^ ' /^ /f -L HARVARD UNIVERSITY LIBRARY OF THE Museum of Comparative Zoology / / - 'T J BULLETIN of The University of Kansas SCIENCE BULLETIN (Continuation of Kansas University Qaarterly) Vol. XXIV LAWRENCE, KANSAS Published Semimonthly Vol. 37 JULY 15, 1936 No. 14 Entered as second-class matter Decemb'er 29, 1910, at the post office at Lawrence, Kansas, under act of July 16, 1894 16-6037 NOTICE TO EXCHANGES The attention of learned societies and other institutions which exchange scientific piibliaations with the University of Kansas is called to the list of publications of this University on the third and fourth pages of the cover of this issue. Those marked "Supply exhausted" cannot be furnished at all; as far as the supply permits the remaining numbers will be furnished gladly to any of our exchanges who may need them to complete their files. Back numbers of the Kansas University Quarterly, as far as pos- sible, will be sent to those of our newer correspondents who are able and willing to reciprocate. Separates are available to specialists. ANNOUNCEMENT The Kansas University Science Bulletin (continuation of the Kansas University Quarterly) is issued in parts at irregular inter- vals. Each volume contains from 300 to 400 pages of reading mat- ter, with necessary illustrations. Exchanges with other institutions and learned societies everywhere are solicited. All exchanges should be addressed to the Library of the University of Kansas. The Kansas University Science Bulletin, Library of the University of Kansas, Lawrence, Kan. EDITORIAL BOARD H. B. HuNGERFORD, Chairman. A. W. Davidson. E. H. Taylor, Secretary. C. M. Baker. J. D. Stranathan. O. 0. Stoland. R. C. Moore. THE KANSAS UNIVERSITY Science Bulletin •DIVERSITY OF KANSAS SCIENCE BULLETIN First Mailing of Volume XXIII — August 15, 1936 First Mailing OF Volume XXIV — February 16, 1938 Vol. XXIV (Whole Series, Vol. 34) published by the "UNIVERSITY LAWRENCE, KANSAS 1936 ^nUlY I'll': c-ara-.^oofor.Y, PRINTED BY KANSAS STATE PRINTING PLANT W. C. AUSTIN. STATE PRINTER TOPEKA 1937 16-6037 NOTICE TO EXCHANGES The attention of learned societies and other institutions which exchange scientific publiaations with the University of Kansas is called to the list "— *-^'"~^~~~--^-^i^oJJnivp.iisit5? on the third and fourth pages of Those marked far as the suppl gladly to any of files. Back number,' sible, will be sei and willing to The Kansas University Science Bulletin (continuation oi tne Kansas University Quarterly) is issued in parts at irregular inter- vals. Each volume contains from 300 to 400 pages of reading mat- ter, with necessary illustrations. Exchanges with other institutions and learned societies everywhere are solicited. All exchanges should be addressed to the Library of the University of Kansas. The Kansas University Science Bulletin, Library of the University of Kansas, Lawrence, Kan. EDITORIAL BOARD H. B. HuNGERPORD, Chairman. A. W. Davidson. E. H. Taylor, Secretary. C. M. Baker. J. D. Stranathan. O. 0. Stoland. R. C. Moore. THE kainsas university Science Bulletin DEVOTED TO THE PUBLICATION OF THE RESULTS OF RESEARCH BY MEMBERS OF THE UNIVERSITY OF KANSAS Vol. XXIV (Whole Series, Vol. 34) />L) th^^^ iljtc( jr/lQ/3 r PUBLISHED BY THE UNIVERSITY LAWRENCE, KANSAS 1936 PRINTED BY KANSAS STATE PRINTING PLANT W. C. AUSTIN. STATE Printer TOPEKA 1937 16-6037 R'l FEB 1 8 1938 I a 'j-j^'T CONTENTS OF VOLUME XXIV No. Page 1. The Synthesis of Pseudodithiobiurets and Their Deriva- tives. H. G. Underwood and F. B. Dains 5 2. On the Constitution of Certain Thiazolidones. V: Phenyl, 0- and p-Tolyl Derivatives. F. B. Dains, Lois Miller Kinsett, C. 0. Holmberg and C. C. Robinson 15 3. On the Reactions of the Thiazolidones. VI: Some Deriva- tives of m-Toluidine. F. B. Dains, Robert D. Coghill and Samuel S. Tihen 25 4. On the Reactions of the Thiazolidones. VII: Some De- rivatives of l-Amino-2-methyl-5-nitro benzene. F. B. Dains and Pauline Garber 33 5. On the Reactions of the Thiazolidones. VIII: Some De- rivatives of 2-Amino-5-nitro-toluene and 2-Amino-3- bromo-5-nitro-toluene. F. B. Dains and Clarence E. Grothaus 37 6. On the Alkylation of the 2-phenylimino-5-dimethyl-Thia- zolidone and the 2-phenylimino-4-phenyl-Thiazoline. Floyd A. Eberly and F. B. Dains 45 7. Some Observations Regarding the Fate of Intravenously Injected Calcium Chloride. A. M. Lands and P. H. Woodard 51 8. Hydrocarbon Secretions and Internal Secretory Systems of the Carduaceae, Ambrosiaceae and Cichoriaceae. Mar- shall W. Mayberry 71 9. The Genus Taphrina. I: An Annotated Bibliography. A. J. Mix 113 10. The Genus Taphrina. II: A List of Valid Species. A. J. Mix 151 11. Two New Sunfish of the Family Centrarchidae from the Middle Pliocene of Kansas. Claude W. Hibbard 177 12. The Family Apioceratidae (Diptera) in North America. Reginald H. Painter I87 13. The Morphology of the Carolina Mantis. Philip Levereault, 205 14. Species of Erythroneura of the Comes Group (Homoptera- Cicadellidae) . R. H. Beamer 26 (3) 4 The University Science Bulletin No. Page 15. Alconeura of the United States (Homoptera, Cicadellidae). Melvin E. Griffith 309 16. A Generic Revision of American Bythoscopinae and South American Jassinae. P. W. Oman 343 17. A Contribution to the Taxonomy of the Subfamily Issinae in America North of Mexico (Fulgoridae, Homoptera). Kathleen C. Doering 421 18. Description of a New Mexican Subspecies of Sceloporus spinosus Wiegmann (Lacertilia). Hobart Smith 469 19. Notes on the Herpetological Fauna of the Mexican State of Sonora. Edward H. Taylor 475 20. Notes on the Herpetological Fauna of the Mexican State of Sinaloa. Edward H. Taylor 505 21. The Lizards of the Torquatus Group of the Genus Scelop- orus Wiegmann, 1828. Hobart Smith 539 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 1. The Synthesis of Pseudodithiobiurets and Their Derivatives H. G. UXDERWOOD and F. B. Dains, Department of Chemistry, University of Kansas. ARYL monopseudodithiobiurets of the type RNHCSNHC (SR') : NH (R represents aryl groups; R' alkyl) are readily prepared by the decomposition of the sulphate salt of a pseudothiourea with sodium carbonate and immediate reaction of the pseudothiourea with a mustard oil, HoNCfSR') :NH + RNCS =: RNHCSNHC (SR') :NH. In previous work^ the alkyl halide addition product of thiourea was decomposed with potassium hydroxide. Diaryl monopseudodithio- biurets of the type RN:C(SR')NHCSNHR (R represents aryl groups; R' alkyl) were prepared by the reaction of a mustard oil with a pseudothiourea in ether solution. The reaction is smooth and the yields are practically ciuantitative, confirming the work of Johnson.^ Johnson found that a monopseudodithiobiuret is converted to a dipseudodithiobiuret by the action of alkyl halides; methyl iodide being used as one of the alkylating agents. Methyl sulphate reacts very smoothly in acetone solution for the conversion of a mono- pseudodithiobiuret into a dipseudodithiobiuret. The sulphate salt first formed is easily decomposed, giving the free base, on treating a cold aqueous solution with sodium carbonate, the yield being quantitative. Pseudodithiobiurets thiohydrolyze readily on warming an alcoholic solution with potassium hydrosulphide. According to the method of Olin and Dains- hydrogen sulphide is slowly bubbled through the alcoholic solution. Good results were obtained omitting the use of hydrogen sulphide; thus 1, 5-diphenyldithiobiuret previously pre- pared by Olin and Dains was obtained by thiohydrolysis of the 1. Johnson, Am. Chem. J., 30, 167 (1903). 2. Olin, Dains, J. Am. Chem. Soc, 52, 3326 (1930). (5) 6 The University Science Bulletin 2-thiolmethyl derivative using only potassium hydrosulphide as the thiohydrolytic agent. Fusion of 1, 5-diphenyl-2-thiolmethyl-pseudodithiobiuret with phenylhydrazine yielded diphenyl-thiosemicarbazide corresponding to the results obtained by Johnson and Bristol^ from 1, 5-diaryl- pseudodithiobiurets. The 1, 5-diaryl-dipseudodithiobiurets were found to react with hydrazine, phenylhydrazine and substituted phenylhy- drazines with the liberation of mercaptan to yield 1, 2, 4-triazols of I I the type R'NC(:NR)NHC(NR)NH (R represents aryl groups; R' hydrogen or aryl groups). Triazols of the above type are also formed by the reaction of hydrazines on oc, oc '-diary 1-thiurets. Ring closure of the substituted guanido-arylthioureas first formed, to the 1, 2, 4-triazols results on treatment with alkali. New thiurets were prepared by the method of Fromm and Schneider* according to the reaction: RNHCSNHCSNH, + L = SC(:NR)NHC(S) :NH.HI + HI While thiurets previously reported have all been of the mono-aryl type our work shows that diaryl thiurets of the type SC(:NR)NHC(S) :NR are readily prepared from 1, 5-diaryl-dithio- biurets; thus 1, 5-diphenyl-dithiobiuret on oxidation in alcoholic solution with iodine yielded 1, 5-diphenyl-thiuret hydroiodide. Condensation of dithiobiurets with acetone and benzaldehyde to form keturets and aldurets was affected by means of dry hydrogen chloride according to the method of Fromm.-'' EXPERIMENTAL Synthesis of Mono- and Dipseudgdithigbiltiets 1 - Phenyl - 4 - thiohnethyl - pseudodithiobiuret, CJir^NHCSNHC {SCHr.) :NH.— This was formed in good yield when methyl isothio- urea sulphate dissolved in an alcohol-water mixture (1 + 1) at 70° was allowed to react gradually with phenyl mustard oil after libera- tion of the free base with sodium carbonate solution. The product on recrystallization from alcohol gave white prism-like crystals melt- ing at 124° (reported 125°).^ Other pseudodithiobiurets from an aryl mustard oil and an alkyl isothiourea sulphate were prepared by this method. 1. Johnson, Am. Chein. J., 30, 1G7 (1903). 3. Johnson, Bristol, Am. Chem. J., 30, 173 (1903). 4. Fromni, Schneider, Ann, 348, 161 (1906). 5. Fromm, Ann., 275, 20 (1893). Underwood and Dains: Pseudodithiobiurets 7 Thiohydrolysis of l-Phenyl-4-thiolmethyl-pseudodithiobiuret to Phenyldithiobiuret, C^H.NHCSNHCSNH,.— The thio ether (4 g.) was dissolved in hot alcohol (50 cc.) together with 2.5 g. of potas- sium hydrosulphide in 5 cc. of water. Hydrogen sulphide was passed into the boiling solution for an hour. The hot solution was filtered and acidified with acetic acid. On cooling, delicate white, feather- like crystals of phenyldithiobiuret melting at 184° (reported 184°)® separated. Analyses: Calcd. for CsHgNsS.rN, 19.89. Found: N, 19.49; 19.79. 1 -m- Tolyl-4-thiob7iethyl - pseudodithiobivret, C.H.NHCSNHC (SCH^) :NH. — Methyl isothiourea sulphate when treated with m- tolyl mustard oil and sodium carbonate solution yielded the pseudo- dithiobiuret. Crystallized from alcohol, it then melted at 87°. Analyses: Calcd. for C,oH,3N3S,,:N, 17.51. Found: N, 17.23; 17.25. l-p-Bromophenyl-4-thioimcthyl-pseudodithiobiuret, BrCJI^NHC SNHC{SCHs):NH. — This was prepared from p-bromophenyl iso- thiocyanate and methyl isothiourea sulphate. Purified from alcohol, it melted at 154°. Analyses: Calcd. for CgHioNgS^Br: N, 13.81. Found: N, 13.50; 13.44. 1 - p-Brojnophenyl-4-thiolethyl - pseudodithiobiuret, BrCf^H^NHC SNHC{SC2H^):NH. — This product, prepared from p-bromophenyl isothiocyanate and ethyl isothiourea sulphate, melted at 147°. Analyses: Calcd. for CioHj.Br^^S.: N, 13.20. Found: N. 13.12; 13.24. 1-Phenyl - 5 - allyl -2 - thiolmethijl-pseudodithiobiuret, C^H^NiC {SCH.,)NHCSNHCH^CH:CH^.— On spontaneous evaporation of a solution of molecular proportions (0.05 moles) of allyl mustard oil and the methyl ether of phenylthiourea in 75 cc. of ether, a crystal- line body was obtained together with gummy material. The crystals were separated by the addition of alcohol and filtering. Recrys- tallized from alcohol, the compound melted at 100-101°. Analyses: Calcd. for CjoH^^NsS,: N, 15.85. Found: N, 15.79; 15.79. Thiohydrolysis of 1 , 5-Diphenyl-2-thiolmethyl-pseudodithiobinret ; 1, 5-Diphenyl-dithiobiuret, C^H,NHCSNHCSNHC,H,.— The thio ether (18 g.) prepared by the method of Johnson^ was dissolved in hot alcohol (100 cc.) together with 10 g. of potassium hydrosulphide 1. Johnson, Am. Chem. J., 30, 167 (1903). 6. Fromm, Ann., 275, 33 (1893). 8 The University Science Bulletin in 20 ce. of water. The hot solution after refluxing for one hour was filtered and acidified with acetic acid. On cooling, fine needles of 1, 5-diphenyl-dithiobiuret melting at 149° (reported 149°)^ sep- arated. 1 - Phenybnethyl - 5 - phenyl-dithiobiuret, C^H^iCH^) iNCSNHC SMHC^H,. — Thiohydrolysis of 1, 5 - diphenyl - 1 - methyl - 2-thiol- methyl-pseudodithiobiuret yielded crystals which melted at 126°. Analyses: Calcd. for C15H15N3S2: N, 13.95. Found: N, 13.36; 13.69. 1 - p - Bromophenyl - 5 - phenyl- J^-thiolmethyl-pseudodithiohiuret, BrC,H,NHCSNHC{SCH,) :NC,H,.—On allowing an ether solution of phenyl thiolmethylpseudothiourea (0.02) and p-bromophenyl iso- thiocyanate (0.02 mol.) to evaporate spontaneously there was ob- tained a crystalline body which on recrystallization from alcohol melted at 140-141°. Analyses: Calcd. for CisH^^BrNgS,: N, 11.05. Found: N, 10.98; 10.97. 1 -p- lodophenyl - 5 - phenyl - 4 - thiolmethyl-pseudodithiobiuret, IC,H,NHCSNHC{SCH,) -.NC^H,.— The product obtained by allow- ing phenyl thiolmethylpseudothiourea to react with p-iodophenyl isothiocyanate crj^stallized from hot alcohol in the form of prisms melting at 126°. Analyses: Calcd. for CigH.JNaS.: N, 9.84. Found: N, 9.90; 9.77. l-7n-Tolyl-5-phenyl-4-thiolmethyl-pseudodithiobiuret, C ^H^NHC SNHCiSCH^):NC^,H^. — Light yellow, transparent plate-like crys- tals melting at 87° resulted on allowing m-tolyl isothiocyanate to react with phenyl thiolmethylpseudothiourea dissolved in ether. Analyses: Calcd. for CieH^^NsS,: N, 13.33. Found: N, 13.34; 13.48. 1 -p- Tolyl-5-phenyl-4-methyl-pseudothiobiuret, C^H^NHCONC {SCH^):NCeH^. — p-To\y\ isocyanate (0.03 mol.) reacted with phenyl thiolmethylpseudothiourea with the evolution of heat. The clear, viscous product first formed solidified on standing. Washed with ether and recrystallized from alcohol the methyl thiobiuret melted at 109-110°. Analyses: Calcd. for C15H16N3OS: N, 14.09. Found: N, 13.98; 14.01. 1, 5-Diphenyl -2,1^- thiolmethyl - dipseiidodithiobiuret, C^H^^N-.C iSCH,)NHC{SCH,):NC^H,. — Methyl sulphate (0.1 mol.) was slowly added to a solution of 1, 5-diphenyl-2-thiolmethyl-pseudo- 2. Clin, Dains, J. Am. Chem. Soc, 52, 3326 (1930). Underwood and Dains: Pseudodithiobiurets 9 dithiobiuret (0.1 mol.) in acetone (100 cc.) and the reaction mixture refluxed on the water-bath for two hours. The light-yellow, gummy residue obtained on evaporation of the acetone, after dissolving in alcohol and treatment with sodium carbonate, jnelded an oil which soon solidified. The product on recrystallization from alcohol melted at 104° (reported 103-104° ).i Analyses: Calcd. for CisHi^NgSg: N, 13.33. Found: N, 13.24. 1 - Phenyl -2 -, 4 - thiohnethyl - dipseudodithiobkiret, CJI^N.C (SCH,)NHCiSCH,):NH.— The free base obtained by the reaction of methyl sulphate with the 4-thiolmethyl derivative crystallized from hot alcohol in the form of long needle-like prisms melting at 123°. A mixed melting point with the starting product was 99-101°. Analyses: Calcd. for CioH^gNgS,: N, 17.54. Found: N, 17.27; 17.60. Synthesis of 1, 2, 4-Triazols Fusion of Phenylhydrazine with 1,5- Diphenyl - 2 - thiolmethyl- pseudodithiobiuret; Formation of Diphenylthiosemicarbazide, C^H^ NHNHCSNHC^H^.—Six grams of the pseudodithiobiuret and 2.5 g. of phenylhydrazine were fused together in an eight-inch Pyrex test tube. The odor of mercaptan was first noted at 90° and after two hours at 110° the reaction mixture had again solidified. The solid recrystallized from alcohol in the form of short, colorless needles melting at 176°. INIixed with diphenyltliiosemicarbazide the melting point was not lowered. l-Phenyl-3, 5-phenylimino-l , 2, 4-triazol, C^H^ NC{:NC^H,)NHC{NH):NC,H,.