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Presented to the University of Manitoba @ Darren Andrew Poilock CIassif ication, reconstructed phylogeny' and zoogeography of Petho Latreill'e (coleopteia: Heteromera: Pythidae ) based on study of }arval, pupal, and adulÈ stages by Darren Àndrerl PoIlock À lhesís presented to the University of Manitoba in fulf i lIment of the thesis requirement for the degree of Master of Science in Department of Entomology Un iversi !y of I'lanítoba. Wi nn i peg ' Man i loba @ Darren Andrew PoIlock' 1988 Perníssíon has been granted L'autorisation a êtê accordée to the National Líbrary of å la Bibliothèque nationale canada to nicrofiln t.hIc du canaCla de nícrofilner thesls and to lend or eel1 cette thèse et de prêter ou copiee of the filnr. de vendre dee exemplaires du fil¡¡. The author (copyríght owner ) L'auteur ( titulaire du droit has reserved ot.her d'auteur) se rêserve leB publ lcat ion ríghts, and autres droits de publ Ícat lon i neither the thesís nor ni la t.hèee ni de longe extensive extractc from ít extralts de ceIIe-ci ne nay be prínted or otherwise clo l vent etre lnprrmea ou reproduceil wlthout hi e/her autrenent reproduits 6anB son written per4i seíon. autorlsation êcr íte. rsBN o-3L5-47944-2 CLASSIFICATION, R,ECONSTRUCTED PHYLOGENY, AND ZOOGEOGRAPHY OF PYTHO LATREILLE (COLEOPTERA¡ HETEROMERA: PYTIIIDAE) BASED ON STUDY OF LARVAL, PUPÀL, AND ADULT STAGES BY DARREN ANDREÍ.I POLLOCK A thesis submitted to the Faculty of Graduate Studies of the University of Manitoba in partial fulfillment of the requirentents of the degree of }ÍASTER OF SCIENCE o 1988 Permissiou has been grairted to the LIBRARY OF THE UNMR- SITY OF MANITOBA to lend or sell copies of this thesis. to the NATIONAL LIBRARY OF CANADA tO MiC¡OfiINì this thesis and to lend or sell copies of the film, and UNIVERSITY lVf ICROFILMS to publish an abstract of this thesis. The author reserves other publication rights, and neither the thesis nor extensive extracts from it may be printed o¡ other- wise reproduced without the autho¡'s written pernrissiorr. I hereby declare lhat I am the sole author of this thesis' I authorize the University of Manitoba to lend this thesis to other irifil"llãr, oi indíuidruis for rhe purpose of scholarly research. Darren Àndrer¡ Polloc k I further authorize lhe university of l,!aniEoba to reproduce lhis thesis ¡v-piróio"opying or by other means,- in-totaL or in part, at. the request ;i ãlh;; i'nåti[utionË or individuats for the purpose of scholarlv research. Darren Àndrelr Pollock The UnÍversity of Manitoba requires the signatures of al1 persons using åi-pl,oioãopyii,9 this thesis. PIease sign below, and give address and date. ÀCKNOWLEDGMENTS I an very grateful for the kind assislance received from the various private collectors and curalors of museum coLlections, whose specimens formed the basis of my study. I am especially indebted to the following curators who loaned me type maleriall L. Jessop (BMNH)r S' Koponen (naru), s. Takagi (HUs), o. Merkl (TMB), s.R. shaw (Mcz)' H' silfverberg (zMH), F. Hieke (ZMHB), and G. scherer (zsl't) . Throughout my study, I obLained gifts of specimens and advice, both of which were invaluable' I should Like to !hank the following for !heir generous contributions: B. and J. Carrr R.C. Crawr R.