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The Evolution of Genitalia and Mating Behavior in Crickets (Gryllidae) and Other Orthoptera

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The Evolution of Genitalia and Mating Behavior in Crickets (Gryllidae) and Other Orthoptera MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 133 The Evolution of Genitalia and Mating Behavior in Crickets (Gryllidae) and other Orthoptera BY RICHARD D. ALEXANDER AND DANIEL OTTE ANN ARBOR MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN NOVEMBER 17, 1967 MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN The publications of the Museum of Zoology, University of Michigan, consist of two series-the Occasional Papers and the Miscellaneous Publications. Both series were founded by Dr. Bryant Walker, Mr. Bradshaw H. Swales, and Dr. W. W. Newcomb. The Occasional Papers, publication of which was begun in 1913, serve as a medium for original studies based principally upon the collections in the Museum. They are issued separately. When a sufficient number of pages has been printed to make a volume, a title page, table of contents, and an index are supplied to libraries and individuals on the mailing list for the series. The Miscellaneous Publications, which include papers on field and museum tech- niques, monographic studies, and other contributions not within the scope of the Occasional Papers, are published separately. It is not intended that they be grouped into volumes. Each number has a title page and, when necessary, a table of contents. A complete list of publications on Birds, Fishes, Insects, Mammals, Mollusks, and Reptiles and Amphibians is available. Address inquiries to the Director, Museum of Zoology, Ann Arbor, Michigan 48104. No. Directions for collecting and preserving specimens of dragonflies for museum purposes. By E. B. WILLIAMSON.(1916) 15 pp., 3 figs. ......... $0.25 No. An annotated list of the Odonata of Indiana. By E. B. WILLIAMSON. (1917) 12 pp., 1 map ............................................... $0.25 No. Notes on American species of Triacanthagyna and Gynacantha. By E. B. WILLIAMSON.(1923) 67 pp., 7 pls. .................................... $0.75 No. Notes on the genus Erythemis with a description of a new species (Odonata). By E. B. WILLIAA~SON. The phylogeny and the distribution of the genus Erythemis (Odonata). By CLARENCEH. KENNEDY.(1923) 21 pp., 1 pl. ................... out of print No. The genus Perilestes (Odonata). By E. B. WILLIAMSONand J. H. WILLIAM- SON. (1924) 36 pp., 1 pl. ............................................. $1.00 No. A revision of the libelluline genus Perithemis (Odonata). By F. RI~. (1930) 50 pp., 9 pls. ................................................ $0.75 No. The genus Oligoclada (Odonata). By DONALDBORROR. (1931) 42 pp., 7 pls. $0.50 No. A revision of the Puer Group of the North American genus Melanofilus. with remarks on the taxonomic value of the concealed male genitalia in the Cyrtacanthacrinae (Orthopetra, Acrididae). By THEODOREH. HUBBELL. (1932) 64 pp., 3 pls., 1 fig., 1 map ................................... $0.75 No. A review of the dragonflies of the genera Neurocordulia and Platycordulia. By FRANCISBYERS. (1937) 36 pp., 8 pls., 4 maps ....................... $0.50 No. The crane flies (Tipulidae) of the George Reserve, Michigan. By J. SPEED ROGERS(1942) 128 pp., 8 ph., 1 map ................................. $1.25 No. The ecology of the Orthoptera and Dermaptera of the George Reserve, Michigan. By IRVINGJ. CANTRALL.(1943) 182 pp., 10 pls., 2 maps. ....... $1.50 No. Monograph of the family Mordellidae (Coleoptera) of North America, north of Mexico. By EMILLILJEBLAD. (1945) 229 pp., 7 pls. .............. $2.0 (Continued on back cover) R/114CALLLANh0US PIJKLICAI IONS VU41 URI 017 /,001,0GI', tINI\ElISI7 1 01 I\IlC~IlIGAh', NO 133 The Evolution of Genitalia and Mating Behavior in Crickets (Gryllidae) and other Orthoptera ANN 1\Kl3Oli MUSI.:LJI\/I 017 ZOOLOGY, 1TNIVERSITI' 01: RIICHIC; IS NOVEMBER 17, 1967 CONTENTS PAGE INI.II~I)U(:I ION ........................................................... 5 (:~~II'ONI:NI.S01: SI:SIIAL ~ITIIAVIOR........................................ 7 TIII:(;I'NI~I<AI . NA.I.UIIF01: COITII.R.L.IONIN ~~KIIIROI'OOS.......................... 7 (:OI~L.A.I.ION IN OK.~IIOP.~ERA................................................. 8 I'R~VIOUSS.ru111rs ....................................................... 'J (;ROUI'S S~IJI)II~I)........ ....................................... 10 1;a111ily Gryllidac ................ ............................. 10 I'a111ily 'l'ct1igolliitl;te ............................................... 11 1:alnily <;ryllacriclitl;le ............................................. 11 I>ESCRII'.I.IONSOF TV~ATINC B~IIA\.IOI<ANI) (;FNI.IRI.ICSIRUC~URE .................. 12 Subfalrlily (;ryllinae .................................................. 