— Four grams of phenylhydrazine mixed with 10.5 g. of 1, 5-diphenyl-2, 4-thiolmethyl-dipseudodithio- biuret first reacted with the liberation of mercaptan at 110°. The reaction temperature was maintained between 110° and 140° until the odor of mercaptan was no longer noticed. The fusion product after two crystallizations from alcohol gave delicate white needles melting at 153-154°. A negative test for sulphur and the analysis (Dumas method) for nitrogen indicated formation of the triazol according to the following reaction : CeH3N:C(SCH3)NHC(SCH3) :NCeH, + CH.NHNH^ = CeH^NC ( : NCeH, ) NHC ( NCeH, ) : NH + 2CH3SH Analyses: Calcd. for CooH^.N^: N, 21.43. Found: N, 21.59. 1. Johnson, Am. Chem. J., 30, 167 (1903). 10 The University Science Bulletin The salt formed with sulphuric acid melted at 190°. Analysis by titration of an alcoholic solution of the salt with standard base in- dicated the addition of one mol. of H2SO4. Analyses: Calcd. for addition of one mol. H0SO4: 23.06. Found: 24.96. Phenyl - anilguanido - phenylthiourea, C^HJSI : C ( NHNHCpH-J NHCSNHCqH^. — A flocculent white precipitate resulted on refluxing an alcoholic solution containing molecular proportions of 1, 5-di- phenyl-thiuret hydroiodide and phenylhydrazine on the water-bath. The reaction product was dissolved in a large quantity of hot alco- hol and filtered free from sulphur. On cooling, white needles of the substituted thiourea melting at 167° separated. Analyses: Calcd. for aoH^gN.S: N, 19.39. Found: N, 19 32. l-Phenyl-3, 5-phenylimino-l , 2, ^-triazol from Phcnyl-anilgiian- ido-phenylthiow'ea. — Ring closure to the 1, 2, 4-triazol resulted on warming three grams of phenyl-anilguanido-phenylthiourea in an alcoholic sodium hydroxide solution (1 g. NaOH, 5 cc. water, 25 cc. alcohol) for one hour at water-bath temperature. The needle-like crystals of the triazol which separated on cooling melted at 153- 154°. A mixed melting point of this product with the triazol from 1, 5-diphenyl-2, 4-thiolmethyl-dipseudodithiobiuret showed no low- ering. Analyses: Calcd. for C^oH^.N^: N, 21.43. Found: N, 21.36. l-p-Bromophenyl-3, 5-phenylimino-l, 2, J^-triazol, BrC,H,NC{:NC,H,)NHC{NC,H,) -.NH.— This triazol was obtained on fusion of molecular proportions of 1, 5-diphenyl-2, 4-thiolmethyl-dipseudodithiobiuret and p-bromo- phenylhydrazine. Mercaptan was first liberated at 100° and on heating at 110° for several hours the odor of mercaptan was no longer noticed. Crystallized from alcohol in the form of fine white needles, it then melted at 190°. Analyses: Calcd. for C^oHieBrN^: Br, 19.68. Found: Br, 19.95. l-{o-Tolyl)-S, 5-phenylimino-l , 2, 4-triazol, C,H,NC{:NC,H,)NHC{NH) :NC,H,.— Molecular proportions of o-tolylhydrazine and 1, 5-diphenyl-2, 4- thiolmethyl-dipseudodithiobiuret were allowed to react for several Underwood and Dains: Pseudodithiobiurets 11 hours at 110°. The resulting triazol recrystallized from alcohol in the form of delicate white needles melted at 174^ A negative test for sulphur and the analysis indicated formation of the triazol. Analyses: Calcd. for CoiHigNgi N, 20.53. Found: N, 20.61. l-ip-Tolyl)-3, 5-phenylimmo-l, 2, J^-triazol, C,H,NCi:NC,H,)NHC{NC,H,) :NH.— This triazol melting at 161° was obtained from p-tolylhydrazine and the above dipseudodithiobiuret. Analyses: Calcd. for C.^HigNg: N, 20.53. Found: N, 20.40. I f 3, 5 - Phenylimino - 1 , 2, I^- triazol, HNC {■.NC,H,)NHC (NH) : NCgHr,. — Hydrazine [0.02 mol. of hydrazine sulphate and potas- sium hydroxide] and 1, 5-diphenyl-2, 4-thiolmethyl-dipseudodithio- biuret in 300 cc. of alcohol were refluxed at water-bath temperature for four hours. Mercaptan was slowly evolved during the reaction and a white flocculent mass of fine white needlelike crystals sep- arated from the reaction solution. Recrystallized from alcohol, it then melted at 250-251° and gave a negative test for sulphur. Analyses: Calcd. for d^HigN^: N, 27.89. Found: N, 27.72. With concentrated sodium hydroxide solution and also with bleaching powder a deep red coloration was observed. The forma- tion of azo compounds is typical of urazols of this type.^ 3, 5-PhenyUmino-l , 2, 4-tnazol from ^, <^'-Diphenyl-thiuret Hy- droiodide. — Hydrazine sulphate (0.02 mol.) and ^, oc'-diphenyl- thiuret hydroiodidc (0.01 mol.) in alcohol (50 cc.) were refluxed together for one hour. An excess of alcoholic sodium hydroxide was then added and the reaction mixture further refluxed for one half hour. The solution was filtered while hot from precipitated sulphur and on cooling the crystals which separated melted at 244°. A slight test for sulphur, the melting point 6° lower than for the above triazol and the low nitrogen analysis indicated traces of the thiuret. Analyses: Calcd. for Ci,H,3N5: N, 27.89. Found: N, 27.26. The Benzoyl Derivative of 3, 5 -Phenylimino- 1 , 2, 4-triazol. — To several grams of the triazol dissolved in pyridine was added an 7. Pellizzari, Gazetta chimica italiana, 21, II, 141 (1891). 12 The University Science Bulletin excess of three molecular proportions of benzoyl chloride. The re- action mixture warmed spontaneously and on addition of water an oily product separated which solidified on cooling. The benzoyl derivative crystallized from alcohol in the form of light yellow needles melting at 136°. Analysis indicated formation of a mono- benzoyl derivative. Analyses: Calcd. for C^iH^.N^O: N, 19.72. Found: N, 19.54. Reaction of p-Tolyl Isocyanate with 3, 5-Phenylimino-l , 2, 4-tn- azol. — Several grams of the triazol were suspended in water-free ether and an excess of three molecular proportions of p-tolyl isocy- anate added. Complete solution of the triazol was effected on warming the reaction mixture. The solid which separated on re- fluxing melted at 188° when recrystallized from heptane. Analysis indicated addition of two molecules of the isocyanate. Analyses: Calcd. for CgoHo.N.O.: N, 18.95. Found: N, 18.99. Synthesis of Thiurets, Keturets and Aldurets ^-m-Tolyl-thiuret Hydroiodide, SCi:NC,H,)NHC {S) -.NH .HI. — Iodine (0.05 mol.) dissolved in alcohol was slowly added to a warm solution of m-tolyldithiobiuret (0.05 mol.) in alcohol. The reaction mixture was refluxed until the color remained constant. The white crystalline product which separated on cooling, after crystallization from hot alcohol melted at 194°. Analysis indicated the retention of one molecular proportion of alcohol of crystalliza- tion. Analyses: Calcd. for CgHiJNsS.. C.HgOH: N, 10.58. Found: N, 10.48; 10.45. o^-p-Bromophenyl-thiuret Hydroiodide, SCi:NCf,H^Br)NHC (S): NH.HI. — This product was obtained in the form of light-yellow needles melting at 237° on oxidation of p-bromophenyl-dithiobiuret with iodine. One molecule of alcohol of crystallization is retained. Analyses: Calcd. for CgH.BrlNsSo.CoHgOH: N, 9.07. Found: N, 9.02; 8.85. ^, ^' - Diphenyl - thiuret Hydroiodide, SC {:NC^H,)NHC (S) : NCJH^.HI. — Oxidation of 1, 5-diphenyl-dithiobiuret in alcoholic Underwood and Dains: PsEUDODiTHiOBirRETS 13 solution with iodine resulted in the formation of bright yellow plate- like crystals, which when recrystallized from alcohol melted at 192°. Analyses: Calcd. for C14H12IN3S2: N, 10.17. Found: N, 10.04; 10.09. ^-m- Tolyldithiodi - c - methyl Keturet, C.H.NHCSNCSNHC I I {CH^)r,. — Five grams of m-tolyldithiobiuret and three grams of ace- tone were placed in a flask and with continual shaking a strong stream of dry HCl gas was introduced. The crystalline mass liquefied and soon resolidified. The reaction product was dissolved in warm 10% sodium hydroxide and filtered, and from the cooled solution the keturet was precipitated as a white powder on acidifying with hydrochloric acid. The keturet crj'stallized from alcohol, in which it is only moderately soluble, in the fomi of white platelike crystals melting at 235-236° with darkening and decomposition. Analyses Calcd. for Ci.H.sNsS^: N, 15.86. Found: N, 15.67; 15.77. ^-m-Tolyl-c-phenyl-dithioalduret, C.H.NHCSNCSNHCH . C^H,. — Dry hydrogen chloride gas effected the condensation of benzalde- hyde with the equivalent of p-tolyldithiobiuret to the alduret. This product, when purified by the method for the corresponding keturet, melted at 189°. Analyses: Calcd. for Ci^H^sNaS.,: N, 13.42. Found: N, 13.44. oc _ ( oc . Naphthyl) - dithiodi - c - methyl Keturet, C^qH^NHCS NCSNHCiCH^)^. — oc-Naphthyl-dithiobiuret condensed with ace- tone in the presence of dry hydrogen chloride gas. When purified and crystalized from hot alcohol, the keturet melted at 225°. Analyses: Calcd. for C16H15N3S2: N, 13.42. Found: N^ 13.42. °^-Phenyl - dithiomethyldi - c- methyl Keturet, C ^H .N :C [SCH .^) NC{SCH,)NC{CH,),.— To a solution of oc _ phenyl - dithiodi - c - methyl keturet (0.03 mol.) in dilute sodium hydroxide was added an excess of methyl iodide (10 g.) in alcohol solution. After several hours at room temperature a crystalline product had separated. Crystallized from hot alcohol the product melted at 134°, Analyses: Calcd. for CiaHi^NsS^: N, 15.05. Found: N, 14.86; 14.91. 14 The University Science Bulletin SUMMARY 1. Methyl sulphate reacts smoothly for the conversion of mono- pseiidodithiobiurets into dipseudodithiobiurets. 2. A method has been described for the synthesis of 1, 2, 4-tri- azols. 3. Several new thiurets, keturets and aldurets have been prepared. i THE UNIVERSITY OP KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 2. On the Constitution of Certain Thiazolidones. V: Phenyl, o- and p-Tolyl Derivatives F. B. DAINS, LOIS MILLER KINSETT, C. O. HOLMBERG and C. C. ROBINSON Contributions from the Chemical Laboratory of the University of Kansas IN previous papers from this laboratory a study has been made of various aryl and acyl substituted thiazolidones. (1) In the following series of papers we wish to put on record the results of other investigations that have been carried out in this same field. When monophenylthiourea is treated \nth chloroacetic acid, chloroacetyl chloride or ethyl chloroacetate there is formed a labile thiazolidone — A. SC(NH)NCsH5C0CH. which quickly rearranges to the stable form B. S-C(NHCeH5)NC0CH, or the tautomeric form C. SCCNCeHJNHCOCH. (2) Wheeler and Johnson found that the sodium salt of the stable form reacted with benzyl chloride with the formation of 2-benzyl-2- phenylamino-4-thiazolidone whose constitution was proven by its synthesis from unsymmetrical benzyl-phenylthiourea and ethyl chloroacetate. Beckurts and Frerich (3) have stated that the monoaryl thiazoli- dones have the structure of the labile form A and that their sodium (1). Walter S. Long and F. B. Dains: Trans. Kan. Acad. Sci., vol. 36 (1933). F. B. Dains and Floyd A. Eberly : J. Am. Ch. Soc. 55, 3859 (1933). John A. Da\ns and F. B. Dains: J. Am. Ch. Soc. 57, 2627 (1933). (2). Wheeler and Johnson: Am. Chem. Jour. 28, 121, 146 (1902). (3). Archiv. d. Pharm. 253, 233 (1915) C. A. 10, 888 (1916). (15) 16 The University Science Bulletin salts gave with ethyl iodide, the 2-ethylimino-3-aryl-4-thiazolidones, results that are in contradiction to the observations of Wheeler and Johnson and also to our own work which confirmed the stable form B as 2-arylamino (or imino-) thiazolidone as postulated by these investigations. In addition, we have found that alkylation of the sodium salts gave mainly the 2-alkyl-arylamino thiazolidones together with yields ranging from 0 to 85% of the 2-arylimino-3-alkyl isomers; the relative yields seemed to depend upon the nature of the alkyl halide and the aryl group. Thus it was found that ethyl iodide and the sodium salt of 2-diodo phenyl thiazolidone gave 85% of the 3- ethyl derivatives and none of the 2-ethyI-2-diodo phenyl compound. (4) These results then are in absolute contradiction to the formula- tions of the reaction by Beckurts and Frerich who not only reversed the position of the aryl and alkyl groups but failed to isolate the isomeric thiazolidones. EXPERIMENTAL DERIVATIVES OF THE 2-PHENYL-THIAZOLIDONE 2-METHYL-2-PHENYLAMINO-THIAZOLIDONE I S-C (NCH3 . CeHr,) NCOCH2 The sodium salts were readily prepared by dissolving the 2-aryl- thiazolidones in hot 5-10% sodium hydroxide solution. The salt which crystallized on cooling was filtered, washed with a little cold water and dried for use. This sodium salt was heated in methyl alcohol solution with methyl iodide in a sealed tube at 100° for eight hours. The same results are achieved by simply refluxing the components in a water bath, or by methylation with methyl sulphate. After removal of alcohol and excess methyl iodide the oily residue was dissolved in ether and the etherial solutions repeatedly ex- tracted with 10% hydrochloric acid. Neutralization of the acid solution gave the thiazolidone I, which crystallized from alcohol in light needles melting at 125°. Its identity was proved by com- parison with the same compound (5) obtained by heating molar quantities of unsymmetrical methyl-phenylthiourea, ethyl chloro- acetate and pyridine (1 mol.) in alcohol solution or by the action of chloroacetyl chloride and pyridine (2 mols.) on the thiourea in cold acetone solution. The addition of pyridine prevents hydrolysis of the thiazolidone and increases the yields. (4). John A. Davis and F. B. Dains : J. Am. Ch. Soc. 57, 2628 (1935). (5). Dixon, J.: Ch. Soc. 71, 620 (1897). Dains et al. : Thiazolidones 17 2-METHYL-PHENYLAMINO-5-BENZAL-THIAZOLIDONE 11 S-C(NCH3.C6H5)NCOC = CHCcHr, This was prepared by boiling for a few minutes a mixture of the thiazolidone (1-pt), benzaldehyde (1-pt), glacial acetic acid (3-pts), acetic anhydride (1-pt) and fused sodium acetate (0.5 pt). The same benzal derivative which was obtained from both prepara- tions (methylation of I and from the unsymmetrical thiourea) was difficultly soluble in alcohol and melted at 180°. These 5-benzal compounds which are useful for the identification of the substituted thiazolidones can be synthesized also by heating the aldehyde and the thiazolidone for thirty minutes at 200-220° or more simply by adding to an alcoholic solution of the components a few drops of 10% sodium hydroxide or of piperidine. Heating the mixture is sometimes necessary. 2-PHENYLIMINO-3-METHYL-THIAZOLIDONE III S-C( NCeHr.) NCH3COCH1; This was isolated in two percent yield from the original ether solution from which acid had extracted I. It was insoluble in dilute acid and the crystals from heptane melted at 60°. For comparison it was synthesized from symmetrical methyl- phenylthiourea and ethyl chloroacetate on refluxing the alcohol so- lution with pyridine (1 mol.) the two preparations being identical. This is the type of compound that Beckurts and Frerich thought they had obtained but with the groups interchanged. HYDROLYSIS When the thiazolidone III was boiled in alcohol solution with 10% hydrochloric acid, the ring was split, yielding thioglycollic acid, methylamine, aniline and a little diphenylurea. When the methy- lene hydrogens at position 5 were replaced by two phenyl groups or a benzal grouping the ring was greatly stabilized. Thus 2-phenyl- amino-3-methyl-5-benzol-thiazolidone IV (made as was II) was dissolved in concentrated sulphuric acid and heated at 115° for fifteen minutes. After pouring the reaction mixture into ice-water, there were isolated aniline and 3-methyl-5-benzal-2-4-tliiazoldione V (m. p. 135°), the formation of which proved the positions of the methyl and phenyl groups. 