À. Cro!¡son' N. Hayashi r S' Hisamatsu, J'F' Larlrence, R' Pettersson, and D.K. Young. Dean of AgriculLure R'C' McGinnis provided part of my financial support during my programme, and for his assistance I am very grateful' I would especially Like to thank R.E' Roughley, who has allowed ne to explore an interesting group of beetles, and also has aIlor¡ed the academic Ìatitude needed to appreciate systemalic research. I]¡ithout his financial and other support, I surely would not have been able to carry out this study. To the other menbers of ny committee, Drs' T'D' Galloway and B. J. Hannr I express rny gratitude for suggestions and guidance. For obtaining many references, I should like to thank profusely, the diligenl workers in the Interlibrary Loan Depârtnent of Etizabelh Dafoe Library. For help tiith translation, I thank RoIf Beutel, Ingolf Àskevold, Germaine Lé9er, and Roger Pettersson' I âm lV - qrateful- to Dr. B. Dronzek and B' Luit for the use of the scannlng electron microscope. Finally, I thank Lisa Reíchert, for helping vith the illustrations and he).ping return foans' but especialLy for her understanding and patience lhrough the past several years' TÀBI.,E OF CONTENTS pa9e x11 xLlL LIST OF FIGI'RES xvll ÀBSTRÀCT " " 1 I. iNÍRODUCTION OF PYIHIDÀE 3 II. CONSTIN'ENTS ÀND SYSTEMÀTIC PLÀCEMENT I III. NÀnJRÀL HISToRY OF PYTHO tÞ IV. HISTORICÀ! REVIEI{ 22 V. MÀTERIÀLS ÀND I'ÍETHODS 22 5.1 MÀTERIÀLS 28 5.2 UETHoDS 28 5,2.1 Generatized methods - a-!! staoes 5.2.2 Generalized nethods - irunature ståqes ' 5,2.3 Generalizedl nethods - adlul! staoe ' ' ' ' 5.3 SPECIES ÀND SPECIES-GRoUP coNcEPTg 36 5.3.1 Speciee conceDt8 5.3.2 SPecies-orouo conceÞt VI . REÀRING METHODS VII. TÀXONOI.IIC CITÀRÀCTERS ÀI¡D TERI'ÍS 7.'1 ÀDI'LT STÀGE 7.2 T.,ÀRVAL SBÀGE q¿ 7.3 EgPÀt STÀGE VIII, CI'ÀSSIFICÀTION 8.1 ÀDI'LT STÀGE 58 8.2 LÀRVÀIJ STÀGE , 64 66 8.3 PUEÀ! srÀGE , 59 8.4 EEyg ro IloRLp SPECIES OF PYrflo 8.4.1 Kev to ailulÈg 69 8,4.2 Kev to mature larvae ' ' ' 73 8.4.3 Kev to knonn ouoae 75 IX. INDIVIDUÀI., SPECIES TREATI{ENTS ' 17 9.1 ryIË9. SEIDLITZI Blair, 1925 ' 17 9.1,1 Adlult stãqê . , ' ' ' ' 1'1 9.1.2 Larval staoe 81 9.1.3 EuEgI staoe . 83 9.1,4 Natural historv 84 9.1 .5 Distribuhion 85 9.1,6 ChoroLooical relationshios 86 9.1.7 PhfÀSS$e!þ relationshios 86 9.2 PYTHO STRICTUS Leconte' 1866 87 9.2.1 ÀÉlult 6taqe . 87 9.2.2 L¡rval staoe 91 9.2.3 PgBI staoe . ' 92 9.2.4 Natural historv 93 9.2.5 Dislribubion 94 9.2,6 Chorotooical relationships ' 94 9. 