12 (;enus Aclrc. lo ..................................................... 12 (;CIIIIS Gryll~t.~......................... ................. 17 (;enus Gryllories .... ................................... 18 C;cnus Morlic.og~yllr~.s ....... ............................. 19 (;cnrls Teleog~-yllus.............. ......................... lY (;cnr~sVnle?.i/irlo~.~rs ............................................. 20 (;~IIIISiMiogry1lrt.s ................. ........................... 21 I co/ila rylo~o//~z...................... 23 (;t'lIllS G~yllO/l.\i~..................................................... 23 SuI)fa~~~ilyNc~nol~iinac ............ .................... 24 (:enus Nerrrohir~s ........ .............................. 24 S11l>k1111ilyI<r:~c:I~ytr~~l~ir~ac . .................................. 25 <;e~rusATIIIYO~I-~ 1111s ....................................... 25 Sul)f;~trrily(.ryllotalp inxc ... ....................... 29 Genera Ncocrtrtilln ant1 (;ryllo/trl/)cr ............................. 29 Sul~Eur~ilyElrcol,tcl.inac .................. ................ 30 (;clllls tIn/)ilIirrs ................................................. 30 II~or11i .\ I TI............................. 31 Sr~l)fa~lrilyI'li;~l;rtlgopsinac ........................................ 32 (;CII~ISAt~r/~lrictr.rr.\/c~ ....................................... 32 Sul)Ial~~ily0cc:lntllin;rc ..... ...... ..... ............ 33 (;CIIIISo~c/lrllllll.\ ................................... 33 S~tl)fal~~ilyTI-igonidiinac ......... ...... ......... 37 (;(.IICS.I (:v~-/osZf~I~nancl I'lryllo/)ul/~~~.~..... ............. 37 Sul)farl~ily Mogol)lisLin;re ........................ .............. 37 (;(.LIIIS (:yclo/)Iilrt~tr ......................... ....... 37 0tl1cr S~l)kt~nilics............... ....................... 37 SII~IMARYANI) CONCL.USIONS...... .... ........ ....... 38 SIIIIIIIILI~~of I%chll:~vioral \':rriatioiis . ...... ......... 34 I3cl1.1vior. (;cnil;~licStl.uclurc. and Hal)i~at ......................... 41 Rcl~.~vior.(;c~iilalic Structure. arid Plrylogcl~). ........................ 42 IIel~avior.(;cnitalic Structure. and Speciation .................. 44 ACKN~I\JI.I-I)(.RII~N.I.S................................................... 44 I.ITERA.I.UI<E(:I 1.1 ..I) ..... ......... ........ 45 l:IGURl,: PAGE 1. C;cnitali;r and copulatiolr of the house cricket, Arlreln rlo~izc,.stirrr . 50 2. Fe~~l;~lcgenitalia, inale genitalia, ant1 s1jcrl1r:~tol)ho1-e~of Gr)~/lrr.ss1). (Mexico), A1iogr)~llusverticalis, and lTcrlr?-i/rclo)11s rrricctrio . 51 3. Fc~ri;llc genitalia, nr;tle gc~rit:tlia, atrtI S~>~~-III~~~~III~IT~of Trlrogi-yllrts co~ft~rtodrr.~ ant1 Clyllodes sigilln/ri.s . 52 4. I.'c~lr;~lcgenitalia, nralc gct~italia,alicl sl)ci~ila~ol)liorcsof ~l~esuljLcrr;u1cat1 (.rickets, Neoctrrlilla hexctrinctyln ;111(1 A~~ri+ogryII~s~rrtrlicrrs . 53 6. Pre-mating and aggressive bcl~avior in il~t~rrogi-yll~rsritt~ticrca a~rtl(;vylltt.s /ti-rtlris 55 7. 1'l.c-n~ating, post-~~~atitrg,;rntl agg~essivcOel~aviol- ill various crickcls . 5G 8. M;rting behavior in scvcrirl crickets . 57 9. Calling, courtsl~ip,and col)ulalion it1 crickets ;~rrtlgrassl~oppers . 58 10. Courtship and cop~tl;~lioliin various Ensifera . 59 11. Iliagra~~rof tllc probable rclatio~lsl~ipsof 111ajo1-groulx of Ortl~ol)tcr:~ant1 ~>ositiolls assulncd during col~ulatiol~. GO 12. Diagrammatic rep1.escrrtatio11 of 1jrol)able cvolutio~~~u-ycl~ar~ges ill 1)ositio11ilrgof tlle bodies and genitalia of ~nalcand fcn~alcOl.tl1o1jtera cluri~~g~r~a~ilig . 61 INTKODUCTION INSEC:TC;ENITAI.IA have bcen used extensively in taxonomic studies because they arc frequently co~nplicatctlin ritructure and ~tsefulin distinguislling species, ;IS well as in asscssi~lgtheir relationships. Little attention has been lx~itl,however, to the questions of how genitalic tliKerenccs arise, in terms of selecti\ic itction, or wllen, in rcgartl to the speciation prot:ess. The opinion has been expressetl that they might freciuently func.tion as repl-oductive isolating nieclianisms Ijetwecn ncwly separatetl species in the manner of "lock-antl-key" tlevices antl nntlcrgo acceleratetl antl exaggerated divergence in that context (1)uI;oru-, 1844; Dobzhansky, 1951). This hypothesis has never been exxnined tl~oroughly,but its attractiveness has been diminislietl by scatteretl ;tccounts of su(:cessi~lmatings between species with different genitalia (Belleim, 19/12; Sllull, 1946), tliscovery t1i;lt genitalic tlifferences are not nearly univcrs;~l;it the spec.ics level, and realization of the inefiicie~lcy or an isol;~tingrneclla~lism wllich can come into play only after a sexually rcsponsii\lc pair has c.onlc togetllel. ant1 procccdctl tllrougll courtshi11 ant1 the initial 11h;ises of a copulatory act (Dobzllansky, 1951; K. I). Alexander, 1962; Alexantler ant1 Moore, 1962; Ehrnlan, 1961). An un(lcrstan(1ing ol the behavior oC insects during c:opulation, including how they use their genitalia,
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