2—6037 18 The University Science Bulletin THE ETHYLATION OF 2-PHENYLAMINO-THIAZOLIDONE The sodium salt and ethyl iodide gave the 2-ethyl-2-phenyI amino-thiazolidone VI (m. p. 118°). It was soluble in acid and identical with the compound obtained from unsymmetrical ethyl- phenylthiourea and ethyl chloroacetate. Each preparation gave the same 5-benzal derivative VII, melting at 165°. The 2-phenylimino-3-ethyl-thiazolidone VIII was formed in small amounts in the above reactions, but was synthesized for com- parison from symmetrical ethyl-phenylthiourea and ethyl chloro- acetate. The crystals from ether melted at 74°. Its 5-benzal de- rivative IX melted at 100°. The analogous n-butyl derivatives X and XI are thick oils. ACETYL DERIVATIVES OF 2-PHENYLAMINO-THIAZOLIDONE Wheeler and Johnson (6) on warming the "labile" phenyl thiazo- lidone with thioacetic acid in benzene solution obtained a com- pound melting at 192°, to which was given the formula 2-acetyl- imino-3-phenylthiazoliclone. The same product (m. p. 192°) was obtained by us when unsymmetrical phenyl-acetyl-thiourea was treated with chloroacetyl chloride in benzene and pyridine solution. This would indicate that it was in fact 2-acetyl-2 phenylamino- thiazolidone, XII. SC(NC,H,.COCH3)NCOCH3, otherwise it would necessitate the rearrangement of the acetyl-phenyl thiourea to the symmetrical form and then reaction. Dixon has found, however, that the symmetrical acetyl-phenyl thiourea failed to react with chloroacetyl chloride, which tends to confirm the above formulation. The acetyl group was readily removed on treatment with cold normal sodium hydroxide, regenerating the 2-phenylthiazolidone and it (XII) was not further acetylated on boiling with acetic anhydride. The analogous 2-methyl-2-phenylamino-thiazolidone also failed to react with acetic anhydride showing the inertness of the sub- stituted amino form ; R2NC = of the thiazolidone. Treatment of the 2-phenylamino-thiazolidone with chloroacetyl- chloride in pyridine and benzene solution gave negative results. THE DIACETYL DERIVATIVE Wheeler and Johnson had obtained a diacetyl compound XIII (m. p. 162°) on warming the 2-phenylthiazolidone with acetic an- (6). Wheeler and Johnson, Am. Chem. Jour. 28, 143 (1902). Dains et al.: Thiazolidones 19 hydride which was confirmed by our experiments. The acetyl groups were easily removed by cold alkali or warming with dilute ammonium hydroxide. The diacetyl compound was heated gently with amilins and from the reaction mixture were isolated acetanilide and the original phenylthiazolidone (m. p. 176°). A possible explanation is that XIII has an acetyl group at 3 and an acetate grouping at 4, viz., SC(NCeH5)NCOCH3C-OCOCH3CH, which would account for the formation of acetanilide. A diacetyl derivative is only possible with a potential grouping -NH-C-OH, wliich is not the case with the di-substituted aminothiazolidones. DERIVATIVES OF 2-p-TOLYL-AMINO OR IMINO- THIAZOLIDONE (m. p. 187°) This was readily made from mono-p-tolylthiourea and chloro- acetic acid or from chloro-acet-p-toluidide on boiling with potassium thiocyanate in alcohol solution. 2-P-TOLYL-2-ETHYL-AMINO-4-THIAZOLIDONE XIV The sodium salt was refluxed in alcohol solution with ethyl iodide for four hours. The oil remaining after steam distillation slowly solidified. Crystallized from alcohol it melted at 124°. The yield was sixty-five percent. Hydrolysis. The compound was heated with dilute hydrochloric at 140° for four hours. The tube opened with pressure due to carbon dioxide and the solution gave a test for thioglycollic acid. It was then made alkaline, extracted with ether and the etherial solution dried with solid potassium hydroxide. Evaporation of the solvent left an oil Ethyl-p-tolylamine, which with phenylisocyanate gave oc -ethyl- oc-p-tolyl-^-phenylurea XV (m. p. 67°) identical with a synthetic specimen of the urea. THE BENZAL DERIVATIVE XVI This was formed when molar quantities of benzaldehyde and XIV were heated at 230°. It was very difficultly soluble in alcohol and melted at 179^ Heated at 140° with dilute acid, it was hydrolyzed into ethyl-p-tolylamine and 5-benzal-2, 4-thiazoldione, XVII (m. p. 242°) (7). This was confirmed by analysis and comparison with the known compound resulting from the hydrolysis of 2-imino-5- benzal-thiazolidone. (7). Andreash, Monats. 10, 75. 20 The University Science Bulletin 2-P-TOLYLIMINO-3-ETHYL THIAZOLIDONE XVIII This was made from cx-ethyl-^-p-tolyl-thiourea and chloroacetic acid and was an oil. This was refluxed for twenty hours in an alcohol-hydrochloric acid solution. p-Toluidine was isolated and also a small quantity of a solid that melted at 140° after recrystal- lization from water and gave figures on analysis that correspond to 3-ethyl-2-4-thiazoldione XIX. The 5-benzal-derivative from XVIII, light yellow crystals from alcohol, melted at 132°, XX. On hydrolysis it gave p-toluidine and 3-ethyI-5-benzal-2-4-thiazoldione XXI (m. p. 97°), thus confirm- ing the structure. BENZYL AND p-NITROBENZYL DERIVATIVES The 2-benzyl-2-p-tolylamino-thiazolidone XXII was made from the sodium salt and benzyl chloride. It crystallized from alcohol in rhombic needles that melted at 121°. On acid hydrolysis at 140° benzyl-p-tolylamine was isolated, which with phenylisocyanate formed the known °= -benzyl- cc-p-tolyl ^-phenylurea. 2-P-NITROBENZYL-2-P-TOLYL-AMINO-THIAZOLIDONE XXIII This crystallized from alcohol in light yellow needles melting at 142°. p-Nitrobenzyl-p-tolyl-amine XXIV (m. p. 66.7°) was one of the products of hydrolysis. This latter compound was also made for comparison from p-nitrobenzyl chloride and p-toluidine. DERIVATIVES OF 2-o-TOLYL-AMINO-THIAZOLIDONE (m. p. 144) This was made in the usual manner, both from mono-o-tolyl thiourea and from the chloro-acet-o-toluidide. The sodium salt melted at 105° with loss of water, then solidified and melted again at 250°. The salt dried at 80° gave figures cor- responding to equal mols. of the thiazolidone and sodium hydroxide. Analyses for C,,H,oN30S,NaOH. Calcd: Na. 9.35; N, 11.38. Found: Na. 9.54; N, 11.15. 2-ETHYL-2-0-TOLYLAMINO-THIAZOLIDONE XXV This was obtained from the sodium salt and ethyl iodide as an oil which, after long standing, ciystallized (m. p. 98°). Hydrogen chloride gas precipitated a salt from a solution of the oil in dry ether (m. p. 165-8°). Analyses for Ci,H,,N,OS-HCl. Calcd: HCl, 10.35. Found: HCl, 10.21. Its constitution was shown by its acid hydrolysis yielding ethyl- o-tolylamine, which was identified by the formation of oc-ethyl-o^-o- Dains et al. : Thiazolidones 21 tolyl-;S-phenyliirea (m. p. 89°) and its identity with a known sample. The 5-benzal-derivative melted at 193° and the isomeric 2-o-tolyl- imino-3-ethyl-thiazolidone at 74°. Proof of the constitution of the 2-aryl-2-alkyl-amino-thiazolidones by their synthesis from the corresponding unsymmetrical thioureas was not found to be practical in several cases. The usual procedure for the preparation of such thioureas by evaporation of a water solution of their hydrogen chloride salts with ammonium thiocyanate failed to give the unsymmetrical thioureas with the following amines; viz., m-tolyl-ethyl, o-tolyl-methyl, o-tolyl-ethyl, o-tolyl- benzyl and p-tolyl-benzyl. 2-METHYL-2-0-TOLYLAMINO-THIAZOLIDONE XXVI This was made from the sodium salt and methj'-l iodide. A 60^ yield of the compound was obtained, which melted at 108°. From the acid hydrolysis at 140°, was isolated mcthyl-o-tolylamine, identi- fied by the foraiation of ^^ -methyl- «:-o-tolyl-/3-phenylurea (m. p. 82°). ^ 2-0-TULYLIMINO-3-METHYL-THIAZOLIDONE XXVII This was formed in the usual manner from oc-methyl-^-o-tolyl- thiourea and chloracetic acid. It was insoluble in dilute acid, melted at 59° and gave a 5-benzal-derivative XXVIII (m. p. 129°). 2-0-TOLYL-2-BENZYL-AMINO-THIAZOLIDONE XXIX The sodium salt was refluxed with benzyl chloride in alcohol solution and then steam distilled; the residual oil solidified after several months standing and was then purified by recrystallization. It melted at 95° and was obtained in 40% yield. Hydrogen chloride gas precipitated a salt XXX from its ether solutions (m. p. 179°). 22 The University Science Bulletin 4-Thiazolidone. Formula. M.p/C. Percent nitrogen. No. Calcd. Found. I 2-methyl-2-phenyl CioHioNoOS 126 13.59 13.58 II 2-methyl-2-phenyl amino-5-benzal C17H14N0OS 180 9.53 9.30 III 2-phenylimino- 3-niethyl C10H10N2OS 60 13.59 13.50 IV 2-phenyliinino- 3-methyl-5-benzal C17H14N0OS 135 9.53 9.55 V 3-methyl-5-benzal- 2-4 thiazolidone C11H9NO2S 135 6.39 6.79 VI 2-ethyl-2-phenyl- amino C11H12N2OS 118 12.73 12.51 VII 2-ethyl-2-phenyl amino-5-benzal C18H16N2OS 165 9.09 9.18 VIII 2-phenylimino- 3-ethyl C11H12N2OS 74 12.73 12.67 IX 2-phenylimino- 3-ethyl-5-benzal CisHioNoOS 100 9.09 8.81 X 2-n-Butyl-2- phenylamino C13H16N2OS oil 11.29 11.29 XI 2-phenylimino- 3-n-butyl C13H16N2S oil 11.29 11.28 XII 2-phenyl-2- acetylamino C11H10N2O2S 192 11.97 11.95 XIII Diacetyl-2-phenyl C13H12N2O3S 162 10.14 10.50 XIV 2-ethyl-2-p- tolylamino C12H19N2OS 124 11.97 11.92 XV cc -ethyl- oc-p-tolyl ;Q-phenylurea C16H18N2O 67 11.02 10.99 XVI 2-ethyl-2-p-tolyl- 5-benzal CigHisONsS 179 8.70 8.74 XVII 5-benzal-2- 4-thiazoldione C10H7NO2S 242 6.83 6.78 XVIII 2-p-tolvlimino- 3-ethyl C12H14N2OS oil 11.97 12.04 XIX 3-ethyl-2-4- thiazoldione C5II7NO2S 140 9.65 9.64 Dains et ALr. : Thiazolidones 23 4-Thiazolidone. Formula. M.p.°C. Percent nitrogen. No. Calcd. Found. XX 2-p-tolylimino- 3-ethyl-5-benzal C19H18X0OS 132 8.70 8.65 XXI 3-ethyl-5-benzal 2-4-thiazoldione Ci2HnN20S 97 6.01 6.10 XXII 2-benzyl-2- p-tolylamino C17H16N2OS 121 9.46 9.47 XXIII 2-p-nitrobenzyl- 2-p-tolylamino C17H15N3O3S 142 12.32 12.40 XXIV p-nitrobenzyl- p-tolylamine C14H14N2O0 67 11.57 11.52 XXV 2-ethyl-2-o tolylamino C12H14N2OS 98 11.97 11.66 XXVI 2-methyl-2-o- tolylamino C11H12N2OS 108 12.72 12.75 XXVII 2-o-tolylimino- 3-methyl C11H12N2OS ■ 59 12.72 12.82 XXVIII 2-o-tolyliniino- 3-methyl-5-benzal C18H16N2OS 129 9.09 8.83 XXIX 2-l)enzyl-2-o- tolylamino C17H16N2OS 95 9.46 9.58 XXX 2-benzyl-2-o- tolyiamino hydrochloride C17H17CIN2OS 179 8.42 8.21 24 The University Science Bulletin SUMMARY The aryl thiazolidones were readily prepared by the action of chloroacetic acid, chloroacetic ester or chloroacetyl chloride on the mono-aryl thioureas or by boiling the chloriacet aryl amide with potassium thiocyanate. Beckurts and Frerich had formulated these compounds as 2- imino-3-aryl-thiazolidones, but the work of Wheeler and Johnson and the investigations in this laboratory have proved that they are really 2-aryl amino or imino thiazolidones. In this paper (using derivatives of aniline and o and p-toluidine) it has been shown that the alkylation of the sodium salts gave mainly the 2-alkyl-2-aryl-amino thiazolidones (A) soluble in dilute acid and a small amount of the isomeric 2-aryl imino 3-alkyl com- pound (B) insoluble in dilute acid. The constitution of type A was proved bj^ synthesis from the unsymmetrical alkyl-aryl thio- ureas and by hydrolysis of the benzal derivative, which gave 5- benzal-2, 4-thiazolidone and secondary aryl-alkylamine. The structure of type B was shown by synthesis from the sym- metrical alkyl-aryl thioureas and the hydrolysis of the 5-benzal compound into a primary aryl amine and 3-alkyl-5-benzal-2, 4- thiazolidone. THE UNIVERSITY OP KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 3. On the Reactions of the Thiazolidones. VI: Some Derivatives of m-Toluidine F. B. DAINS, ROBERT D. COGHILL and SAMUEL S. TIHEN Contribution from the Chemical Laboratory of the University of Kansas SOME years ago Beckurts and Frerich (1) investigated the action of potassium thiocyanate on chloroacet-m-toluidide. Interpreting their results in the light of the work of Wheeler and Johnson (2), they obtained (a) the thiocyanate (m. p. 97°), (b) the "labile" form, i. e., the 2-imino-3-m-tolylthiazolidone (m. p. 136°), and (c) the stable 2-m-tolylamino-thiazolidone (m. p. 161°). To this latter (m. p. 161°), Beckurts and Frerich gave the 2-imino- 3-m-tolyl structure, basing their formula on the fact that on boiling the compound with 25% hydrochloric acid, 3-m-tolyl-2-4-thiazol- dione was produced. This does not of necessity prove the contention, since it has been found that the thiazolidone ring with an unsubstituted methylene grouping may open and close again after the loss of either ammonia or the aryl amine. Our own experiments confirmed the formulation of Wheeler and Johnson and proved definitely that the tolyl group was at position 2 and not 3. We found that the thiazolidone which was made both from mono-m-tolylthiourea and from chloroacet-m-toluidide was decidedly stable. Heating with various amounts of acid either gave no results or only m-toluidine. In no case were we able to isolate the 3-m-tolyl-thiazoldione (m. p. 90°). EXPERIMENTAL Sodium Salt. This was made by heating the thiazolidone with 10% sodium hy- droxide and filtering. From the cold solution the salt crystallized (1). Archiv. d. Pharm. 253. 233-65 (1915). (2). Wheeler and Johnson: Am. Chem. J. 28, 121-146 (1902). (25) 26 The University Science Bulletin out in good yield. This was filtered off at the pump, washed with a little cold water and dried for use. 2-ETHYL-2-m-TOLYLAMINO-THIAZOLIDONE I The sodium salt was refluxed in alcohol solution with ethyl iodide for several hours and then the alcohol and excess ethyl iodide re- moved by steam distillation. The oily residue was taken up in ether and the ether solution repeatedly extracted with 10% hydro- chloric acid. From the acid solution after neutralization with sodium carbonate was precipitated I, which, crystallized from heptane, melted at 106°. Beckurts and Frerich synthesized this compound (m. p. 106°) by a similar procedure, but had given to it the erroneous structure, 2-ethylimino-3-tolyl-thiazolidone, based evidently on their idea that the original thiazolidone was a 2-imino-3-tolyl compound. EVIDENCE FOR THE AMINO STRUCTURE FOR I Molar quantities of the thiazolidone I and benzaldehyde were dissolved in hot alcohol with the addition of a few drops of piperi- dine. From the solution there slowly crystallized the 2-ethyl-2-m- tolylamino-5-benzal-thiazolidone II fm. p. 185° from heptane). Hydrolysis. On long heating at 125° in 50% sulphuric acid. I was hydrolyzed into 5-benzal-2-4-thiazoldione (m. p. 243°) and ethyl- m-tolylamine. No evidence of m-toluidine was found. These re- sults showed that the ethyl and tolyl groups are joined to a nitrogen at position 2. 2-m-TOLYLIMINO-3-ETHYL-THIAZOLIDONE III The ether solution from which I had been extracted contained a small amount of an acid insoluble oil which failed to crystallize after months' standing. When its alcohol solution was treated with benzaldehyde and piperidine, there was obtained the same benzal- derivative IV (m. p. 127°) that was made from a synthetic specimen of III. oc-Ethyl-y8-m-tolylthiourea from m-tolylisothiocyanate and ethyl- amine, was heated in alcohol solution with ethyl chloroacetate and pyridine. From the reaction mixture was isolated III in 80% yield. The crystals from ether melted at 57°. This condensed with benzal- dehyde yielding the benzal-compound IV (m. p. 127°). Both specimens were identical as shown by a mixed melting point and analysis; thus proving that the 2-3-isomer was formed in small amount in the original reaction. Dains et al.: Reactions of Thiazolidoxes VI 27 HYDROLYSIS PRODUCTS OF III This was refluxed for twenty hours in an alcohol solution con- taining 3.5% hydrochloric acid. After evaporating off most of the solvent, the solution was made slightly alkaline and distilled with steam which carried over m-toluidine. The residual solution was concentrated and a solid crystallized out, containing no sulphur and melting at 110°. Analysis showed 24.35% and 24.40% nitrogen. Diethylurea melts at 109-10° and contains 24.15% nitrogen. There Avas no lowering of the melting point when mixed with a specimen of diethylurea made by disulphurizing diethylthiourea with yellow mercuric oxide. The diethylurea must be due to the intermediate formation of 3-ethyl-2-4-thiazoldione, described in the previous paper and then further hydrolysis into thioglycollic acid and ethyl carbonic acid and then diethylurea. Definite proof of the structure of III was afforded in the hy- drolysis of the benzal-compound IV. Heating with 50% sulfuric acid gave m-toluidine and 3-ethyl-5-benzal-2, 4-thiazoldione V (m. p. 97°). 