2.7 Phvlooenetic relationshios 94 95 9.3 PYTHO KOLI|ENSIS C. Sahlberg, 1833 9.3. 1 Àdlult 6taae . 95 9.3.2 Larval ståqe ' 100 - v11 9.3.3 Puoa!. staoe . 101 9. 3.4 Natural hiatory l03 9. 3. s Distribution 103 9.3.6 Choroloqical relationships 104 9.3.7 Phvlooenetic relationships 104 9.4 PlEg NMLIS Lewis, 1888 105 '105 9.4. 1 Àclult staqe . 9.4.2 Larval staoe 110 9.4.3 Puoal staoe . 112 9.4.4 Natural historv 112 9.4, 5 Díslribution 112 9.4.6 9þ¡-9I9S-!Sê! relationshios 113 9.4.7 PhyLgSlEË-g retationshios 113 9.5 ryIË9. NIGER KirbY, 1837 114 9.5.1 ÀdluLt staoe . 114 9. 5.2 L,arval staoe 120 121 9,5.3 PUpeI slaqe . 9. 5.4 Natural hi storv 122 9, 5.5 Di stribution 123 9. 5.6 Choroloqícal relationshi¡s 124 9. 5. ? Phyl-gggtg!!-q relatíonshipE 124 125 9.6 9ITH9. ÀBIETICOLÀ J. Sahlberg' 1875 . : ' ' ' 12s 9.5. 1 Àdult staoe . 9.5.2 L,arval Etaoe 130 9.6.3 Plpel staqe . 9.6.4 Natural Hisbore 133 171 9. 6.5 Di str ibution - vI11 - 9.6.6 Choroloqical relationshios 134 9.6.7 Phvlooenetic relationshios 134 135 9.7 EIIË9. JEZOENSIS Kôno, 1936 9.7.1 Àdult staoe . 135 9.?.2 Larva1 g9¡gg 139 9.7.3 Puoa1 6taqè . 140 9.7 .4 Natural hi sÈorv 140 9.7. 5 Distr ibuÈion 141 9. 7.6 Chorolooical relationshios 141 9. 7. 7 PjylgSgqÊlj-q relationshios 141 142 9.8 PY'rH9. E!å!Ug (oIivier, 1795) 9.8. 1 Àdult BÈaoe . 143 9.8.2 Larval gLagg 149 9.8. 3 Puoal staoe . tJt 9.8.4 Natural hi6lorv 1q) I Ea 9,8.5 Di sÈribuLion '1 9.8. 6 Chorolooical relat ionshios 53 9.8.7 PhvloqeneÈic relationshios 153 154 9.9 ryrHg. DEPRESSUS (Linnaeus, 1767) I EC 9.9. 1 Àdu1t staoe . 9.9.2 Larval staqe 1s8 9,9.3 Elpd staqe . 160 9.9.4 þlglel historv tbt 9.9. 5 DisLribulion 162 9,9.6 Choroloqical relationshios 162 9.9.7 Phvloqenetic relationships 162 9.10 SPECIES OF I'NCERTÀIN PLÀCEMENT 163 164 X, PHYLOGENY '1 164 0,1 PYTHINÀE ÀS À MONOPHYLETIC IJNIT 10.2 PYTHo ÀS à I,foNoPHyLETIC tNiT 165 10.3 EIË98 tb5 169 10.4 ÀNÀLYSIS OF CHÀRÀCTERS 10.4. 1 L,arval character6 159 10.4.2 Àdlult characters 171 177 XI . ZOOGEOGRÀPHY 17't 11.1 !N!89Duc'!19!. '1 181 1.2 ZOOGEOGRÀPHY AND EVOIWION OF PYTHINÀE 186 11 .3 ZOOGEOGRÀPHY OF CT¡ÀDES OF PYTHO 186 1 1 .3.1 !¡!¡-g.d!sË-e!. 11,3.2 P-. gs!-tlElzå-sroup . 187 11.3.3 P. kolsensis- and !. qlgg¡-groups 187 11.3.4 P. leÞressus-group . 190 '1 1.3.5 PleisÈocene events . 191 193 XII. CONCII'DING REI.ÍÀRKS 196 XIII. REFERENCES CITED ÀPPENDIX " ' Page À. IOCÀLITY RECORDS 264 À.1 ryrHg. SEIDEITZI À.2 P!Eg. srRrcns 275 276 A.3 PYTHO KOII{ENSIS 278 À.4 E!IHq NrVÀLrs A.5 PlËq NIGER . 278 290 4.6 BEIH9 ÀBIETICOÌ,À 292 A.7 PYTHO JEZOENSIS À.8 PYIHO PLÀNUS 293 À.9 ElËg. pEPRESSUS 317 T.,IST OF TÀBIJES Table ' Page 1. selected features of samples of seidlitzi Measurements of !. go f rom Ghost Danr ÀLberta and LanieL, ouebäc . '-. l-l-l-ll . .) Measurenents of selected features of sampLes of g. strictus f rom various localities in eastern North Amer ica 90 J.
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