshwari 5>2 2>1 T8I10T12 M planata 9 151215 ;>2:>2i>2 1W11W1 M maculicornis M pseudoplanata 3 7 44-T M manganensis 0>U>2 2>1 5 12': III M formosana 2 4 57 ': 2>1J>30>1 H § I | M pompilina 0>10>M>CO31911 M naga M rufipodus - 1 i i i - m andreasi Fig. 1. Strictconsensustreefor 14 species ofMacrophyaderived from the 7 most parsimonioustrees calculated from the data in Table 1 ; outgroup = Tenthredorufipodus Saini and Bharti,1996 . Characterofthe ingroup have been optimized by fasttransformation as implemented in PAUP. Character numbers are above the hashmarks; state changesareshown belowwith the respective primitiveand derived conditions reported bya '>'. 7 treeswere found using the computer program PAUP version 4.0, with the shortest tree requiring 28 steps, a consistency index ( CI ) of0.50, a retention index ( Rl) of0.55 and rescaled consistency index ( RC) of 0.28. 56 Halteres, Vol.3, 2011 M. pseudoplanataonthe basisofaverage length ofbody classificationoftheTenthredinini(Hymenoptera, Symphyta, offemale is 10mm Character5, state 0). M. verticalis, Tenthredinidae). Contributions of the American ( M. brancucciiand M. regia due to absence of median Entomological Institute, Gainesville 29(2): 1-135. foveagotseparated from M. gopeshwari, M. planata, M. Hartig, T. 1837. Die Familien der Blattwespen und Holzwespen. maculicornisand M. pseudoplanata. M. brancucciiand Berlin Haude und Spener. M. regia are sister species and got isolated from M. : verticalisdue to characters6 and 9. M. gopeshwarican Malaise, R. 1945. Tenthredinoidea ofSouth-EasternAsiawith a be distinguished from M. planata, M. maculicornisand general zoogeographical review. Opuscula Entomologica, M. pseudoplanata duetoaverage length ofbody below Lund supplement 4: 1-288. 10mm (Character5, state2)and labrumwith subtruncate anterior margin (Character 7, state 1). M. planata on Naito, T. 1988. Systematic position of the genus Rocalia basis ofcharacters8, 10and 12 gotisolated from sister (Hymenoptera,Tenthredinidae)feedingonfernspores,with species M. maculicornisand M. pseudoplanata. Dueto description of a new species from Japan. Kontyu 56(4): 798-804. triangularly incision in clypeus, labrum with slightly emarginated anterior margin and mesepisternum Rohwer, SA. 1913. Notesonthefeeding habitsofadultsawflies obtusely raised M. manganensisgotseparated from M. (Hymenoptera: Symphyta). Proceedings of the formosana, M. pompilina, M. nagaand M. rufipodusand Entomological Society ofWashington 15: 148-149. all these species showed monophyly as indicated by homologue 6. M. naga and M. rufipodusat the base of Ross, H.H. 1937.Agenericclassificationofthe Nearcticsawflies cladogram got separated from closely related species (Hymenoptera: Symphyta). Illinois Biological Monographs, M. pompilina due to characters 2, 4, 5, 7 and 10. M. 34: 173. naga and M. rufipodus on basis ofcharacters 1, 9 and Saini, M.S.2007. IndianSawfliesBiodiversity(Keys,Catalogue& 11 got separated from each other. So, extremely specializedformsdescended bygradual changes leads ITlelnustthrraetidoonsL).vDoel.hrIIa.dSuunb:faBmiislhyenTeSnitnhgrhe,dMianihneanedrSaanPsalGSeinnugsh to accumulation of certain appropriate features which Publishers. represents body organization acquired to become complex so as to meet requirements which also Saini,M.S.andBharti,H. 1996.Additionoffivenewspeciestothe underlies the biological maxim. Indian TenthredoLinnaeus (Tenthredinidae, Hymenoptera) from North-westIndia.AnnalsofEntomology14(2):61-68. References Saini, M.S., Bharti, H. and Singh, D. 1996.Taxonomicrevisionof Abe, M. and Smith, D.R. 1991. The genus-group names of genus Macrophya Dalhbom (Hymenoptera, Symphyta, Symphyta (Hymenoptera) and their type species. Esakia, Tenthredinidae; Tenthredininae) from India. Deutsche Fukuoka 31: 1-115. Entomologische Zeitschrift 43(1): 129-154. Benson,R.B. 1938.Ontheclassificationofsawflies(Hymenoptera, Saini, M.S. and Kaur, L. 2010. Phylogenetic analysis of Indian Symphyta).TheEntomologist'smonthlyMagazine75: 110- species of genus Macrophya Dahlbom (Hymenoptera: 113. Symphyta;Tenthredinidae:Tenthredininae). Halteres2:24- Cameron,P. 1882. A monograph of the British phytophagous 28. Hymenopteravol. I. London:TheRaySociety. Saini, M.S. and Vasu, V. 1997. Present position of Macrophya Dahlbom, A.1835. Conspectus Tenthredinidum, Siricidum, Dahlbom from India (Hymenoptera, Tenthredinidae; Oryssinorum, Scandinaviae,quasHymenopterumfamilias. Tenthredininae). Journal ofEntomological Research21(3): Hafiniae: Kongl Swenska Wetenskaps Academiens 237-243. Handlingar 1-16. Swofford, D.L. 2000. PAUP: Phylogenetic analysis using Gibson,A.R 1980.Arevision ofthe genus Macrophya Dahlbom parsimony version 4.0. USA: Sinauer Associates, (Hymenoptera:Symphyta:Tenthredinidae)ofNorthAmerica. Sunderland, Massachusetts. Memoisofthe Entomological Society ofCanada 114: 166. Westwood,J.0. 1840. SynopsisoftheGeneraofBritish Insecta. Goulet, H. 1996. Revision ofthe Nearcticspecies ofthe arcuata In: Westwood, J O. (ed.). An Introduction to Modern group of the genus Tenthredo with notes on the higher Classification of Insects, 2: 158.