2-METHYL-2-m-TOLYLAMINO-4-THIAZOLIDONE VI This was the acid soluble product when the sodium salt was methylated. The crystals from heptane melted at 104°. With the aid of piperidine it condensed with benzaldehyde giving the 5-benzal- derivatives VII (m. p. 173°). Heating with 50% sulfuric acid broke the ring of VII and there was isolated from the reaction product after making alkaline and distilling with steam, methyl-m-tolylamine. This was identified by its reaction product with phenylisocyanate, viz. : «= -methyl- cc-m- toIyl-yS-phenylurea (m. p. 76°). Anal. Calcd. for C,5Hi6N,0;N, 11.67. Found: 11.90. 2-m-TOLYLIMINO-3-METHYL THIAZOLIDONE VIII This was present in the original ether extract from which the acid soluble VI had been extracted. It failed to ciystallize, but its presence was proved by the formation of a benzal-derivative IX (m. p. 167°). oc-methyl-yff-m-tolylthiourea from methyl amine and m-tolyl iso- thiocyanate melted at 105°. N. Calcd. 15.53. Found: 15.65. This with ethyl chloroacetate gave a 70% yield of the 2-m-tolylimino-3- methyl-thiazolidone VIII. Crystals from ether, melting at 65°. The benzal-derivative from this melted at 167° and was identical with the one above (IX). 28 The University Science Bulletin Its constitution was proved and in consequence that of VIII, by its hydrolysis into m-toluidine and the known 3-methyl-5-benzal 2, 4-thiazoldione X. (m. p. 133°). DERIVATIVES OF TRIBROMO-m-TOLUIDINE (1-AMINO-3-METHYL-2, 4, 6-TRIBROMO-BENZENE) A satisfactory method for its preparation was to add to a dilute solution of m-toluidine hydrochloride. With constant stirring the calculated amount of bromine dissolved in acetic acid. It melted at 101°. Calcd. N, 4.07. Found: 3.82. CHLOROACET-TRIBROMO-m-TOLUIDIDE XI This was prepared in 70% yield by heating on a water bath a mixture of tribromo-m-toluidine moistened with alcohol and an excess of chloroacetyl chloride in a flask fitted with an air condenser. It was purified by recrystallization from benzene and then melted at 196°. When this was boiled in alcohol solution with potassium thio- cj^anate it gave a product which could not be purified completely. However, the sodium salt of the tribromo-tolyl-thiazolidone was ob- tained by treating the above product with 10% sodium hydroxide and filtering the hot solution. The sodium salt crystallized out on cooling. When this was treated with methyl iodide in the usual manner, a methyl derivative XII insoluble in dilute acid was ob- tained. It melted at 143° after repeated crystallizations from ben- zene. Lack of time prevented a more accurate determination of its constitution, but it is doubtless the 2-(tribromo-m-tolyl)imino-3- methyl thiazolidone. This is based upon its insolubility in dilute acid and its analogy to the behavior of 2, 4-diiodophenyl-thiazoli- done, which gave an 85% yield of the 2-ar5d-3-alkyl-thiazolidone and none of the 2-2 isomer. (3) For the sake of record, the following data are included in this paper. The results are incomplete owing to the departure of the junior authors from the Unive^sit3^ cc-n-Amyl-/?-o-Tolylthiourea XIII from amylamine and o-tolyl isothiocyanate, melted at 70°. The thiazolidone from this thiourea was an oil, but the 2-o-tolylimino-3-amyl-5-benzal thiazolidone XIV melted at 80°. °^-n-Amyl-^-m-tolylthiourea XV. This was made from m-anyl amine and m-tolyl isothiocyanate and when crystallized from ether, (3). John A. Davis and F. B. Dain.s : ,Iour. Am. Ch. Soc. 57, 2628 (1935). Dains et al. : Reactions of Thiazolidones VI 29 melted at 46°. The thiazolidone was also an oil, but its benzal de- rivative XVI melted at 68°. Only m-toluidine was isolated from its hj^drolysis products. The above sti-ucture is assigned from analogy in the behavior of symmetrical n-amyl-phenyl thiourea. (4) (4;. John A. Davis and F. B. Dains: Jour. Am. Chem. Soc. 57, 2629 (1935). 30 The University Science Bulletin Compound. Formula . M.p.°C. Percent nitrogen. No. Calcd. Found. I 2-Ethyl-2-m-tolyl- amino-thiazolidone C12H14N2OS 106 11.97 12.00 II 2-Ethyl-2-m-tolyl- amino-5-benzal- thiazolidone CioHisNoOS 185 8.70 8.76 III 2-m-Tolylimino-3- ethyl-thiazoUdone C12H14N2OS 57 11.97 11.90 IV 2-m-tolylimino-3- ethyl-5-benzal- thiazolidone C19H18N2OS 127 8.70 8.60 V 3-ethyl-5-benzal 2, 4-thiazoldione C12H11NO2S 97 6.01 5.85 VI 2-methyl-2-m-tolyl- amino thiazolidone C11H12N2OS 104 12.73 12.85 VII 2-methyl-2-m-tolyl- amino-5-benzal- thiazolidone CisHifiNoOS 173 9.10 8.99 VIII 2-m-tolyl-imino-3- methyl-thiazolidone C11H12N0OS 65 12.73 12.78 IX 2-m-tolylimino-3- methyl-5-benzal- thiazolidone C,sHi6N20S 167 9.10 9.14 X 3-methyl-5-benzal- 2, 4-thiazoldione CiiHoNOoS 133 6.39 6.40 XI Chloro-acet-+ri- bromo-m-toluidide CoHtBi-sCINO 196 3.33 3.21 XII 2-tribromo-m-tolyl- imino-3-methyl- thiazolidone CiiH9Br3N20S 143 6.12 5.87 XIII or. -n-Amyl-Q-o- tolyl thiourea C13H20N2S 70 11.87 11.96 XIV 2-o-Tolyl-imino- 3-n-a myl-5-benzal- thiazolidone C22H24N0OS 80 7.69 7.62 XV oc -n-amyl-^-m-tolyl- thiourca Ci3H2oN20S 46 11.87 11.98 XVI 2-m-tol yl-imi no- 3-n-amvl-5-benzal- t^hiazolidone C22H24N20S 68 7.69 7.77 Dains et al. : Reactions of Thiazolidones VI 31 SUMMARY This is a study of the constitution and reactions of 2-m-tolyl- amino-4-thiazolidone. Beckurts and Frerich had stated that this was a 2-imino-3-m-tohi-thiazolidone which our results have shown to be incorrect. In addition ethylation of the sodium salt gave both isomers. (A), the 2-ethyl-2-m-tolyl-thiazolidone, was the main product. Its ben- zal derivative was hydrolyzed into 5-benzal-2, 4-thiazoldione and ethyl-m-tolylamine. (B), the 2-m-tolyl-imino-3-ethyl-thiazolidone, was identified by means of its 5-benzal-compound which on hy- drolysis gave m-toluidine and 3-ethyl-5-benzal-thiazoldione. The methylation products of the sodium salt afforded corre- sponding results. The sodium salt of tribromo-m-tolyl thiazolidone gave almost entirely the 2-alkyl-3-aryl thiazolidone, thus showing the effect of the heavy tribromo radical. THE UNIVEESITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 4. On the Reactions of the Thiazolidones. VII: Some Derivatives of l-Amino-2-methyl-5- nitro benzene F. B. D.\INS and PAULINE GARBER Contribution from the Chemical Laboratory of the University of Kansas TWO thiazolidones, derived from amines in which the amino and nitro groups are meta to eacli other, viz., m-nitroaniline and 1- amino-3-nitro-4-methyl-benzene have been investigated by Beckurts and Frerich (1) and Walter S. Long and F. B. Dains (2). In each of these cases, on alkj'^lation of the sodium salt only the 2-aryl-2- alkyl-amino-thiazolidones were isolated and none of the 2-aryl- imino-3-alkyl isomers. In order to study the effect of the meta position, there has been chosen a derivative of o-toluidine in which the nitro group is meta to the amino and para to the methyl. Our results which are set forth in the following paper show that both isomers are formed on alkylation of the thiazolidone and that there is no inherent hindrance in such a meta position. EXPERIMENTAL The chloro-acet-nitro toluidide used was easily made as follows: A mixture of the nitro-toluidine (1 mol.) and pyridine (IVrs mols.) was dissolved in acetone in a flask cooled with ice water. Chloro- acetyl chloride (1 mol.) was added slowly. After standing for several hours, the contents of the flask were poured into water and the precipitate filtered, washed and dried. (1). Beckurts and Frerich, Archiv. d. Pharm. 253, 233-65 (1915). (2). Walter S. Long and F. B Dains: Trans. Kan. Acad. Sci. Vol. 16, 119-24 (1933). (33) 3—6037 34 The University Science Bulletin 2(5-NITRO-2-METHYL-PHENYL)-AMINO-4-THIAZOLIDONE I Molar quantities of the chloro-acet-compound and potassium thio- cyanate were boiled in alcohol solution for twenty hours — no effort being made to isolate any intermediate products. Water precipitated a crude mass which was purified by crystallization from glacial acetic acid. The brownish crystals melted at 186°. Analyses: Calcd. for C10H9N3O3: N, 16.74. Found: 16.25. Th.e ring was completely disrupted when I was heated with 50% sulfuric acid at 150° with the formation of ammonia and the nitro- toluidine (m. p. 110°). In the reaction flask a small amount of nitrotoluidine sulfate had crystallized out. It melted at 203° and contained 11.02% nitrogen (calcd. 11.2%). A preparation of the pure sulfate melted at 213-16°. 2(5-NITRO-2-METHYL PHENYL) -2-ETHYL-AMINO-4- THIAZOLIDONE II The sodium salt which was made by the action of hot 10% sodium hydroxide on I, was heated with ethyl iodide in alcohol solution for eight hours and then the excess ethyl iodide and alcohol removed by steam distillation. The gummy residue was dissolved in ether and the solution extracted several times with dilute hydrochloric acid. Neutralization of the acid solution gave a solid which was purified by recrystallization from heptane. The blunt yellow crystals melted at 85-7°. Analyses: Calcd. for C10H13O3N3S: N, 15.06. Found: 15.13. When the compound was hydrolyzed and the solution made alka- line, ammonia was set free and a precipitate formed which purified from alcohol formed yellow crystals melting at 81-2°. Analyses: Calcd. for C9Hi.,N20,: N, 15.56. Found: 15:86. This was {5-nitro-2-methyl-phenyl) -ethyl amine III, which W. MacCullum has reported (3) as melting at 81-2°. The formation of ammonia and the secondaiy amine thus proved the constitution of II. All efforts to condense the thiazolidone with benzaldehyde were unsuccessful. (3). W. MacCallum, J. Oh. Soc. 67.247 (1895). Dains et al.: Reactions of Thiazolidoxes VII 35 2-(5-XITRO-2-METHYL PHENYL) -IMINO-3-ETHYL-4- THIAZOLIDONE IV The ether residue from which II had been extracted was purified by crystallization from heptane. The yellow crystals, insoluble in dilute acid, melted at 124-5° and it is interesting to note that five times as much of the imino isomer IV was isolated as of the amino form II, Analyses: Calcd. Ci^H.aN.OsS: N, 15.06. Found: 15.29. On hydrolysis it gave the original nitrotoluidine (110°) and a volatile amine characterized by the carbylamine reaction, thus proving that the aryl and alkyl groups were joined to different nitrogens. Molar (luantities of this thiazolidone and benzaldehyde in hot alcohol solution condensed immediately on the addition of a few drops of dilute sodium hydroxide. The 2-nitrotolyl-imino-3-ethyl- 5-benzal-thiazolidone V melted at 140°. Analyses: Calcd. for CigHj^NgOgS: N, 11.44. Found: 11.10. The position of the two groups was indicated by the hydrolysis products: viz., the original nitrotoluidine and the known 3-ethyl-5- benzal-2, 4-thiazoldione (4) (m. p. 95°) VI. Analyses: Calcd. for C12H11NO2S: N, 6.00. Found: 6.27. 2-(5-NITRO-2-METH\T.-PHENYL)2-METmT-AMIXO-4- THIAZOLIDONE VII Methylation of the sodium salt gave a black tarrj'- residue which was taken up in ether. The acid extract on neutralization precipi- tated a product which cr^^stallized from heptane in long reddish- orange needles, melting at 101°. Analyses: Calcd. for C11H11N3O3S: N, 15.85. Found: 15.75. Like its analogue II, this also failed to condense with benzalde- hyde. From the hydrolysis reaction-mixture was isolated ammonia (but no alkyl amine) and the 5-nitrotolyl-methylamine VIII orange (4). F. B. Dains and Floyd A. Eberly: J. Am. Ch. Soc. 50. 3861 (1933). 36 The University Science Bulletin crystals from alcohol (m. p. 109-10°) (5). The melting point in literature was recorded as 107.5°. Analyses: Calcd. for C.,H,oN,0.: N, 16.88. Found: 17.07. 2-(5-NITRO-2-METHYL-PHENYL)-IMINO-3-METHYL-4- THIAZOLIDONE IX This was isolated in somewhat larger amount than VII from the original methylation product, since it was insoluble in dilute acid. The needles from heptane melted at 134.5°. Analyses: Calcd. for CiiH.iNoOsS: N, 15.85. Found: 15.72. It was broken down by heating with acid into methylamine and the nitrotoluidine (m. p. 110°). Its 5-benzal derivative, X, which was readily foraied, melted at 188°. Analyses: Calcd. for CigHi^NgOgS: N, 11.90. Found: 11.68. Incomplete experiments with benzyl chloride and the sodium salt indicated the formation of the two possible benzyl isomers. The 2-nitrotolyl-2-benzyl amino-thiazolidone melted at 160° and on hydrolysis gave the 2-mtrotolyl-benzyl amine (m. p. 124°) (6). SUMMARY Two derivatives of aryl thiazolidones, previously investigated, in which a nitro group was meta to the amino grouping, had failed to give the 2-arylimino-3-alkyl isomers. For this reason a study of the thiazolidone from l-amino-2-methyl-5-nitrobenzene was under- taken in order to ascertain whether there was any special hindrance in such a meta position. Our work has proved that the original compound has the structure 2-(5-nitro-2-methyl-phenyl) amino-4- thiazolidone and that the sodium salt on ethylation gave both the 2-nitrotolyl-2-ethyl (A) and in larger amount, showing the effect of the less basic grouping, the 2-nitrotolyl-imino-3-ethyl thiazolidone (B). Their constitution was shown by their hydrolysis product; A yielding ammonia and a secondary amin; B, an alkylamine and nitrotoluidine. It was noted that the 2-2-isomers failed to condense with benzaldehyde, while the 2-3 isomers easily formed benzal de- rivatives. (5). Gnehm and Plunier, Ann. 304, 99. (C). Ber. 35, 338 (1902). THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 5. On the Reactions of the Thiazoliclones. VIII: Some Derivatives of 2-Amino-5-nitro-toluene and 2-Amino-3-bromo-5-nitro-toluene F. B. DAINS and CLARENCE E. GROTHAUS IN the previous papers (1) it has been shown that the alkylation of the sodium salts of the 2-aryl-imino- or amino-thiazoli- dones gave usually a mixture of the 2-alkyl-2-aryl-amino- and the 3 - alkyl-2-arviimino-thiazolidones, S-C(NRR')N-COCH, and I ; I SC(NR)NR'COCH.>, the amount of each varying with nature of the I I aryl group. The results of our investigations show the inaccuracy of the in- terpretation of Beckurts and Frerich, that the alkylation of the sodium salts gave a 2-alkyl-imino-3-aryl-thiazolidone. They also confirm the work of Long (loc. cit.) and point out that the substi- tution of nitro, and bromo and nitro groups in the aryl nucleus in- creases the amount of the 3-alkyl isomers. EXPERIMENTAL PREPARATION OF 2-(4-NITRO-2-METHYL-PHENYLAMINO)- 4-THIAZOLIDONE (I), 12 3 4 5 SC ( NC6H3CH3NO2) NCOCH2 ! I This compound was made by refiuxing the chloracetyl derivative of 2-amino-5-nitro-toluene and an equivalent amount of potassium thiocyanate together in dilute alcohol solution. The thiazolidone melted at 172°. Beckurts and Frerich and Walter S. Long prepared (1). Beckurts and Frerich, Archiv. d. Pharni. 353, 233-65 (1915). Wheeler and Johnson, Am. Ch. J. 28, 121-35 (1902). Walter S. Long and F. B. Dains, Trans. Kan. Acad. Sci. SO. 119-24 (1933). F. B. Dains and Floyd A. Eberlv, J. A. C. S. 55, 3859 (1933). John A. Davis and F. B. Dains, J. A. C. S. 57, 2627 (1935). (37) 38 The University Science Bulletin this same compound similarly. Its preparation is described in this connection as it was used in this investigation as a starting material in the synthesis of several other derivatives. The sodium salt of this derivative was easily prepared by adding the thiazolidone to a hot 10% sodium hydroxide solution. On cool- ing, the sodium salt crystallized out. HYDROLYSIS OF 2-(4-NITRO-2-METm'L-PHENYLAMINO) -4-THIAZOLIDONE Five grams of the thiazolidone were heated with 50% sulfuric acid at 150° for three hours. When poured into water and neu- tralized with sodium hydroxide, a yellow precipitate was isolated which melted at ]26-127° after crystallization from alcohol. A mixed melting point proved this to be 2-amino-5-nitro-toluene. The alkaline solution was found to contain ammonia, formed evidently by breaking down of the heterocyclic ring. 2-(4-NITRO-2-METHYL-PHENYLAMINO)-5-BENZAL-4- THIAZOLIDONE (II), SCCNCtHcNOo) NCOC = CHC0H5 This derivative was prepared by treating the thiazolidone (5 grams) with benzaldehyde (4 c.c.) in alcoholic solution, to which a few drops of sodium hydroxide had been added. After standing several days the product precipitated out. Recrystallization from benzene yielded crystals melting at 230°. In attempting to hydrolyze the compound, no results were ob- tained using alcoholic sodium hydroxide. With 50% sulfuric acid, the derivative yielded 2-amino-5-nitro-toluene and an unidentified tarry residue. 2-(4-NITRO-2-METHYL-PHENYL)-2-METHYL-AMIXO-4- THIAZOLIDONE (III), SC(NCH3.C7H6N02) r^NCOCHo Twenty grams of the sodium salt of 2-(4-nitro-2-methyl-phenyl- amino-4-thiazolidone) were refluxed with methyl iodide for eight hours in alcoholic solution. The addition of water precipitated the product as a tarry mass which solidified to a grainy precipitate on standing. On extracting with 10% hydrochloric acid, about one half to two thirds of the product dissolved, reprecipitating on addition of ammonia. When crystallized from heptane, fine lemon yellow needles melting at 158° were formed. The structure was proved by Dains et al. : Reactions of Thiazolidones VIII 39 hydrolysis of the compound. The residue was shown to be 2-{4- nitro-2-methyl-phenylamino) -3-methyl-4-thiazolidone (see descrip- tion) . HYDROLYSIS OF 2-(4-NITRO-2-METHYL-PHENYL)-2-METHYL- AMINO-4-THIAZOLIDONE This was effected by heating to 150° for six hours in 50% sulfuric acid, the reaction product going into solution. On addition of sodium hydroxide, a yellow precipitate formed which melted at 137° when crystallized from alcohol. By a mixed melting point this was shown to be 4-nitro-2-methyl-phenyl methyl amine. The alkaline solution was distilled, a volatile base being carried over which was collected in hydrochloric acid and when analyzed proved to be ammonia. Since a secondary amine and ammonia were obtained, the stmcture of the compound is shown to be that assumed. If the methyl group were attached to a different nitrogen than the aryl group, methyl amine should have been obtained on hydrolysis instead of ammonia. 2-(4-NITRO-2-METHYL-PHENYL)-2-METHYL-AMINO-5- BENZAL-4-THIAZOLIDONE (IV), SC(NGH3C7H6N02)NCOC = CHC,;H.-. I ! This wv^s prepared by refluxing 2-(4-nitro-2-methyl-phenyl)-2- methyl-amino-4-thiazolidone with benzaldehyde in a solution of acetic acid and acetic anhydride. When crystallized from acetic acid or benzene, it melted at 238-240°. Attempts to hydrolyze in acid and alkali failed except that small quantities of 5-nitro-2-methylamino-toluene were isolated. 2-(4-NITRO-2-METHYL-PHENYLIMINO)-3-METHYL-4- THIAZOLIDONE (V), SC(XC7H,;N02)NCH3COCH2 The portion of the product from the methylation of 2-(4-nitro- 2-methyl-phenylamino)-4-thiazolidone, which was insoluble in hy- drochloric acid proved to be this compound. When crystallized from heptane, fine lemon colored crystals melting at 93° were obtained. Hydrolysis of the Derivative: This compound was hydrolyzed in the same manner as its isomer. 2-Amino-5-nitro-toluene was identi- fied as one of the products. The acid solution from hydrolysis was made strongly alkaline with sodium hydroxide and distilled. The 40 The University Science Bulletin distillate yielded methyl amine which was proved by the isonitrile test and analysis. These results indicated that the methyl group was in the three position on the heterocyclic ring. 2-(4-NITRO-2-METHYL-PHENYLIMINO)-3-METHYL-5- BENZAL-4-THIAZOLIDONE (VI) This was prepared by treating 2-(4-nitro-2-methyl-phenylimino')- 3-methyl-4-thiazolidone with bcnzaldehyde in the manner previously described. The small yellow crystals melted at 197°. 2- (4-NITRO-2-M ETH YL-PHEN YL ) -2-ETHYL AMINO-4- THIAZOLIDONE (VII) Our ethylation of the sodium salt of the nitrotolyl thiazolidone confirmed the work of Long, who obtained the 2- and 3-ethyl de- rivatives and proved their structure. Beckurts and Frerich (loc. cit.) had isolated the 2-nitrotolyl-2-ethyl-thiazolidone (m. p. 129°) but had assigned it to the erroneous structure 2-ethylimino-3-(p) nitrotolyl-4-thiazolidone. When this derivative was hydrolyzed with 50% sulfuric acid, 5-nitro-2-ethylamino-toluene melting at 98° was isolated and identi- fied. 2-(4-NITRO-2-METHYL-PHENYLIMINO)-3-ETHYL-4- THIAZOLIDONE (VIII) In the ethylation just described, the portion of the product which was insoluble in 10% hydrochloric acid was purified and found to melt at 86°. When hydrolyzed by heating with 50% sulfuric, 2-amino-5-nitro- toluene and ethyl amine were isolated and identified. These results indicate the compound to have the structure assumed. 2-(4-NITRO-2-METHYL-PHENYLIMINO)-3-BENZYL-4- THIAZOLIDONE (IX) This compound was obtained by refluxing the sodium salt of the monosubstituted thiazolidone with benzyl chloride (excess) in al- coholic solution for eight hours. The excess benzyl chloride was removed by steam distillation. None of the product dissolved in 10% hydrochloric acid and no evidence of the 2-benzyl isomer was obtained. It separates as small yellow needles melting at 98° when crystallized from a mixture of chloroform and ether. Hydrolysis: To effect hydrolysis, it was necessary to employ a 75% solution of sulfuric acid. The mixture was heated for six hours at 150°. From this solution was identified bv means of a mixed Dains et al.: Reactions of Thiazolidones VIII 41 melting point, 5-nitro-2-amino-toluene. A volatile amine was iso- lated from the solution which gave a positive isonitrile test and yielded a benzoyl derivative which melted at 102°. This benzoyl derivative of benzyl amine is recorded in the literature (2). These results substantiate the assumption that the benzyl group is attached to the nitrogen of the heterocyclic ring. 2-AMIXO-3-BROMO-5-NITRO-TOLUENE Since this product was used at the starting material for a series of compounds, it was made in considerable quantities. This was effected by suspending the nitro toluidine in water and slowly adding bromine with constant stirring. When crj'stallized from alcohol, it formed dark gold prisms, melting at 180°. This compound is reported in the literature (3). CHLORO-ACETYL DERIVATH^E OF 2-AMIXO-3-BROMO-0- XITRO-TOLUENE (X) Preparation of this compound was attempted by treating the toluidine with chloracetyl chloride in acetone and pyridine. When this failed, dry benzene was used instead of acetone without results. Finally the amine was heated directly with a large excess of chloro- acetyl chloride and the mixture heated to fuming at intervals. The mass was allowed to stand several days and then washed with water. It crystallized from alcohol in white crystals which melted at 202°. 2-(4-NITRO-6-BROMO-2-METHYL-PHENYLAMIXO)^- THIAZOLIDOXE (XI) 2-Methyl-4-nitro-6-bromo-chloracetanilide (50 grams) was treated with potassium thiocyanate (18 grams) in the usual manner, yield- ing the thiazolidone which melted at 223° when crystallized from alcohol. Hydrolysis: This was brought about by heating with 50% sul- furic acid. From the solution was isolated 2-amino-3-bromo-5- nitro-toluene. Ammonia was also found to be a product of hy- drolysis. 2-(6-BROMO-4-XITRO-2-METHl'L-PHEXYLIMIXO)-3-METHYL -4-THIAZOLIDOXE (XII) The sodium salt of the thiazolidone (XI) was treated with methyl iodide in the regular manner. The major part of the product was insoluble in dilute acid. It separated from benzene as light cream colored crystals, melting at 204°. (2). Blacher, Ber. 28, 424. (3). Ber. 13, 964 (1880). 42 The University Science Bulletin Hydrolysis: This was effected by heating a small portion of the compound in a sealed tube with 25 c. c. of 10% hydrochloric acid at 140° for eight hours. The hydrolysis yielded only one product, a volatile amine which could be identified. A good isonitrile test and its analysis proved it to be methyl amine. This substantiated the structure assumed as it is not probable that methyl amine would be obtained if the methyl group were attached to the other nitrogen. 2-(6-BROMO-4-NITRO-2-METHYL-PHENYLAMINO)-5-BEXZAL -4-THIAZOLIDONE (XIII) The thiazolidone {XI ) was heated with benzaldehyde in a solution of acetic acid and acetic anhydride and then allowed to stand for several days. It came down from acetic acid solution as fine fluffy white crystals which had not melted at 250°. 2-(6-BROMO-4-NITRO-2-METHYL-PHENYLIMINO)-3-ETHYL -4-THIAZOLIDONE (XIV) This derivative was made by refluxing the sodium salt of the thiazolidone (XI) with ethyl iodide. The product was found to be insoluble in dilute acid. The purified product from benzene melted at 164°. None of the 2-ethyl derivative was formed. Hydrolysis: This was effected in the same manner as the cor- responding methyl derivative. Ethyl amine was identified from the hydrolysis by the isonitrile test and analysis as the hydrochloride salt. 2-(6-BROMO-4-NITRO-2-METHYL-PHBNYLIMINO)-3-BENZYL -4-THIAZOLIDONE {X\) In this case the sodium salt of the thiazolidone (XI) was allowed to react with benzyl chloride. The compound was found to be in- soluble even in concentrated hydrochloric acid. It was crystallized from a mixture of benzene and heptane and melted at 172°. Hydrolysis: This hydrolysis was also carried out with 10% hydrochloric acid in a sealed tube. Benzyl amine was isolated as the hydrochloric salt and analyzed. This amine gave a positive isonitrile test. 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IZ Iz: 1^ ^ ^ ^ " ^ ^ V PQ « m « m ^ <-■ CO *-t IC rH O 2 S S m Tl 71 -^ ^ 05 -H -H — T-t rH X rH 1-* iH iH i O t U: HH W w ffi tr w K W w K w o t- - H c/: rH OC ^ 1 t- HH c iH I- W t- »-t t— » r-t 1-H H rH ^ "H iH {-S r- i rH 1— 1 rH C u o o 6 o c O C o o O o O O y-l e ^ o o a; O 3 Q 6 3 3 "o 'o Hi o 3 Q- c c N 0. c c s 'x. -J 'X rt 2 a, c c s S3 ■a r H c c c c C c c 1 .£ c SI .2 1 I .2 "o .2 1 irt< I Ti rfl -faH> a; S] .2 i >> d c •4^ c x> ^ '^ _2 ^ IS 1 0) « ^ X t 1 o ■rf- 1 ■^ 1 3 > a, CO 1 'c .£ _E t c5 1 :0 ^ d z P o a, S O 1 1 K C It; 1 ■c c 's c: 1 > £ V 1 Y £ C^ 1 'c limino)-2-methyl c C i 3 > a, 'c 1 c 'c C C '£ c c o ' 1' o A _g S c X. 1 1 T" 1 'o' g ■g ">> aj a. >> T ■> T 1 '> •> ">> '> ~> >> '> 1 r" ■> > s c rC X -£5 ^ e e ^ fl c cl c c p £ ^1-^ o a^ a) a c 0. Q. 03 a. 0, rC c s g a a C H ^ 1 c 1 1 U c L, C ' ^ 1 C<) "^ '^ 1 f-^ 1 1 1 I 1 > ■^ ^ 1 > ■) '>^ > ■^ T^ '> ■> > 1 > % CD C o 2 o 2 -c -c ,:= xi X rC X ^ -c C ;- M HH HH ■+J +i •*^ hh; • rH • f-l a "l 0. 0) a. (U 0. a. a. fr c '2 rH t>H fl £ E £ s e ^ £ 1 ^ r c 1 ^ ■* t 4 1^ 1 c| <=^ "^^ CJ =? — 1— I— 1— s > 1— 1 > > > >< y. 1— 1 X 1-H HH > HH > X 44 The University Science Bulletin SUMMARY The thiazolidones from (p)nitrotoluidme and the 6-bromo-(p)- nitrotoluidine and a number of other derivatives have been prepared. The methylation and ethylation of the (p)nitrotolyl thiazolidone gave the 2-alkyl-2-nitroaryl-amino and in larger amount the 2-aryl- imino-3-alkyl thiazolidones. However, with benzyl chloride only the 3-benzyl derivative was obtained. In the case of the bromo-nitro-tolyl thiazolidones from the action of methyl iodide, ethyl iodide, and benzyl chloride on the sodium salt, only the 3-methyl, 3-ethyl, and 3-benzyl isomers were isolated, thus showing the effect of the bromine and nitre substituents. THE UNIVERSITY OP KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 6. On the Alkylation of the 2-phenylimino-5-dimethyl- Thiazolidone and the 2-phenylimino-4-phenyl- Thiazoline FLOYD A. EBEIILY and F. B. DAINS Conlribution from the Chemical Laboratory of the University of Kansas IN a recent paper (1) a study was made of the 2-phenyliraino-4- diplienyl-thiazolidone in order to ascertain the effect of substi- tuting tlie methylene hydrogens at 4- with phenyl groups — and it was found that the compound behaved normally, yielding on alkyla- tion derivatives with alkyl groups at positions 2- and 3-. The fol- lowing paper records some additional observations on the analogous dimetiiylthiazolidone where the phenyl groups were replaced by methyl. In this case, however, only the 2-phenyl-2-methyl-amino- thiazolidone was isolated and none of the 3-methyl isomer. On the other hand, alkylation of the 2-piienylimino-4-phenyl-thiazoline gave only the 3-methyl compound and none of the 2-methyl isomer. EXPERIMENTAL PREPAR.4TI0N OF THE ETHYL ESTER OF ALPHA-BROMO ISOBUTYRIC ACm. (CH3)2CBrCOOC.>Hr, Isobutyric acid was converted into bromo-butyryl bromide on treatment with bromine and red phosphorus. Since the acid bromide could not be distilled without decomposi- tion even under reduced pressure, it was treated with absolute alcohol and converted into the ester. From sixty grams of this acid an eighty-gram yield of the ester boiling between 160-70° was obtained. (1). Ebcrly and Dains, Jour. Am. Chem. See, vol. 58, 2544, 1936. (45) 46 The University Science Bulletin 2-PHENYLIMINO (OR AMINO)-5-DIMETH\T.-4-THIAZOLIDONE, I SC(NC6H5)NHCOC(CH3)2 Monophenylthiourea (15 gms.), dimethyl-bromoacetic ester (20 gms.), and pyridine (8 gms.), in acetone (100 c. c), were refluxed for four hours with little effect ; the acetone was then replaced with benzene and the refiuxing continued for twelve hours. Removal of the benzene by steam distillation left a gummy res- idue. This was soluble in hot sodium hydroxide solution (10%) and on cooling, the crystalline sodium salt of the thiazolidone sep- arated. From a solution of the sodium salt, in hot water, the free base was precipitated by the addition of hydrochloric acid. Excess of acid dissolves the thiazolidone, but it can be reprecipitated from the acid solution by making it alkaline with ammonia. From alcohol it crystallized in colorless needles, melting at 172°. Analyses: Calcd. for C^Hi.N.OS: N, 12.72. Found: N, 12.87, 12.48. METHYLATION OF THE SODIUM SALT OF I. FORMATION OF 2-PHENYL-2-METHYL-AMINO-5-DIMETHYL THIAZOLIDONE II SC(NCoH5CH3)NCOC(CH3)2 I [ The dry sodium salt was heated with an excess of methyl iodide in absolute alcohol for several hours. Dilution of the reaction mix- ture with water and evaporation of the excess methyl iodide and alcohol gave an oil that slowly solidified. It was completely soluble in dilute acid, thus showing the absence of the 3-isomer. Crystallized from hot dilute alcohol, it formed colorless prisms that melted at 145°. Analyses: Calcd. for CioH^.N^OS: N, 11.97. Found: N, 12.05. Its structure was proven by its synthesis from unsymmetrical methyl-phenylthiourea. Equivalent amounts of the thiourea, ethyl bromoisobutyr&te and pyridine were refluxed in alcohol solution. From the reaction mixture was isolated the above thiazolidone II. 2-PHENYLIMINO-4-PHENYL-THIAZOLINE III SC(NCcHr,)NHC(CGH.-,)CH I I It will be noted that while this compound has the grouping SC(NC6H5)NH, which is found in the preceding thiazolidone, it differs in having no oxygen at 4- and a double bond between carbons Eberly and Dains: Alkylation of Thiazolidone 47 4 and 5. In consequence, the chemical behavior of the compound is distinctly modified. PREPARATION Twenty grams of phenacyl bromide were dissolved in alcohol and added slowly w^th constant shaking to a hot solution of monophenyl- thiourea in alcohol. The reaction was immediate, as indicated by the vigorous boiling of the alcohol, and was completed by heating for thirty minutes. The hydrobromide of the thiazoline separated on cooling; the free base was obtained by the addition of ammonia to the hot alcohol solution of the salt and dilution with water. The thiazoline is moderately soluble in hot alcohol, from which it deposits colorless crystals melting at 138°. Analyses: Calcd. for Cj^Hj^KS: N, 11.11. Found: N, 11.02, 11.07. The salts of this base with hydrochloric or hydrobromic acid are slightly soluble in hot water, but readily in hot alcohol. All at- tempts to prepare a sodium salt of this thiazoline analogous to those of the thiazolidones were fruitless. However, the results of methyla- tion and the action of carbon disulfide point to a 2-phenylimino structure. The work is in correlation with that of Traumann (2) who found that chloro-acetone or dichloro-ether reacted with monophenyl- thiourea giving an analogous 2-phenylimino-thiazoline and that of Naf (3) with dichloro ether and the monoalkjdthioureas. METHYLATIOX OF 2-PHEXYLIMINO-4-PHENYL-THIAZOLINE. FORMATION OF 2-PHENYLIMINO-3-IMETHYL -4-PHENYL THIAZOLINE IV SC . (NCcHr.) NCHsC . CgH-,-CH I [ Ten grams of the free base III and eight grams of methyl iodide were heated for five hours at 140-50°. Extraction of the reaction product with hot water and purification by crystallization from the same solvent gave a poor yield of the thiazoline IV. The plates melted at 171°. Analyses: Calcd. for C.eHi^N.S: N, 10.52. Found: N, 10.56. 10.43. Its structure was confirmed by its synthesis from symmetrical raethyl-phenylthiourea and phenacyl bromide. (2). Ann. 249, 31 C 1888. (3). N'af. Ann. 265, 127 (1891). 48 The University Science Bulletin In contradiction to the thiazolidones, the compound is extremely stable. It was not hydrolyzed by heating with 75% sulphuric acid or 20% hydrochloric acid at 160°. The presence of a phenylimino group at position 2- was indicated by the formation of phenyl isothiocyanate when the compound was heated with carbon disulphide at 190-200° for four hours. (4) 2-PHENYL-2-METHYL-4-PHENYL THIAZOLINE V SC(NCH3C6H5)NC(C6H5)CH This was obtained as a salt by the action of phenacyl bromide on unsymmetrical methyl-phenylthiourea in hot alcohol solution. The free base crystallized from alcohol in thick needles melting at 82°. Analyses: Calcd. for C.eHi^N^S: N, 10.52. Found: N, 10.37. This is the usual type of product resulting from the alkylation of sodium salt of the 2-phenylamino-thiazolidone, — but none was iso- lated in the methylation of the thiazoline. 2-PHENYLIMINO-3-ALLYL-4-PHENYL THIAZOLINE VI SC(NC(iH5)X(C3H5)C. (C6H5)CH Phenacyl bromide and symmetrical phenyl-allylthiourea were found to react with the formation of the above thiazoline which melted at 117°. Analyses: Calcd. for C^sH.gN.S: N, 9.55. Found: N, 9.35. Heating with carbon disulfide at 190-200° gave a nearly quanti- tative yield of phenyl mustard oil, thus showing the position of the phenyl and allyl groups. It is of interest to note that when the same allyl-phenyltliiourea was treated with chloroacetyl chloride, a 2- allylimino-3-phenyl thiazolidone was formed — the position of the groups in the two cases being reversed. SUMMARY The methylation of the 2-phenylimino-5-dimethyl-4-thiazolidone gave only the 2-phenyl-2-methyl-amino derivative with no evidence of the 3-methyl isomer. A sodium salt could not be obtained from the 2-phenylimino-4-phenyl thiazoline, but heating at 140° with meth}^ iodide gave in small yield the strongly basic 3-methyl de- rivative, whose constitution was proven by its synthesis from the (4). Grfifenliaspn, J. pr. Ch. (2) 75. 192. (1907). Eberly and Dains: Alkylation of Thiazolidone 49 symmetrical methyl-phenylthiourea. The position of the phenyl group was shown by the formation of phenyl isothiocyanate on heating with carbon disulphide. Symmetrical allyl-phenylthiomea and phenacyl bromide formed the 2-phenyl-3-allyl-thiazoline. In the previous paper it was found that the same thiourea and chioro-acetyl chloride gave a thiazolidone with the allyl group at 2- and the phenyl group at 3-, the positions thus being reversed. 4 -6o;{7 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15. 1936 [No. 7. Some Observations Regarding the Fate of Intrave- nously Injected Calcium Chloride A. M. LANDS and P. H. WOODARD, Department of Physiology, University of Kansas Abstract: The problem which the authors of this paper have investigated is the immediate fate of a soluble calcium salt following its injection into the blood stream of normal anesthetized dogs. The method of study is simi)ly the repeated determination of serum calcium at varying intervals of time over a total period of approximately two to three hours immediately fol- lowing the injection. Other workers are agreed that such injected calcium leaves the blood stream very rapidly at first and less rapidly later, but that approximatelj' the normal calcium level is attained in the time mentioned. Tliis finding is confirmed by the authors. They report that 69 percent leaves in the first five minutes and 95 percent has left in three hours. There has been some luck of agreement as to whether or not this disappearance is partly or wholly accounted for through excretion by the intestine and kidneys. Since the large intestine is believed to play the major role in excretion of calcium, tests were made to recover any excreted salt by washing out loops of the organ, but only small quantities were obtained, quite insufficient to account for the disappearance from the blood stream. Also, following the removal of the kidneys alone, the large intestine and kidneys, or the kidneys and both large and small intestine, the authors find still an early rapid disappearance of injected calcium from the blood. In case of the removal of the large intestine, however, the later disappearance of injected calcium is hindered somewhat. To test out the proposed idea that the bones are mainly responsible for the disappearance, animals were prepared in two waj's: (1) Insofar as pos- sible, the vessels supplying the bone were ligated, leaving only the soft tis- sues; and (2) heart -lung preparations were made. In both these cases, the disappearance curve closely resembled that of intact animals. These findings are discussed in connection with a theory proposed by Gre- ville that there is at first a rapid diffusion of calcium into the tissue spaces until some kind of an equilibrium is reached, after which the disappearance from the blood is slower. Perhaps this equilibrium involves some sort of binding of the calcium in an un-ionized form. It appears, however, that (51) 52 The University Science Bulletin neither excretion by the intestine or kidneys nor absorption by the bones plays any significant part in these processes. The work suggests that the large intestine may have a greater "binding" affinity for the injected calcium than some other tissues. THE calcium of whole blood is reported to be 5 to 9 mgm. percent (Lyman 1917, Kramer and Tisdall 1921, Jones 1921, Alport 1924), most if not all of which is found in the plasma (Abderhalden 1911, Leiboff.1930, Greville 1931, and others). The normal varia- tion of serum calcium is 9 to 12 mgm. percent, the value usually being nearer 10 mgm. percent (Lyman 1917, Matz 1925, Kramer and Tisdall 1921, Kramer and Rowland 1920, Watchorn 1924- '25, Esau and Stoland 1930, and many others). There is still consid- erable uncertainty as to the forms of calcium held in the blood stream. While that is not within the scope of the present discus- sion, it would seem that only a small portion of the total is ionized (Cushny 1920, Cameron and Moorhouse 1925, Richter-Quittner 1921, Brull 1930, and Benjamin and Hess 1933). A great deal of interest has been manifest in the fate of injected calcium salts and the restoration of the calcium balance of the body. Voorhoeve (1911) appears to have been one of the first to study the tissue calcium after the administration of calcium salts. He reports the administration of calcium (as the chloride or lactate, several grams per day) increased the calcium stock of the tissues and blood, even in well-fed adults. Kost (1918), who treated grow- ing young rabbits with calcium chloride for a week, found insig- nificant increases in the blood calcium and in the liver, somewhat more in the spleen, kidney and stomach, still more in the bone, muscle and intestine. Heubner (1918) found the only significant increase after calcium injection in the kidneys. Hetenyi (1924) could account for only half of the injected calcium in the blood and excretions at the end of three hours. After lethal doses he found the calcium content of the bones increased by 2 percent. Taylor and Fine (1930) could find no significant increase in the urinary excretion after calcium injection in cats nor could they recover any significant amount by washing out loops in the intestine. That injected calcium rapidly leaves the blood stream has been show^n by Heubner (1918), Hetenyi (1924), Salveson, Hastings and Mc- intosh (1924), Rothlin (1929), Taylor and Fine (1930), Gersch- mann (1930), and Greville (1931). While it is generally agreed that injected calcium rapidly leaves Lands and Woodard: Calcium Chloride 53 the blood stream, there is no general agreement regarding its fate. It occurred to the authors that more knowledge of the mechanism for disposing of an excess of calcium, produced by the intravenous injection of one of its soluble salts, might give some information regarding the normal metabolism of this ion. Much of the early work reported was carried out before adequate methods of calcium analysis had been devised. The purpose of the present investiga- tion was to study again carefully the immediate fate of intrave- nously injected calcium chloride, making use of recent advances in the technique of calcium. The work is divided into the following parts: 1. The rate of disappearance from the blood stream of the cal- cium excess following intravenous injection of calcium chloride in the normal animal. 2. The rate of disappearance of calcium excess after the removal of certain visceral organs which might be concerned in its dis- appearance. 3. The rate of disappearance of calcium excess in preparations containing only soft tissues. INIethods Adult dogs kept on the ordinary laboratory diet consisting of whole-wheat bread, milk and hamburger were used in these experi- ments. The animals were under ether anesthesia throughout the experimental period (90 to 180 minutes) ; the ether being admin- istered from an ether bottle through a tracheal cannula. One ca- rotid was cannulated to draw blood for analysis and the femoral vein exposed for injections. For the first part of the experiment the normal dogs were simply anesthetized and after securing a sample of normal blood the re- quired amount of calcium chloride was injected, following which samples of blood were drawn at stated intervals. The results of the analysis of these samples for calcium are recorded in Table I. In a second group of dogs a similar procedure was carried out with the exception that in each case the kidneys were removed after the withdrawal of the sample of normal blood. A third group was studied after both the large intestine and kidneys were removed and a fourth group following the removal of the small intestine, large intestines and both kidneys. Averages of these results are shown in Table I. Some data w^th respect to the relative impor- tance of the skeletal and soft parts of the body was obtained from studies of the disappearance rate of calcium from one group of dogs 54 The University Science Bulletin in which an effort was made to tie off most of the vessels leading to the bony skeleton. Results are shown in Table V. In a final group of animals the disappearance rate of calcium was studied from the heart-lung preparation. Results are shown in Table VI. table I. — Concentration in mg. calcium per 100 c.c. of blood and in percent of the amount injected at stated periods following the intravenous injection of calcium chloride (60 mgm. per kg. body weight) into normal and partly eviscerated animals. Time after injection of calcium chloride. Normal (average of 3 animals). After removal of kidneys (average 6 animals). After removal of large intestine and kidneys (average 3 animals). After removal of large and small intestines and kidneys (average 7 animals). Mg., 100 c.c. Percent injected. Mg., 100 c.c. Percent injection. 11.2 21.9 11.1 19.9 10.3 23.0 10.7 23.9 5 minutes .... 69 62 15 minutes .... 18.8 77 17.2 19.8 20,7 71 30 minutes .... 16.4 85 15.4 18.3 18.9 76 60 minutes. . . . 14.7 90 14.5 18.2 18.3 78 90 minutes .... 13.8 92 13.7 17.4 17.9 79 120 minutes. . . . 13.5 93 13.3 16.2 17.4 80 150 minutes .... 13.4 12.8 93 95 12.2 12.6 16.5 16.1 83 180 minutes .... 15.4 84 In all cases serum calcium was determined by the method of Clark and Collip (1925), a modification of the earlier method of Kramer and Tisdall (1931). The calcium is precipitated directly from clear blood serum by means of ammonium oxalate. The cal- cium oxalate precipitated is taken up in sulphuric acid and the oxa- late radical oxidized with a standard solution of potassium perman- ganate (n/100) . The amount of permanganate required to complete the oxidation of the oxalate carried down with the calcium enables the computation of the calcium present, one cubic centimeter of n/100 potassium permanganate being equivalent to 0.2 mgm. of calcium. At the present time this method is widely used for the determina- tion of blood calcium and with ordinary care has been found to give quite accurate results. Greville (1931) made a careful investiga- tion of this method. Calcium salts were added to horse serum and Lands and Woodard: Calcium Chloride 55 run in quadruplicate with a maximum error of less than 2 percent. Recoveries of 98.4 and 97.6 percent are reported with 0.576 mgm. and 0.289 mgm. of added calcium, respectively. The serum calcium of rabbits after the injection of calcium salts were determined both by Clark and Collip technique and the wet combustion method of Widmark and Vahlquist (1931). From this evidence it may be con- cluded that the method as modified by Clark and Collip is adequate and convenient for the determination of serum calcium. We see from the data (Table I), first column, that immediately after the injection of calcium salts the serum calcium is raised to high values (15 to 22.4 mgm. percent). Powers, Bowie and Howard (1930) using the congo red technique of Hooper, Smith and Whipple (1920) . found the blood volume of adult dogs to be 11.28 c.c. per 100 gms. body weight, the plasma volume being 6.18 c.c. Assuming this plasma volume, we find the calcium injected should give 45.2 mgm. percent if all of it were contained in the plasma. However, at the end of 5 minutes we find not 45.2 mgm. but 21.9 mgm. percent, 57 percent having disappeared from the plasma during the first five minutes after injection. Similar results were obtained by Heubner (1923) in the cat, by Hetenyi (1925) in humans, Rothlin (1929) in rabbits, Taylor and Fine (1930) in cats, Gerschmann (1930) in dogs and Greville (1931 ) in rabbits, all of whom report a rapid disap- pearance of intravenously injected calcium salts. Where has this calcium gone? Is it possible the corpuscles have taken up a part of the injected calcium? Abderhalden (1911) could find no calcium in the blood corpuscles of human blood. Guillaumin (1930) reports varying ciuantitics up to 2 mgm. per 100 c.c. of blood. Heubner and Rona (1919) found up to 7 mgm. percent in the cor- puscles following the injection of calcium salts. Greville (1931) found in rabbits, after the injection of calcium salts, the quantity of calcium carried down with or inside the corpuscles on centrifuging to be negligible. Results of determination made in this laboratory on whole blood, using the rapid combustion method of Bolliger (1932) compared with serum calcium determinations made by the method of Clark and Collip (19251, are in agreement with the find- ings of Greville (Table 11). A large portion of the injected calcium has left the blood stream. 56 The University Science Bulletin TABLE II. — Analysis of whole blood for calcium. Rapid combustion method of Bolliger Serum calcium by method of Clark and Collip., mgm. percent. Analj-sis of whole blood. Calcium in rngm. percent, assuming the total in the plasma. 10.2 9.8 17.4 17.7 22.8 24.0 23.2 24.0 22.9 21.5 Is the injected calcium excreted through the intestine and kidneys sufficiently rapid to account for its disappearance from the blood stream? That calcium is normally excreted by the kidneys and the epithelium of the large intestine has been well established by Rey (1894-'95), Patterson (1908), Cushny (1918), Nelson (1916), Hus- band, Godden and Richards (1923) , to mention only a few. Bergeim (1926) has definitely shown that in the rat there is an absorption of calcium along the small intestine and an excretion by the epithe- lium of the large intestine. Hetenyi (1924) believes calcium is nor- mally excreted, for the most part, by the large intestine, the ratio of large intestine to kidney excretion varying from 10 to 1, to 20 to 1. They state further this ratio may be influenced in favor of the kidneys bj^ many factors. Stewart and Percival (1927) found the calcium excretion by the large intestine of the cat to be 0.33 to 0.9 mgm. in a three-hour period. During the same period the kid- neys excreted 0.14 to 0.25 mgm. Walsh and Ivy (1927) studied calcium excretion in dogs with pouches (Thiry-Vella). Daily wash- ings of the upper jejunum gave 0.32 to 1.2 mgm. of calcium, lower ileum 2.0 to 8.5 mgm., the entire colon 1.8 to 15 mgm. daily. It is evident there are many difficulties to an accurate study of normal calcium excretion. The fecal calcium represents unab- sorbed calcium as well as excreted calcium. Bergeim (1926), as mentioned above, has shown both absorption and excretion take place in the intestine. There is little or no agreement as to the relative importance of these two processes in normal animals. Similarly there are many factors which complicate the study of urinary excretion. BruU (1930) found the excretion of calcium in the urine to be proportional to the rate of urine secretion, the ex- creted calcium acting like a "no-threshold" substance. Not only Lands and Woodard: Calcium Chloride 57 kidney activity but the composition of the diet may alter urinary calcium. Bogert and Kirkpatrick (1922) found base-forming diets caused a decrease in the urinary calcium and an increase in the fecal excretion, while the reverse was found true of acid-forming diets. Stehle (1917) found an increased excretion of calcium in the urine of dogs after the ingestion of hydrochloric acid. But M^hat of the excretion following the injection of calcium salts? Rey (1894-'95) stated that after intravenous injection of calcium salts 53 percent could be recovered from the colon in three days. Hetenyi (1924) found after intravenous injection of calcium salts slightly more fecal calcium than in control animals. There was little or no effect on urinary output of calcium. Salveson, Hastings and Macintosh (1924) found fecal excretion of calcium during the six days following a large injection of calcium chloride lower than that before the injection. Taylor and Fine (1930) studied calcium excretion in the cat. They observed that injected calcium chloride disappeared rapidly from the blood stream, but did not find any change in the excretion through the kidneys or intestine. They state further that evisceration or tying off the kidneys had no ef- fect on the rate of disappearance of calcium salts. In view of the rapid disappearance of injected calcium salts immediately after injection (60 to 70 percent during the first five minutes) and the general lack of agreement regarding the excretion- of calcium salts, the authors considered it worth while to reinvestigate the possibility of excretion playing a role in this rapid disappearance from the blood stream. In a series of six dogs, the kidneys were removed. All experi- ments were performed under conditions previously described. Cal- cium chloride (60 mg. per kilogram body weight) was injected in- travenously. Samples of blood were drawn at regular intervals and the serum analyzed for calcium. The data in Tabic I, second column, strongly suggests that the kidneys play a minor role, if any, in the removal of the injected calcium within the experimental period (three hours). The possi- bility of a compensatory increase in intestinal excretion after re- moval of the kidneys will be considered in another place. Hetenyi (1924) recovered 9 to 18 percent of the injected calcium from the urine after three hours. Brull (1930) found no increased output of calcium salts in the urine formed by a heart-lung-kidney prepa- ration. He believes the urinary output of calcium is dependent upon the rate of urine formation and not upon its concentration in 58 The University Science Bulletin the blood stream. Taylor and Fine (1930) injected calcium intra- venously in cats. They could not account for its disappearance through excretion by the kidneys. These observations justify the conclusion that the kidneys play no important role in the rapid dis- appearance of intravenously injected calcium salts. In three dogs the large intestine as well as the kidneys were removed. The averages of the serum calcium determinations are shown in the table. (Table I, column 3.) Examination of the data shows a consistently slower rate of dis- appearance in the animals with the large intestine and kidneys re- moved, this difference becoming more marked the longer the time after injection. As in groups previously reported, there is a rapid disappearance during the first few minutes following injection of calcium chloride. However, the slower process, the restoration of the calcium balance, is the portion most influenced. In some manner, the removal of the large intestine hinders the return of serum calcium to the normal level. This will be dis- cussed later. There is necessarily some loss of blood in the re- moval of a part of the intestinal tract. In these experiments that was not considerable. To determine the effect of this on the rate of disappearance of injected calcium the following experiment was run : A dog was anesthetized and prepared for injection and bleeding as previously described. Before injecting calcium chloride the ani- mal was bled 250 c. c. of blood and then a disappearance of injected calcium determined as previously described. The following results were obtained: TABLE III Time, in minutes, follow- Serum Ca. IVIgm. ing injection percent N 11.3 5 23.4 30 17.5 50 16.1 60 15.9 90 15.0 120 14.5 150 13.7 180 12.7 Using the values for blood volume of the dog found by Powers, Bowie and Howard (1930), 250 c.c. in this animal represents 17.8 percent of the circulating volume. Inspection of the above data shows the disappearance of calcium to be normal. It is unlikety that in any evisceration experiments here reported there was a loss of blood as great as that reported here. While it is impossible to Lands and Woodard: Calcium Chloride 59 say if impaired circulation may not have been in part responsible for the slower disappearance from the blood stream, yet it may be pointed out that the exposure and manipulation of the viscera in removing the kidneys did not alter the rate of disappearance. Is the difference in the rate of disappearance a general result of the loss of tissue or a specific effect from removal of the large in- testine? Would the removal of the small intestine as well as the large intestine alter this rate of disappearance further? To give some information regarding this possibility the small intestine was removed (up to the tail of the pancreas) as well as the kidneys and large intestines. The averages of serum calcium after the injec- tion of calcium chloride are given in the table. (Table I, column 4.) The rate of disappearance of injected calcium here is slightly slower than that reported in column 3. Considering the mass of tissue removed with the removal of the small intestine, it would seem there is a great difference in the affinity of tissues for calcium if the difference observed is due to some tissue "binding" of calcium. There are three rather obvious possibilities to be considered: 1. That the intestines excrete a portion of the calcium excess and that this excretion is considerable immediately after injection. 2. That these tissues take up and hold a portion of the calcium excess. 3. That both 1 and 2 occur. We will consider the first of these possibilities in the next series of experiments to be reported. As wc see from an earlier review of the literature on calcium ex- cretion by the intestine, increases in the rate of excretion of calcium following the injection of one of its soluble salts was reported by Rey (1894- '95) and by Hetenyi (1924) more recently. Stewart and Percival (1929) studied the normal calcium excre- tion in cats by isolation of segments of the intestine and washing out the excreted material at intervals. Similar studies were made by Walsh and Ivy (1927- '28) in the dog using Thiry-Vella pouches. Both report a small normal excretion. Taylor and Fine (cf. above) using the technique of Stewart and Percival, studied the excretion of calcium in cats after the intravenous injection of calcium chlo- ride. They state, "The disappearance cannot be accounted for by excretion through the bowels or kidneys." They believe the in- jected calcium does not leave the body. Since the removal of the 60 The University Science Bulletin intestines does alter the rate of disappearance of injected calcium salts, it is imperative that the possibility of their excretion be studied. The method of study used here is essentially the same as that used by Taylor and Fine. The kidneys were removed and the large intestine cannulated, carefully cleaned of all contents and washed with warm physiological saline until absolutely clear. A last por- tion of the washings was saved and analyzed for calcium. After three hours from the time of calcium injection (60 mgm. CaClg per kilogram body weight was injected in all cases) the large intestine was again washed out with 500 c.c. of warm physiological saline, the saline being run through the intestine three times. Ten cubic centi- meter portions of the clear washings were taken for analysis. To this was added 2 c.c. of 4 percent ammonium oxalate and after standing 24 hours in the ice box centrifuged and titrated according to the technique of Clark and Collip (previously described). The results from all dogs in which calcium excretion was studied are shown in tabular form below. (Table IV.) TABLE IV. — The excretion of intravenously injected calcium by the epithelium of the large intestine. Recovery experiments Before injecting calcium. Serum calcium. Calcium excreted, large intestine, in 180 minutes. Number of Experiment. 5 minutes after injecting calcium. 180 minutes after injecting calcium. 27 29 33 mgm. 11.8 12.6 10.3 10.4 11.2 Control mgm. 21.4 21.5 23.0 18.9 16.3 ingm. 13.3 14.8 mgm. 7.6 1.4 2 0 37 11.8 14.3 1.43 38 2.8 44 Trace Walsh and Ivy (1927-'28) have shown that repeated washing increases the calcium ex- creted into the large intestine. In the experiments here reported, the calcium excreted was calculated by substracting the amount in the last portion of the first washing from the amount obtained after three hours, on the assumption that this represents the amount excreted due to the washing out. While these values are a little higher than those of Taylor and Fine (1930) who reports 0.77 mgm. in one experiment on a normal dog (studied excretion in the cat) yet they are in agreement in that they show only a small amount of the injected calcium has been excreted in the experimental period. Examination of Table I shows Lands and Woodard: Calcium Chloride 61 a difference in serum calcium between normal dogs and those in which the kidneys and large intestine had been removed of 2.6 mgm. percent. In the case of a dog of average size (12 kilograms) there should be about 741 c.c. of plasma which would contain, after in- jection, a total of 19.2 mgm. of calcium more than the normal amount. This represents the loss attributable to the large intestine. We see that the greatest recovery reported here is 7.6 mgm. with 3.06 mgm. as an average value. If the intestine excretes as much as 19.2 mgm. of calcium it was not recovered from the washings in our experiments. The small amount excreted in our experiments is wholly inadequate to explain the rapid disappearance that oc- curs during the first five minutes after injection. Since the kidneys were removed in these experiments, we must conclude a large por- tion of the injected calcium is retained in the body. Since the calcium leaves the blood stream and since it is retained in the body we have the possibility of: 1. Diffusion into the soft tissues. 2. Deposition in the bones. 3. Both diffusion into the soft tissues and deposition in the bones. Greville (1931) suggested that both processes do occur. He be- lieves that there is first a rapid diffusion into the tissue spaces until there is some form of equilibrium between the calcium in these spaces and the raised blood calcium. The calcium then disappears from the tissue spaces and blood stream, the rate of disappearance at any given instant being proportional to the excess present. If this be correct, the first part of the disappearance curve would take place in the complete absence of the skeletal system. The amount of calcium leaving the blood stream would be proportional to the amount of calcium injected and the amount of soft tissues taking part in the reaction. The next series of experiments will consider this possibility. In all dogs used in these experiments the thorax was opened under ether anesthesia and the animal kept alive by artificial respira- tion. The blood vessels to the extremities, all vessels to the head except the internal carotids and all vessels to the dorsal body wall were tied off. The intercostals were ligated from the opened thorax. The vessels to dorsal abdominal wall were exposed by an incision into the abdominal cavity through the dorsal wall. These were then dissected from other tissue near their origin from the ab- dominal aorta and ligated. This procedure was found to disturb the abdominal viscera the least. 62 The University Science Bulletin Calcium chloride in a 5 percent solution was injected into the jugular or subclavian vein near the heart. Blood was drawn at regular intervals thereafter and analyzed for calcium as previously described. Table V gives the results of seven experiments in which the above technicjue was employed. It is obvious from the examination of this data that there is a rapid disappearance of injected calcium from the blood stream during the first five minutes after injection and that the bony tissues play no role in this. In the past it was generally believed calcium salts decreased the permeability of the capillary wall. That this is not the case has been shown by Yana- gawa (1916), Bayliss (1918) and Smith and Mendel (1920). That calcium does penetrate the wall of the capillaries has been shown by Arnold and Mendel, who demonstrated its appearance in oedema fluid and serous transudates. Thus, there is little doubt but that table v. — The disappearance of intravenously injected calcium chloride after tying off the blood vessels to the skeleton (including some skeletal muscle) Number of Experiment. Calcium injected per 100 c.c. plasma, mgm. Calcium in mg. percent 5 minutes after injection. Calcium leaving blood, in percent injected. 5 minutes. 30 minutes. 60 minutes. 180 minutes. 36 66 55 47 50 38 45 50 25.4 19.0 27.8 31.8 28.2 30.5 38.0 77 83 64 57 55 55 44 78 40 87 41 78 43 72 76 45.. 58 (145 min.) 40 69 48 50 there is a rapid diffusion out into the tissue spaces of a large amount of the injected calcium. It would seem that during the first few minutes some equilibrium is reached between the blood and tissues after which only slow changes in the concentration to- ward normal occur. Certainly the nature of these processes whereby calcium is taken out of circulation and held by the soft tissues in- vites more study. This study is only a beginning in the study of tissue calcium. Future work must deal with the problems of the normal content of various tissues and the tissue spaces, the forms in which it is held in the tissues and the role it plays in normal cell metabolism. Lands and Woodard: Calcium Chloride 63 Having shown that there was a rapid disappearance of injected calcium in preparations containing only soft tissues, it would be interesting to see if a similar thing would occur in preparations containing only a few organs. The heart-lung preparation of Star- ling offers an ideal opportunity to do just that. The data from three satisfactory preparations is given in the table below. (Table VI.) T.\BLE VI. — The disappearance of calcium added as calcium chloride to a Starling heart-lung preparation NUMHEK OF EXPEKIMENT. 28 32 53 Serum Percent Calcium Normal calcium injected injected serum m mgm. calcium per calcium percent leaving 100 c.c. m mgm. 5 minutes blood plasma. percent . after first 5 injection. n)inutes. ingm. mgm. mgm. 43 7.8 33.8 39.5 28.8 8.4 23.0 49.3 42.0 8.0 27.0 54.7 At other periods, calcium in mgm. percent. After 4.J mill., 2S nism. After 75 min., 20.0 ingni. After 53 min., IS. 2 ingni. A comparison of the disappearance curves obtained in normal dogs with those in which the skeleton had been tied off from the circulation and with the heart-lung jireparations shows in general the same contour, particularly during the first few minutes after injection. There is first a rapid fall immediately after the injection of calcium chloride and then a very slow decrease. It will be seen from an examination of both Tables V and VI that there is still some further loss of blood calcium after the first few minutes, al- though this takes place more slowly. In experiment 53 (heart- lung) there was a loss of 8.8 mgm. between the 5th and 53d min- utes and in number 43 (soft tissue preparations) there was a loss of 3.8 mgm. between the 5th and 145th minutes. While it has been shown that the bone trabcculae stores calcium on a high calcium diet (Bauer, Aub and Albright 1929) this process may be much slower than the rate of disappearance here observed. These ex- periments show that there is a rapid loss of injected calcium salts from the blood stream, this taking place in the complete absence of bone tissue. The data here reported is in general agreement with the hy- pothesis of Greville (1931), who suggested, on the basis of the normal disappearance curve, that there is a rapid diffusion into the tissue spaces until some kind of an equilibrium is reached between these and the blood stream. It may be further stated, that there is a 64 The University Science Bulletin rapid diffusion into the tissue spaces until some kind of an equilib- rium is reached between these and the blood stream. It may be further stated, that there is a rapid diffusion into the tissue spaces immediately after injecting a hypertonic calcium chloride solution followed by some slower process which is capable of further reduc- ing the blood calcium below the level resulting from the rapid dif- fusion into the tissue spaces. Just what this process is, it is not possible at the present time to say. It does not involve the skele- ton, although the bones may be necessary before calcium equilib- rium can be regained. Our knowledge of the calcium content of the soft tissues is all too meager. That it is very variable was first shown by Parhon, Dumitresco and Nissipesco (1909) and since confirmed by Rona and Heubner (1918). The calcium content of the tissues was re- ported increased following high calcium intake by Voorhoeve (1911). Similar findings are reported by Kost (1918) for some organs (blood, liver, spleen, kidney and intestine). Huebner and Rona (1923) made a careful analysis of the tissues of cats for calcium after chronic and acute poisoning with calcium chloride. They could find no significant increase in their calcium content. How- ever, since their animals were not killed for analysis until 2 to 18 hours after the last calcium injection, it is possible that the calcium equilibrium of the tissues had been restored to or near normal. Hecht (1924) and Jungmann and Samter (1924) report similar findings after calcium chloride, acetate or hexose phosphate. Yet, examination of their data shows increases in the skin in some in- stances, large amounts of calcium in the small intestine and the kidneys, particularly after the injection of calcium hexose phos- phate. Hetenyi (1924) believes he can account for 78 percent of the injected calcium in the bones at the end of three hours. There is no general agreement regarding normal calcium content of the tissues or the effect of injection of calcium on that content. Hav- ing demonstrated the rapid disappearance of the calcium excess into the soft tissues from the blood stream, a preliminary study was made of the calcium content of some of these in an attempt to de- tect this increase w^hich must have occurred. Method Various methods of tissue analysis for calcium have been pub- lished. A few of the more recent ones are: Jansen (1918), Barral and Barral (1928) and Widmark and Vahlquist (1931). Most of the earlier methods required incineration to destroy the organic Lands and Woodard: Calcium Chloride 65 material. Widmark and Vahlquist (cf. above), however, devised a technique of analysis in which the destruction of the organic material was carried out in solution by the addition of sulphuric and nitric acids. Calcium was determined by the ordinary ox- alate-permanganate method after the complete oxidation of the or- ganic material. The method used in this laboratory makes use of the Bolliger (1932) technique for the rapid digestion of bio- logical materials. The material (usually 4 to 7 gms.) is oxidized with perchloric and nitric acids until practically colorless, neutral- ized to methyl red with ammonium hydroxide and made up to 25 c.c. volume. To a 10 c.c. portion in a conical centrifuge tube, 2 c.c. of 4 percent ammonium oxalate is added, mixed and allowed to stand over night. This is then centrifuged for 15 minutes at about 2,000 revolutions per minute. After washing (cf. the method of Clark and Collip, 1925) the calcium oxalate is dissolved in 3 c.c. of approximately normal sulphuric acid and titrated with n/100 po- tassium permanganate solution, each cubic centimeter of this being equivalent to 0.2 mgm. of calcium. A few studies were carried out to determine the calcium content of both normal tissues and those following the injection of calcium chloride. However, such wide variations were found in normal values that, considering tlie amounts of calcium which were injected, the data obtained following calcium injection was not very significant. It suggests that there is somewhat greater accumulation of calcium in certain organs, viz.: the heart, lungs and small and large intestine. Examination of Table I, column 3, shows that the removal of the alimentaiy tract below the pancreas greatly interferes with the re- turn of the serum calcium to normal levels although there appears to be a rapid diffusion from the blood stream during the first few min- utes following injection (62 percent of the injected calcium being removed during the first few minutes following injection, which com- pares favorably with controls in which 69 percent was found to have left the blood stream during the same period) . Whereas, in the intact animal 95 percent had left during the first three hours, in those without kidneys or alimentary tract below the pancreas only 84 percent had been removed from the circulation. Analysis of the large and small intestines for calcium in the experiments here re- ported suggest the calcium content of these may be significantly higher thirty minutes after injection (when the experiments were terminated). "While more study is required on this point, yet there is a suggestion from this and previous data given that these organs 5—6037 66 The University Science Bulletin are capable of "binding" a portion of the excess injected and thus holding it until it can be stored in the bones or excreted. It would thus be rendered biologically inactive (nonionized and nondifTusi- ble). It is not improbable that this exists in some equilibrium with the blood stream so that its elimination by storage in the bones or excretion will gradually restore the calcium content of the soft tissues to normal. Discussion and Summary Benjamin and Hess (1933) in a recently published article suggest there are four forms of calcium in the blood stream, two diffusible and two nondiffusible. Of the diffusible calcium, about two thirds is in the form of an adsorbable calcium-phosphorus complex; the re- mainder contains the calcium ion. Of the nondiffusible calcium, about one quarter is adsorbable and the remainder is probably bound to the plasma proteins. In hypercalcemia induced by the sub- cutaneous injection of calcium lactate, there was an increase in the serum calcium from a normal 11.0-11.6 mgm. percent to 15.3 mgm. percent. The protein bound fraction was not significantly altered (normal 4.3 mgm. percent was increased to 4.6 mgm. percent), but there was an increase in the adsorbable complex from a normal of 3.9-4.9 mgm. percent to 7.2 mgm. percent. The supposedly ionized portion was increased from a normal value of 2.9 to 3.5 mgm. per- cent. Thus we see there is a disproportionately large formation of some nonionized calcium complex following the injection of a solu- ble calcium salt. Brull (1930), making use of a vivi-diffusion tech- nique, finds a similar normal partition in the blood stream. He finds a large portion of the calcium added as calcium chloride rapidly be- comes nondiffusible. He further shows that the addition of an ionized salt of calcium to the heart-lung preparation causes an im- mediate reduction of systolic and diastolic volume and an increase in the duration of the contractions, results which would be expected from many experiments which have been performed on the heart of lower animals. However, this effect rapidly disappears and normal heart action is restored. (We have noticed a similar response in our heart-lung experiments.) Since no such accommodation as this is known to occur with the isolated heart, it suggests that the injected calcium is rapidly deionized by the organism. Miss Benjamin (cf. above) has found such a form and has shown that it is increased after the injection of calcium lactate, increased more than other forms of calcium found in the blood stream. Lands and Woodard: Calcium Chloride 67 Thus we are led to construct the following picture of the events which occur following the injection of calcium chloride: There is a rapid diffusion of the injected calcium into the tissue spaces accom- panied by the deionization of the injected calcium, possibly part of the deionized portion being represented by the calcium complex of Benjamin and Hess. Excretion plays little or no role in the rapid disappearance observed and it would seem from the work of others (Rey 1894-'95, Hetenyi 1924, Salveson, Hastings and Macintosh 1924, Taylor and Fine 1930, and others) that a portion may be re- tained for several days after injection. Brull (1930) suggests there is an excretable form of calcium (not ionized). Excretion would then depend upon the production of this form rather than the pres- ence of an excess calcium in the body fluids. After this rapid diffu- sion and the resultant equilibrium the calcium content of the blood and tissue spaces is considerably higher than normal. Reactions of a slower nature then become apparent — reactions which are capable of disposing of this excess. That bone, particularly cancellous bone, does take up and hold a portion of the excess calcium absorbed on a high calcium diet has been demonstrated in a series of clever ex- periments carried out by Bauer, Aub and Albright (1929). This suggests itself as the logical site for the storage of excess calcium removed from the blood and tissues, although a small amount may be excreted. Since calcium disappearance curves obtained from animals containing only soft tissues demonstrate both the rapid diffusion and the slower decrease in calcium which becomes ap- parent after this rapid diffusion, the deposition in the bones would appear to be the terminal phase of a complex reaction in which the tissues probably play an important role. SUMMARY 1. Intravenously injected calcium chloride rapidly leaves the blood stream. Following the intravenous injection of 60 mgm. of this salt per kilogram body weight as a 5 percent solution, 69 per- cent was found to have left the blood stream during the first five minutes and 95 percent had left within 3 hours. 2. This rapid disappearance does not depend upon excretion, since it occurs in the absence of the intestine and kidneys. This is further confirmed by attempts to recover excreted calcium from the large intestine. Only small amounts could be obtained 3 hours after injection. 68 The University Science Bulletin 3. Removal of the large intestine in some way retards the res- toration of the normal level of serum calcium after the injection of calcium chloride. The probable significance of this is discussed. 4. The rapid disappearance of injected calcium chloride observed in the intact animal can be observed in preparations containing no bone tissue. It is concluded that this reaction depends upon the blood and soft tissues alone, although the skeleton may be necessary for the restoration of the normal level of calcium in the blood and tissues. 5. Some preliminary analyses of various organs for calcium fol- lowing the intravenous injection of calcium chloride lend further support to the concept that a large amount of the calcium excess may have diffused into the tissue spaces. BIBLIOGRAPHY Abderhalden. 1911. Qjjoted by Leiboff. Alport. 1924. Biochem. Journ., 18, 455. B.'iRRAL AND Barral. 1928. Compt. rend. Soc. Biol., 99, 519. Bayliss. 1918. Quoted by Greville. Bauer, Aub and Albright. 1929. Journ. Exper. Med., 49, 145. Benjamin and Hess. 1933. Joura, Biol. Chem. 100, 27. Bergeim. 1926. Journ. Biol. Chem., 70, 35; 51. BoGERT AND KiRKPATRicK. 1922. Journ. Biol. Chem., 54, 375. Bollinger. 1932. Australian Journ. of Exper. Biol. Med. Sc, 10. 57. Brull. 1930. Arch. Intern. Physiol., 32, 138. Cameron and Moorhouse. 1925. Journ. Biol. Chem., 63, 687. Clark and Collip. 1925. Journ. Biol. Chem., 63, 461. CusHNY. 1920. Journ. Physiol., 53, 391. CusHNY. 1918. Textbook of Pharmacology and Therapeutics, page 560. Esau and Stoland. 1930. Amer. Journ. Physiol., 92, 25. FoHSTEE. 1837. Zeitschr. Biol., 9, 297; 381; ibid., 1876, 12, 464. Gerschmann. 1930. Compt. lend. Soc. Biol., 104, 413. Greville. 1931. Biochem. Journ., 25, 1931. Goitein. 1906. Arch. F. D. ges. Physiol., 115, 118. Guillaumin. 1930. Bull. Soc. Chem. Biol., 12, 1269. Hecht. 1924. Biochem. Zeitschr., 144, 270. Hetenyi. 1924. Zeitschr. f . d. ges. Exp. Med., 43, 1924. Heubner and Rona. 1918. Biochem. Zeitschr., 93, 187. 1923, Ibid., 135, 248. Hooper, Smith and Whipple. 1920. Amer. Journ. Physiol., 51, 205. Husband, Godden and Richards. 1923. Biochem. Journ., 17, 707. Jones. 1921. Journ. Biol. Chem., 49, 187. Jungmann and Samter. 1924. Biochem. Zeitschr., 144, 265. KosT. 1913. Quoted by Heubner and Rona. (1918). Kramer and Tisdall. 1921. Journ. Biol. Chem., 47, 475. Kramer and Howland. 1920. Joura. Biol. Chem., 43, 35. Leiboff. 1930. