HARVARD UNIVERSITY LIBRARY OF THE Museum of Comparative Zoology 0, "7 L4 4474 “al a , ene o gs f Pf mage / } / j = y UNIVERSITY OF KANSAS SCIENCE BULLETIN ae pnten gral | MUS. COMP, 7G0L | Lamaifiir | ; TAK 10C 4 } JAN = 4.1961 | ' | JARIFARK AQUARD UNIVERSITY a UNIVERSITY OF KANSAS PUBLICATIONS University of Kansas Science Bulletin - Vol. XLI December 23, 1960 Lawrence, Kansas ANNOUNCEMENT The University of Kansas Science Bulletin (continuation of the Kansas University Quarterly) is issued in parts at irregular inter- vals. Each volume contains from 500 to 1,800 pages of reading matter, with necessary illustrations. Exchanges with other institu- tions and learned societies everywhere are solicited. All exchanges should be addressed to THE UNIVERSITY OF KANSAS SCIENCE BULLETIN, LIBRARY OF THE UNIVERSITY OF KANSAS, LAWRENCE, Kan. PUBLICATION DATES The actual date of publication (i. e., mailing date) of many of the volumes of the University of Kansas Science Bulletin differs so markedly from the dates borne on the covers of the publication or on the covers of the separata that it seems wise to offer a corrected list showing the mailing date. The editor has been unable to verify mailing dates earlier than 1982. Separata were issued at the same time as the whole volume. Vol. Vol. XX—October 1, 19382. XXXII—Nov. 25, 1948. XXI—November 27, 1934 XXXIII, Pt. I—April 20, 1949. XXII—November 15, 1985. Pt. II—March 20, 1950. XXIII—August 15, 1936. XXXIV, Pt. I—Oct. 1, 1951. XXIV—February 16, 1938. Pt. Il—Feb. 15, 1952. XXV—July 10, 19389. XXXV, Pt. I—July 1, 1952. XXVI—November 27, 1940. Pt. II—Sept. 10, 1953. XXVII, Pt. I—Dec. 30, 1941. Pt. III—Nov. 20, 1958. XXVIII, Pt. I—May 15, 1942. XXXVI, Pt. I—June 1, 1954. Pt. II—Nov. 12, 1942. Pt. II—July 15, 1954. XXIX, Pt. I—Tuly 15, 1948. XXXVII, Pt. I—October 15, 1955. Pt. II—Oct. 15, 1943. Pt. II—June 29, 1956. XXX, Pt. I—June 12, 1944. XXXVIII, Pt. I—Dec. 20, 1956. Pt. II—June 15, 1945. Pt. II—March 2, 1958. XXXI, Pt. I—May 1, 1946. XXXIX—November 18, 1958. Pt. II—Nov. 1, 1947. XL—April 20, 1960. Beitr Ss So a Ne Rurus H. THomMpson Editorial Board............4: R. H. THomeson, Chairman CHARLES MICHENER Paut ROoFEe Davin PARETSKY WortTHIE H. Horr ParkE H. Wooparp, Secretary UNIVERSITY OF KANSAS SCIENCE BULLETIN > 180% 1866 DEVOTED TO THE PUBLICATION OF THE RESULTS OF RESEARCH BY MEMBERS OF THE UNIVERSITY OF KANSAS VoLUME XLI UNIVERSITY OF KANSAS PUBLICATIONS LAWRENCE, DECEMBER 23, 1960 PRINTED IN THE STATE PRINTING PLANT TOPEKA, KANSAS 1960 10. MUS. Ceesp 7661 LIBRARY JA : ~4 1961 | VARD GHIVERSITy Contents of Volume XLI Keys to Subfamilies, Tribes, Genera and Subgenera of the Gerridae of the World, Herbert B. Hungerford and Ryuichi Matsuda, Morphology, Evolution and a Classification of the Gerri- dae (Hemiptera-Heteroptera) ...... Ryuichi Matsuda, Serological and Chemical Studies of Gamma-irradiated Ovalbumin, Charles A. Leone and M. Elizabeth Wesley, A Seasonal Survey of the Vertical Movements of Some Zooplankters in Leavenworth County State Lake, Kan- Sashes er ees ee Jerry C. Tash and Kenneth B. Armitage, The Genus Penstemon in New Mexico, Gladys T. Nisbet and R. C. Jackson, The Biology of Nomia (Epinomia) triangulifera with Comparative Notes on Other Species of Nomia, Earle A. Cross and George E. Bohart, A Revision of the Genus Iva L. .......... R. C. Jackson, Lung-Flukes of Snakes, Genera Thamnophis and Coluber, lid, AKATISAS: LOR ya en ee ec tae ae Betty Lan Stewart, A Survey of the Periotic Labyrinth in Some Representa- tive, Necent@hepules’=.) 49.2204 Irwin L. Baird, Observations on the Morphology of the Inner Ear in Certain Gekkonoid Lizards ...... David W. Hamilton, PAGE THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vor, XE] DECEMBER 23, 1960 [No. 1 Keys to subfamilies, tribes, genera and subgenera of the Gerridae of the World* BY HERBERT B. HUNGERFORD and RyvuicHt MATSUDA Apstract: This paper presents keys for the identification of subgenera, genera, and higher categories of the five subfamilies of the waterstrider family Gerridae. The groupings are as follows: Rhagadotarsinae with two genera and four subgenera; Trepobatinae with thirteen genera; Halobatinae including the tribe Halobatini with two genera, and the tribe Metrocorini with seven genera and two subgenera; Ptilomerinae with eight genera and two subgenera; Gerrinae including the tribe Cylindrostethini with three genera, the tribe Charmatometrini with three genera, the tribe Gerrini with ten genera and eight subgenera, and the tribe Eotrechini with four genera. When we began this and the following detailed study by the junior author we had available in the Francis Huntington Snow Entomological collections of the University of Kansas eighty-nine percent of the known genera and subgenera of the Gerridae of the world. The collections also contained the types of many species described by Hungerford and his students (Anderson, Shaw, Kenaga, Kuitert), by Drake and Harris, by Kirkaldy and by Torre- Bueno. There were paratypes of species described by Drake and Harris, Esaki, China, Drake, Hussey, Usinger, Hoberlandt and a number of others. We also had specimens determined by Kiritschenko and Poisson which were most helpful. | Of the genera not represented in our collection we have been able to borrow for study representative species of seven genera through the kindness of Dr. W. E. China and Dr. E. S. Brown of the British Museum, Dr. T. Jaczewski of the Polish Academy of Sciences, Dr. P. Basilevsky of the Royal Museum of Belgian Congo, Dr. A. Collart of the Royal Institute of Natural Sciences of Belgium, Drs. A. Soos and E. Halasfy of the Hungarian National Museum, Dr. R. Poisson of the University of Rennes, France, Dr. S. Miyamoto of the Kyushu University, Japan, and Miss L. C. Chen of the National Taiwan University, to all of whom we are most grateful. Of the three genera we have not seen, the type of one has never been located at any museum ( Rheu- matotrechus himalayanus Kirkaldy ). During our studies of the Gerridae we have published many papers * Contribution number 1,047 from the Department of Entomology, University of Kan- a guns study was made possible with the aid of a grant from the National Science oundation. (3) 4 THe UNIVERSITY SCIENCE BULLETIN describing new genera, subgenera and species, raising subgenera to genera, reducing genera to subgenera or synonymizing genera so that the information could be used in reporting the present study. As we bring them to a close we recognize fifty-three genera and sixteen subgenera in the Gerridae and the possibility of at least another subgenus, a question we cannot settle with- out more material. The keys which are given below are followed by six plates illustrating all but three of the recognized genera. There are sixty-four of these wash- drawings. The figures referred to beyond these sixty-four are to be found in the succeeding paper by the junior author (Morphology, Evolution and a Classification of the Gerridae); certain page numbers also refer to the latter work. These structural drawings are useful in understanding the keys and must be studied before undertaking to use the keys. Key TO SUBFAMILIES 1. Metacetabular suture in wingless forms dorsally continues to posterior margin of mesonotum, forming secondary definitive intersegmental suture between mesonotum and metanotum.* Without anteriorly produced primary intersegmental suture lat- erally in front of metathoracic spiracle (figs. 78-81, 107) 1. Metacetabular suture in wingless forms dorsally not continuous to posterior margin of mesonotum, not forming secondary de- finitive intersegmental suture between mesonotum and meta- notum (exception Cylindrostethini *). Primary intersegmental suture anteriorly produced laterally in front of metathoracic spiracle, or the suture lost laterally or rarely dorsolaterally lost (figs. 72-77, 107) bo 2. First abdominal ventrite present (fig. 96) Rhagadotarsinae Lundblad, p. 319 2. First abdominal ventrite absent (fig. 94)..Trepobatinae Matsuda, p. 330 3. Metacetabular suture dorsally reaching anterior end of first abdominal tergite and intersegmental suture between mesono- tum and metanotum always distinct laterally. Metacetabular region thus always appears to be divided into two areas Gh ioc heay OATS Lem es Patt Sel open ee at ee oe Ptilomerinae Bianchi, p. 260 3. Metacetabular suture not reaching anterior end of first ab- dominal tergite (except for a few genera of Halobatinae). Intersegmental suture either lost laterally or rarely retained lat- erally. Metacetabular region thus never divided into two areas Gee RTC: PTAT (9 Tis Ses BERPER greats nery NO aye io ter gree Oe POR da 4 4. Metasternum clearly present, at least about one tenth as long as mesosternum in length......... Gerrinae Amyot and Serville, p. 160 4. Metasternum represented by a very short transverse, sub- triangular plate rarely reaching metacetabular region laterally, or rarely by omphalium only.............. Halobatinae Bianchi, p. 289 *In Cylindrostethini the metacetabular region is divided into two areas but the metacetabular suture dorsally reaches the intersegmental suture (posterior margin of mesonotum ), not the anterior end of the first abdominal tergite. In the genera of Halo- batinae, in which the metacetabular suture reaches the anterior end of the first abdominal tergite, the intersegmental suture is lost laterally, thus the metacetabular region is not divided into two areas. bo bo to Key TO THE GERRIDAE OF THE WORLD 5 Key To GENERA OF RHAGADOTARSINAE Distal tarsal segment of front leg cleft at tip, with claws arising from base of the cleft. Eighth abdominal segment of both sexes elongate and nearly cylindrical. Males with a longitu- dinal depression on venter of seventh and eighth segments Chet Sine ries ie, arene hae, Rhagadotarsus Breddin 2, p. 322 Distal tarsal segment of front leg not cleft at tip but claws arising preapically from beneath. Eighth abdominal segment of both sexes not elongate as described above (fig. 39) Rheumatobates Bergroth 3, p. 326 Pronotum in wingless forms short, one quarter the length of head. Mesothorax slightly wider than long. Hind coxae visible LromMMAapoOve Ces sl)). 245s. sss R. (Rhagadotarsus) Breddin, p. 326 Pronotum longer, one half the length of head. Mesothorax plainly longer than wide. Hind coxae not visible from above’ (HgV SS) al fae ee Ty ey R. (Caprivia) China, p. 326 Anterior femur of male may be stout but lacks tuft of stout hairs on anterior margin.......... R. (Rheumatobates) Bergroth, p. 329 Anterior femur of male stout, with a tuft of stout hairs on mimicyatore imnentenbay (Git, SYYL)) ao one meoogceo one R. (Hynesia) China, p. 329 Key To GENERA OF TREPOBATINAE Hind femur distinctly longer than length of body. Second and third antennal segments swollen at distal ends and often bearing conspicuous spine clump in male. Hind tarsus with elongate, knife blade shaped claws at or before middle of second tarsal segment i( fig: (57)! 2 es Metrobates Uhler, p. 362 Hind femur as long as or shorter than length of body. Second and third antennal segments without apical modifi- cations mentioned above. Hind tarsus with claws not as GesCcribedsabovewsee tics neo: oars eh on a oe eee eee 2 Intersegmental suture between mesonotum and metanotum Caninatecan loam 49) pees Sere eee Hynesionella Poisson, p. 359 Intersegmental suture between mesonotum and metanotum MOE CATUINATEG @ Mehsud Whe ERC UTR Mees Ren, cree sy Sere eee 3 First antennal segment considerably longer than second and third segments together (fig. 52) Trepobatoides Hungerford and Matsuda, p. 343 First antennal segment at most as long as second and third SECMents tO TetMerys thaws en uee Shae ented hs Pee aaa ee eee nae 4 First middle tarsal segment over twice as long as second middle.tarsallsegient #48. ¢ 05. <2 a Sects ene ae 5 First middle tarsal segment distinctly less than twice as long as second middle: tarsal segment, 4)... 440s eee ne ae 6 Head between eyes greatly widened posteriorly in female. Eyes overlapping less than half of propleuron. Male front tibia strongly arched. (figs. 62, 63) ...... Rheumatometra Kirkaldy, p. 364 Head between eyes subquadrangular, very slightly widened posteriorly in female. Eyes overlapping most of propleuron. Male front femur and tibia not arched (fig. 53) Metrobatopsis Esaki, p. 367 90 90 to bo THE UNIVERSITY SCIENCE BULLETIN Third antennal segment over twice as long as second segment, and distinctly longer than first segment.................... a Third antennal segment less than one and a half times as long as second, distinctly shorter than first................. 8 First antennal segment nearly straight, not reaching beyond eyes. Anterior margins of first and second abdominal tergites completely absent. Metanotum without median longitudinal sulcus. South East Asia (fig. 51)........... Cryptobates Esaki, First antennal segment distinctly curved near base and reach- ing behind eyes. Anterior margin of first and second tergites recognizable laterally. Median longitudinal sulcus of metano- tum normally absent (fig. 59).......... Telmatometra Bergroth, Omphalium#distinetly present. .-..............00..454.65% 9 . Omphalium absent or occasionally vestigial................. 10 Omphalium very conspicuous, located on strongly anteriorly produced anterior margin of metasternum (fig. 58 ) Stenobates Esaki, Omphalium not conspicuous, anterior margin of metasternum not strongly produced anteriorly (fig. 56) Rheumatometroides Hungerford and Matsuda, Mesopleura without distinct longitudinal stripe (see fig. 61) Naboandelus Distant, Mesopleura with distinct black, or yellow, or brown longi- tudinal stripe eer inc Men. Loner ta ee VE eel a tal Middle tibia distinctly shorter than length of body (fig. 55) Ovatametra Kenaga, Middle tibia about as long as or a little longer than length of body.) 3s SS een eee ee, Meee See cD OP BG Nec rgiee 12 Eyes not extending beyond middle of propleura in side view. Hind tibia distinctly less than twice as long as tarsus (fig. 60) Trepobates Uhler, Eyes extending beyond anterior half of propleura. Hind tibia over twice as long as tarsus (fig. 50) ....Halobatopsis Bianchi, Key To Tripes, GENERA AND SUBGENERA OF HALOBATINAE Clypeus with basal margin well defined, anterior margin of head not smoothly rounded. Marine in habitat Halobatini Bianchi 2, Clypeus with basal margin obliterated or lost. Anterior mar- gin of head broadly and smoothly rounded. Fresh water in habitat fie oof cay a cee eee Metrocorini Matsuda _ 3, Long hairs confined to middle tibia (fig. 40). . Asclepios Distant, Long hairs present on tibia and first tarsal segment of middle leg Gigs AU 3.8 coe, eee eee Halobates Eschscholtz, Metanotum with lateral longitudinal elevation * reaching inter- segmental suture between mesonotum and metanotum. Male third antennal segment has stiff hairs on margins (fig. 44) Esakia Lundblad, * Appears to be forward continuation of the abdominal connexivum. 340 . 348 337 . 346 294 . 301 297 . 299 . 316 oS LSS) fs Kry TO THE GERRIDAE OF THE WORLD Metanotum with lateral longitudinal elevation not reaching intersegmental suture between mesonotum and metanotum. Male third antennal segment without stiff marginal hairs... .. . 4 Eyes overlapping anterolateral angles of mesonotum......... 5 Eyes not overlapping anterolateral angles of mesonotum..... . 6 Posterolateral angles of metacetabula simple. Male without a tubercle either on mesosternum or on inner margin of front LEMUR ig CA I. Ae PS: Ventidius (Ventidius) Distant, Posterolateral angles of metacetabula bilobed. Male with a tubercle on mesosternum and on inner margin of front femur (fig 243%. vee Ventidius (Ventidioides) Hungerford and Matsuda, Metasternum represented practically by a small omphalial pore Gi Sl) Rae Ne ee a Shee: Eurymetropsiella Poisson, Metasternum represented either by a transverse subtriangular lobe, or posteriorly produced conical plate bearing omphalium, 7 Metasternum represented by a conical plate produced pos- teriorlyaGhiee 49). Sates DY. Bea eee Eurymetropsielloides Poisson, Metasternum represented by a transverse subtriangular plate, its anterior margin more or less strongly produced anteriorly.. 8 Mesonotal region with a median longitudinal and lateral paired Obluquenblackstripese <. 0. toca a soe oN eee ne 9 Mesonotal region predominantly black, without black longi- tudinal stripes. Male pygophore bifurcates apically (fig. 46) Eurymetropsis Poisson, Body flattened and often lustrous above. Female seventh abdominal segment with ventral apical margin simply con- CAVE nf tt) a ley. RCRA SiUt Letra Nepean Eurymetra Esaki, Body flattened and dull in color above. Female seventh abdominal segment with ventral apical margin excessively developed and modified in various shapes, hiding eighth seg- MEN GAA 74 oe Peet) een tyiet dora Me URE Rae, Metrocoris Mayr, KEY TO GENERA AND SUBGENERA OF PTILOMERINAE Hind tarsal’ segments fused: 7. 505 te ee ») Hind tarsal segments distinct from each other .............. 4 Lateral longitudinal suture of mesonotum distinct. Female abdomen withdrawn into thoracic cavity (fig. 28 ) Potamometra Bianchi, Lateral longitudinal suture of mesonotum absent. Female abdomen is not withdrawn into thoracic cavity ........... 3 Hind coxa without a spine. Seventh segment of female without lateral lobes (fig. 33) Ptilomera (Proptilomera) Hungerford and Matsuda, Hind coxa with a spine. Seventh segment of female with conspicuous lateral lobes (fig. 36) Ptilomera (Ptilomera) Amyot and Serville, Anterior margin of head rounded. First antennal segment shorter than three following segments together (fig. 32) Rheumatogonus Kirkaldy, 313 315 . 307 . 309 311 305 302 271 270 270 283 8 THe UNIVERSITY SCIENCE BULLETIN 4. Anterior margin of head not rounded. First antennal segment about as long as or longer than three following segments to- Pethler” oan os ee Tee ee se Se patien § erten hie Ta 5) 5. First hind tarsal segment shorter than second. Female seventh abdominal segment without narrow, long and spinous process. Metanotum in female without median elevated process on Mini SIMA OT Tes 1 ees ME CNRS OIE EE Pe Ss a 6 5. First hind tarsal segment twice as long as second. Female seventh abdominal segment with narrow, long and spinous process. Metanotum in female with median elevated process on, ‘hindkamargin, hacia ec foe oe Pleciobates Esaki, p. 286 6. Middle and hind tarsi with distinct claws (fig. 30) Potamometropsis Lundblad, p. 281 Middle and hind tarsi without distinct claws ............... 7 Hind coxa long and cylindrical, twice as long as basal width. Female abdomen telescoped into thoracic cavity and_ its metacetabula with a fingerlike process on inner rear margin (Gaokerspeeey © Otay) Wee eet cok eee? Potamometroides Hungerford, p. 278 7. Hind coxa shorter, not cylindrical, basally broader. Female abdomen not telescoped into thoracic cavity, its metacetabula without, 7a) projecting sprocess: «406 1 .2cet a. se ae 8 8. Front femur without or with one or two indefinite dorso- lateral bands. Caudal margin of pronotum straight or con- cave. Mesothorax with sides converging cephalad and antero- lateral angles sloping, not prominent. Female venter normal (fig. D9) mes’. pheele ceeeias sesh utes a Rhyacobates Esaki, p. 273 8. Front femur with two dorsolateral longitudinal black bands. Caudal margin of pronotum faintly undulate, its median lobe slightly produced caudally. Mesothorax with sides nearly parallel and anterolateral angles prominent and _ transverse. Female venter with well-demarcated flattened area (fig. 31) Heterobates Bianchi, p. 276 SS Key to TRIBES OF GERRINAE 1. Metacetabular suture connected dorsally with dorsal posterior margin of mesonctum (intersegmental suture ) Cylindrostethini Matsuda, p. 217 1. Metacetabular suture not connected with intersegmental suture dorsally: ted \.08 nell Vas at A i se gee mR NE ana 2 2. Anterior margin of first abdominal tergite straight Charmatometrini Matsuda, p. 233 2. Anterior margin of first abdominal tergite bisinuate (flattened Weshapéd)) ie rt: ea a eee ee er... ene 3 3*, Pronotum prolonged primitively. Connexival spine present primitively. Apical segment of endosoma always provided Wwithaventral plates...) seen Gerrini Amyot and Serville, p. 163 * These two characters indicate fundamental differences in evolutionary trends. For separation of the genera of Gerrini and Eotrechini, see the key to genera of Gerrinae. Kry TO THE GERRIDAE OF THE WORLD 3*. Pronotum not prolonged. Connexival spine absent primi- tively. Apical segment of endosoma with very poorly de- veloped ventral plate or without it, and apical plate greatly developed iy citeet eA Ae een ee ite erase Eotrechini Matsuda, p. Kry To GENERA OF GERRINAE + 1. Hind leg longer than middle leg.....:....:-:....---.-... 2 1. Hind leg nearly as long as or shorter than middle leg........ . 3 2. Metasternum with omphalial groove present. Claws arising preapically. Gigantic in size (fig. 3) Gigantometra Hungerford and Matsuda, p. 2. Metasternum without omphalial groove. Claws arising apically. Moderatedinesizes( fig. 2o))* sac ames. 28 nee oe Eotrechus Kirkaldy, p. 3. Metacetabular suture reaching dorsally to intersegmental suture between mesonotum and metanotum....... ......... 4 3. Metacetabular suture not reaching dorsally to intersegmental suture between mesonotum and metanotum................ 6 4. Body strongly flattened and short, without omphalial groove (EDO) ao Pacts ce See 3. amen: Platygerris B. White, p. 4. Body not flattened, cylindrical or at least not short. With OmphalialeenoOVvers aN a a.): Pee ee se, ok repens eae 5 5. Abdominal spiracles located closer to anterior margins than to posterior margins of segments.** Male pygophore not rotated. Body more or less cylindrical (figs. 17, 18) Cylindrostethus Fieber, p. 5. Abdominal spiracles located at middles of segments.* Male pygophore rotated. Body shorter (fig. 19) Potamobates Champion, p. 6G Omphatialteroove spresents;- sag ser tt. Ace cee oe ee ri 63 sOmphalialveroovetabsent..)5. Ut 40...) ere es eee 10 7. Pronotum relatively short. Mesosternum about twice as long as metasternum (fig. 9). .Gerriselloides Hungerford and Matsuda, p. 7. Pronotum long. Mesosternum at least five times as long as IME tASLELTAUITTN gi tates SAE ee ona PWR Ary Sag POM PRA ye aA 8 8. Middle femur longer than middle tibia..............5.2..... 9 8. Middle femur shorter than middle tibia (fig. 23) Brachymetra Mayr, p. 9. Pronotum with four black longitudinal stripes, marginal ones confluent posteriorly. First tarsal segment of front leg shorter thansecomen (toe 22)) ee ee ere esata Eobates Drake and Harris, p 9. Pronotum without black longitudinal stripes, concolorous brown. First tarsal segment of front leg longer than second (fie (OR) e ye ee alg no Charmatometra Kirkaldy, p. 10. First tarsal segment of middle and hind legs shorter than second. Claws arising from near middle of second segment ang conspicuous (fig26))--.. 4565.2: os Onychotrechus Kirkaldy, p. 243 Al 249 231 224 228 187 240 242 238 251 + Rheumatotrechus Kirkaldy is not included. ** In Potamobates thomasi Hungerford the spiracle is placed closer to the anterior margin than to the posterior margin of each segment. 10 10. te 17: 18. 18. Tue UNIVERSITY SCIENCE BULLETIN First tarsal segment of middle and hind legs longer than second. Claws inconspicuous and arising from near apex of Second sepments ic. Ure ee oe be bes ee oe tders detach 11 Mesonotum with paired oblique depressions near anterior mar- gin. Paramere greatly developed (fig. 27) Chimarrhometra Distant, Mesonotum without paired oblique depression near anterior TUVALU 2s. xo ee MRICS ES PCat: Bane OPA or th yteite-ae 12 Hind coxa distinctly longer than wide. Pronotum not pro- longed! (fig: 24) ccamcioaa sant oh ckam cay eae ie Amemboa Esaki, Hind coxa shorter than wide or as wide as long. Pronotum pro- longed! impmiost species mete = tee toe ares tian het arse 13 Hind tibia less than one fourth as long as hind femur. Male suranal plate with conspicuous spinous process on each side (Goce HOMIES) bot yn. 2 eso ake et ae eee Gerrisella Poisson, Hind tibia over one-third as long as hind femur. Male suranal plate without conspicuous spinous process................-; 14 Pronotum with a median black longitudinal stripe........... 15 Pronotum with a median yellow longitudinal stripe.......... 18 Pronotum not prolonged (fig. 16) Tenagogonus (Tenagometra) Poisson, Pronotuna jroloncedwes taerrar et. cents 4 ee scr ae 16 Rostrum with third segment not reaching onto mesosternum (Gita OW) he eenene ene ater Tenagogerris Hungerford and Matsuda, Rostrum with third segment reaching onto mesosternum...... 17 Male abdomen short, 7th, 8th and 9th segments together at least as long as four preceding segments. Male without con- nexival segment produced into triangular flattened plate or NexiVal spe. 44s ieee Tenagogonus (Tenagogonus) Stal, Male abdomen not reduced, 7th, 8th and 9th segments together shorter than four preceding segments. Male 7th connexival segment produced into triangular flattened plate or spinelike process 4(dig.ch4)" Jessie aor Tenagogonus (Limnometra) Mayr, Mesopleuron with two large white spots. Legs and antennae in male about three times longer than in female (fig. 1) Tenagometrella Poisson, Mesopleuron without two large white spots. Legs and an- tennae nearly equal in length in both sexes................. 19 Second antennal segment as long as or longer than third, or a littlesshortensthamuthirdesteeteet: <.- fois ena ecw che ote 20 Second antennal segment much shorter than third (fig. 2) Tachygerris Drake, Pronotum shiny in most species. Pronotum either with a median longitudinal yellow stripe and lateral short yellow stripes, or the lateral stripes alone, or with a large yellow spot alone on anterior lobe........-.-.----+- +--+ +0022 00: 21 Pronotum dull, always with a median yellow longitudinal stripe which is obliterated on posterior lobe and often with a pair of large black spots on either side of median yellow longitudinal stripe - 24,.9e20 sod. oc oor ee 22 p. 254 256 189 213 209 - 214 202 Key TO THE GERRIDAE OF THE WoRLD 15 Pronotum with median yellow longitudinal stripe reaching posterior margin of pronotum and always with a pair of short concolorous lateral stripes one on either side of median longi- tudinalcstripes (igsa)).5. 5-900: Limnogonus (Limnogonus) Stal, p. 200 Pronotum with a large median yellow spot on anterior lobe (fig. 8)... Limnogonus (Limnogonellus) Hungerford and Matsuda, p. 200 First antennal segment considerably shorter than second and (ovina! GSAT WOYUNE, aw acaco dg auveecooodccoenusowuse 23 First antennal segment longer than or equal to or slightly shorter than second and third segments together............. 24 Relatively broad and short species. Hind femur about as long as middle femur. Metathoracic spiracle placed more than its own length from pronotum. Pronotum not fully prolonged (fig) Ree nn ee ee Eurygerris Hungerford and Matsuda,* p. 194 More slender and more elongate species. Hind femur distinctly longer than middle femur. Metathoracic spiracle placed less than its own length from pronotum. Pronotum fully pro- longed/a( fig. 14)" pega ees. ee Gerris (Limnoporus) Stal, p. 184 Hind tibia about three times (never over 3.2 times) as long as first tarsal segment. First antennal segment about equal to or a little shorter than two following segments together * (fig. 13) Gerris (Gerris) Fabricius, p. 179 Hind tibia at least four times as long as hind first tarsal seg- ment. First antennal segment about equal to or a little longer than two following segments together ** (fig. 15) Gerris (Aquarius) Schellenberg, p. 175 *In E. mexicanus (Champion) the posterior lobe of pronotum is almost fully pro- longed. ** The antennal character is not satisfactory for determining New World Aquarius. to 10. JAI Tue UNIverRsiItry SCIENCE BULLETIN Ficures 1-12 Tenagometrella longicornis (Poisson), female. Length of body: 12.4 mm. Tachygerris quadrilineatus (Champion), male. Length of body: 6.85 mm. Gigantometra gigas (China), male. Length of body: 31.9 mm. Tenagogonus (Limnometra) femoratus (Mayr) male. Length of body: 18.0 mm. Eurygerris fuscinervis (Berg), male. Length of body: 6.4 mm. Tenagogonus (Tenagogonus) albovittatus Stal, male. Length of body: 6.55 mm. Limnogonus (Limnogonus) hyalinus (Fabricius), male. Length of body: 8.53 mm. Limnogonus (Limnogonellus) hesione (Kirkaldy ), female. Length of body: 6.15 mm. Gerriselloides brachynotus (Horvath), female. Length of body: 7.6 mm. Gerrisella settembrinoi (Poisson), winged male. Length of body: 5.2 mm. Gerrisella settembrinoi (Poisson), wingless male. Length of body: 4.5 mm. Tenagogerris euphrosyne (Kirkaldy ), wingless female. Length of body: 6.4 mm. KEY TO THE GERRIDAE OF THE WORLD 13 Ficures 1-12 2 Tachygerris quadrilineatus | Tenagometrella longicornis vey 4 Tenagogonus (_imnometra) femoratus 6 Tenagogonus(Tenagogonus) 8 Limrogonus ( ee 7 Limnogonus(Limnogonus) hesione hyalinus 5 Eurygerris fuscinervis I] Gerrisetla settembrinoi 12 Tenagogerris euphrosyne 9 Gerriselloides brachynotus IO Gerrisella settembrinoi 14 Tue UNIVERSITY SCIENCE BULLETIN Ficures 13-24 Gerris (Gerris) thoracicus Schummel, winged female. Length of body: 11.4 mm. Gerris (Limnoporus) rufoscutellatus (Latreille), winged male. Length of body: 13.6 mm. Gerris (Aquarius) paludum Fabricius, winged male. Length of body: 15.1 mm. Tenagogonus (Tenagometra) sp., wingless female. Length of body: 6.1 mm. Cylinorostethus palmaris Drake and Harris, wingless male. Length of body: 16.2 mm. Cylindrostethus productus Spinola, winged female. Length of body: 27.0 mm. Potamobates unidentatus Champion, wingless male. Length of body: 8.7 mm. Platygerris depressus B.-White, wingless male. Length of body: 5.8 mm. Charmatomeira bakeri Kirkaldy, wingless female. Length of body: 13.5 mm. Eobates vittatus (Shaw), wingless male. Length of body: 7.5 mm. Brachymetra kleopatra Kirkaldy, wingless male. Length of body: 8.0 mm. Amemboa fumi Esaki, wingless female. Length of body: 4.1 mm. Key TO THE GERRIDAE OF THE WORLD 1155 Ficures 13-24 \ j MM x! Hie: 15 Gerris(Aquarius) A - s(Limnoporus) 13 Gerris (Gerris) thoracicus 14 Co aaah poludum 18 Cylindrostethus productus : 20 Platygerris depressus I6 Tenagogonus (Tenagometra) sp, |9 Potamobates unidentatus = 2 3 Brachymetra kleopatra 24 Amemboa fumi 22 Eobotes vittatus 16 34. 35. Tue UNIVERSITY SCIENCE BULLETIN Ficures 25-35 Eotrechus kalidasa Kirkaldy, winged male. Length of body: 10.2 mm. Onychotrechus sakuntala, Kirkaldy, wingless female. Length of body: 6.5 mm. Chimarrhometra orientalis (Distant), wingless male. Length of body: 7.2 mm. Potamometra berezowskii Bianchi, wingless female. Length of body: 16.2 mm. Rhyacobates takahashii Esaki, wingless female. Length of body: 9.2 mm. Potamometropsis werneri Hungerford, wingless female. Length of body: 8.2 mm. Heterobates dohrandti Bianchi, wingless male. Length of body: 7.1 mm. Rheumatogonus burmanus, Distant, wingless female. Length of body: 6.8 mm. Ptilomera (Proptilomera) himalayensis Hungerford and Matsuda, winged male. Length of body: 10.4 mm. Potamometroides madagascariensis Hungerford, wingless male. Length of body: 6.5 mm. Potamometroides madagascariensis Hungerford, wingless female. Length of body: 5.8 mm. (as for coxa) KrEY TO THE GERRIDAE OF THE WORLD 17 Ficures 25-35 25 Eotrechus kalidasa 26 Onychotrechus sakuntala 29 Rhyocobates takahashii 30 Potamometropsis werneri : ; 3| Heterobates dohrandti 35 Potamometroides 33 Ptilomera(Proptilomera) 34 Potamometroides madagascariensis 32 Rheumatogonus himalayensis madagascariensis burmanus 18 44, 45. Tue UNIVERSITY SCIENCE BULLETIN Ficures 36-45 Ptilomera (Ptilomera) werneri Hungerford and Matsuda, winged male. Length of body: 11.0 mm. Rhagadotarsus (Rhagadotarsus) kraepelini Breddin, wingless male. Length of body: 3.1 mm. Rhagadotarsus (Caprivia) hutchinsoni China, wingless female. Length of body: 5.4 mm. Rheumatobates rileyi Bergroth, wingless female. Length of body: 2.6 mm. Aschepios apicalis Esaki, wingless male. Length of body: 2.6 mm. Halobates sobrinus B.-White, wingless male. Length of body: 4.0 mm. Ventidius (Ventidius) malayensis Hungerford and Matsuda, wingless male. Length of body: 3.9 mm. Ventidius (Ventidioidis) kuiterti Hungerford and Matsuda, wingless male. * Length of body: 2.4 mm. Esakia kuiterti Hungerford and Matsuda, wingless female. Length of body: 2.2 mm. Metrocoris strangulator Breddin, wingless male. Length of body: 5.05 mm. Key TO THE GERRIDAE OF THE WORLD 19 Ficures 36-45 39 Rheumatobates rileyi 37 Rhagadotarsus (Rhagadotorsus) kraepelini 38 Rhagadotarsus ( Caprivia) Ptilomera(Ptilomera) 36 wernerl hutchinson 40 Asclepios apicalis idioi Esokio kuiterti 43 Net DoCctlle As au 45 Metrocoris strangulator bo S 46. 47. 53. 54. THE UNIVERSITY SCIENCE BULLETIN Ficures 46-54 Eurymetropsis carayoni Poisson, wingless male. Length of body: 5.2 mm. Eurymetra natalensis (Distant), wingless female. Length of body: 4.9 mm. Eurymetropsiella schoutedeni Poisson, wingless female. Length of body: 3.8 mm. Eurymetropsielloides milloti Poisson, wingless male. Length of body: 3.5 mm. Halobatopsis platensis (Berg), wingless male. Length of body: 3.5 mm. Cryptobates raja (Distant), wingless male, genital segments removed. Length of body: 3.2 mm. Trepobatoides boliviensis Hungerford and Matsuda. Length of body: 3.8 mm. Metrobatopsis flavonotatus Esaki, wingless female. Length of body: 2.4 mm. Hynesionella omercooperi Hungerford and Matsuda. Length of body: 2.35 mm. Key TO THE GERRIDAE OF THE WORLD OA Ficures 46-54 Eurymetra natalensis = ; : a 48 Eurymetropsiella schoutedeni Say 52 Trepobatoides boliviensis 53 Metrobatopsis flavonotatus 5 4 Hynesionella omercooperi bo bo 63. 64. THE UNIVERSITY SCIENCE BULLETIN FicuRES 55-64 Ovatametra minima Kenaga, wingless male. Length of body: 2.1 mm. Rheumatometroides browni Hungerford and Matsuda. Length of body: 3.2 mm. Metrobates hesperius Uhler, wingless male. Length of body: 4.1 mm. Stenobates biroi (Esaki), wingless male. Length of body: 4.05 mm. Telmatometra whitei Bergroth, wingless female. Length of body: 4.2 mm. Trepobates pictus (Herrich-Schaeffer ), wingless male. Length of body: 3.3 mm. Naboandelus bergevini Bergroth, wingless female. Length of body: 2.55 mm. Rheumatometra philarete Kirkaldy, wingless female. Length of body: 3.15 mm. Rheumatometra philarete Kirkaldy, wingless male. Length of body: 2.3 mm. Hermatobates weddi China, wingless male. Length of body: 3.6 mm. Kry TO THE GERRIDAE OF THE WORLD 23 Ficures 55-64 57 Metrobates hesperius 5G Siewoates: bits 61 Naboandelus bergevini 59 Telmatometra whitei 63 Rheumatometra philarete 64 Hermatobates weddi 62 Rheumatometra philarete i _t. oe ein. 4 SAKE avivaiT THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Voi. XLI! DECEMBER 23, 1960 [No. 2 Morphology, Evolution and a Classification of the Gerridae (Hemiptera-Heteroptera ) * BY Ryvuicut MatTsupA The University of Kansas Apstract: This work is a study of the morphology, evolution and classifica- tion of the Gerridae of the World. Fifty genera and sixteen subgenera out of fifty-three genera and sixteen subgenera known were examined. In the section on morphology it is attempted to establish homologies and a terminology for as many external structures as possible. In the section on the structural evolution the process of evolutionary change of each structure is traced, and its taxonomic significance is discussed. The postembryonic de- velopment of the antennal and leg segments has been studied in representative species of each major group to see how the different proportional lengths of antennal and leg segments are realized ontogenetically; how the ontogenetic growth patterns for these segments have been carried over to adult phylogeny; and how the growth patterns themselves have evolved. It was found that (1) the antennal and leg segments show roughly a simple allometric growth, with either an appreciable increase or decrease in growth ratio at the final stage of development; (2) often lengths of the leg and antennal segments of adults in a great majority of species within a genus fall roughly on the growth lines for the corresponding segments in a representative species of the same genus, indicating that species within a genus share very similar growth patterns for corresponding segments; (3) a hypothetically primitive growth ratio for the antennal segments (k= 1.142) is suggested; (4) a process of development of the proximo-distal gradient in growth ratios for the antennal segments in the phylogeny of the Gerridae is traced; (5) for certain segments, such as the hind tibia, there is evidence in many genera that the growth patterns vary among species of a genus, thus forming the secondary phylogenetic allomorphic lines; (6) as a result of the formation of the secondary allomorphic slope for the hind tibia, which is always steeper than that for the hind femur, the tibia is shorter in relation to the femur in the smaller species of a given genus, and _ this tendency occurs in most major groups of the Gerridae; (7) since there is a striking tendency toward smaller body size in the structurally more specialized forms at all taxonomic levels, and the congeneric species often appear to have * The contribution number 1048 from the Department of Entomology, The University of Kansas. This study was made possible with the aid of a grant from the National Science Foundation. (25) 26 Tue UNIVERSITY SCIENCE BULLETIN very similar allometric growth patterns for corresponding antennal and leg segments, the lengths of antennal and leg segments in the early postembryonic developmental stages in larger and primitive species roughly approximate the lengths of the same in adults of related but phylogenetically more advanced forms. In the light of the knowledge gained from the study of evolution of the leg and antennal segments, the taxonomic status of all groups of the Gerridae (subfamilies, tribes, genera and subgenera) is evaluated. The arrangement of genera in the proposed new classification of the Gerridae follows. Trepobatinae is described as a new subfamily and Hermatobatinae is excluded from the Gerridae. (1) Gerrinae. Gerrini, including Gerris (Gerris s. str., Aquarius, Limnoporus), Gerriselloides, Gerrisella, Gigantometra, Tenagogerris, Eurygerris, Limnogonus (Limnogonus s. str., Limnogonellus), Tachygerris, Tenagogonus (Tenagogonus s. str., Limno- metra, Tenagometra), Tenagometrella. Cylindrostethini, including Cylindrostethus, Potamobates, Platygerris. Charmatometrini, including Charmatometra, Brachymetra, Eobates. Eotrechini, including Eotrechus, Onychotrechus, Chimarrhometra, Amemboa, Rheumatotrechus(?). (2) Ptilomerinae, including Ptilomera (Ptilomera s. str., Proptilomera), Pota- mometra, Rhyacobates, Heterobates, Potamometroides, Potamometropsis, Rheu- matogonus, Pleciobates. (3) Halobatinae. Halobatini, including Asclepios, Halobates. Metrocorini, including Metrocoris, Eurymetra, Eurymetropsiella, Eurymetro- psielloides, Eurymetropsis, Ventidius (Ventidius s. str., Ventidioides), Esakia. (4) Rhagadotarsinae, including Rhagadotarsus (Rhagadotarsus s. str., Ca- privia), Rheumatobates (Rheumatobates s. str., Hynesia). f (5) Trepobatinae, including Trepobates, Telmatometra, Trepobatoides, Halobatopsis, Ovatametra, Rheumatometroides, Stenobates, Cryptobates, Na- boandelus, Hynesionella, Metrobates, Rheumatometra, Metrobatopsis. CONTENTS PAGE INBSTIRA CTS Mice ca cytes eee RAD ER Roe ERE CR ERTL caesecLe ea her oie Ct mR er es 25 TINTRODUCTION go's, ocheneshocdeb ee Re oe oe Ee REE oe 31 EXTERNAL. MORPHOLOGY: saditsaiie ako Oe each Spot aon > OE exniea cil 33 Mee Hea Gace = Hai ARR a Le ce ee eS 33 (hey thorax bie siweks: Bes MOD. ers Oh RSA ae Seen te Set 34 Dhetprothorax: 225g ae. AAU BEE PRS eee 35 The mesothoraxand: wingibases.- she 2h Aaa ee es ae 35 Rhewitetathoraxws Law. SEP Pee ee ee ee ee 37 "Thesabdomenty ts -s: 8): eek ae. ee ee ee eerie 39 Phe, ,pregénital .segments tak. Aen eee eee eee ee 39 lw Dhesfrst abdominalasegmentine Bisse eee se 40 2. The second to seventh abdominal segments ............. 41 The genital: segmentsiziaien ese eee ee ee ee 41 1. Origin of the male external genitalia in Hemiptera ...... 41 SruDY OF THE GERRIDAE OF THE WORLD oF PAGE 2. The male external genitalia of the Gerridae ............. 43 3. Origin of the female external genitalia in Hemiptera ...... 45 4. The structural plan of the female external genitalia in Hemiptera, with special reference to Gerridae......... 45 ELVOLURIONHORS SERUCTURES! (te Seen ttre: Sethe eh. cen ae Aen Ar eaem oie 47 I. EvoLurioN OF THE STRUCTURES OTHER THAN THE LEGS AND ANTENNAE Wop yates cae cata eR ono ce ee each TORE ee 47 the sshapesofthe: bodye2c, 4. os Ghia @ Sho Se Ve eee ee eee AT HE @heac tier Fe het eT on Pee, Meee ono Mt ee Cue ts Sylar eae ee 51 Rhershaper of inesncad we ketscl kwh omelet. mare Sane 51 DRS CUCS BRE RUN NEE A i AO ad cht eo Et Ais oa 52 REM CLI CUS LIA RE Eine ins cee Rie nae lore eg Mee oe 53 The mandibular and maxillary plates and the labrum........ 53 DRC PTOSENITIVag ae ses ye cae de e went tt ea 54 Pee PLOL Or axes ise oie, au. site Aatencr ohs ars tare sate Oa sd cet te nn ee 54 TREY DON OLUTNIE: Brae mien, ors ae ee Ee aE EO Te 54 he amesothoraxce, 2ee:) 21) Ae ce COMER n Wek thon nce 56 The lateral longitudinal suture of the mesonotum........... 56 The median longitudinal sulcus of the mesonotum........... 56 CRIME SOSLETIIUIUN Staats oe AN eae ers Cee a EE 57 The intersegmental suture between the mesonotum and MELAN OTL cl eet toa a ar eee 57 he pametathoraxy isles aeons Sth Oe AAR eee 58 Mheemetathoracicespinacle 2) Wo ee ee 58 The median longitudinal sulcus........5.5....50.01.<02:5- 59 The lateral longitudinal suture of the metanotum............ 59 Ne sNetasternunn a5) 1 PRA aay eae Beis): epee 59 Bhewomphaliinve: os Oh fe OA Ee Ee 61 Thesfirst;abdominalesegment. ......-.5...6-. 0. dee ee 61 ithe frstiabdominal: tenvites 4 42e8e snes eee 61 The connexival part of the first abdominal segment ........ 62 The first abdominal ventrite..............0200 0c e ceases 62 The second to sixth abdominal segments...................... 63 The second to sixth abdominal tergites and ventrites........ . 63 The location of the abdominal spiracles................... 64 The ventral longitudinal suture of the connexivum.......... 65 Thexseventh abdominal ‘seoments ../. 6s q5024. noo catsoee eee oe 66 dhereighthvabdominalisegment... ..... 5.045 5a eee ee 70 The ninth abdominal segment 72 The ninth tergite (suranal plate) in the male............... 72 Te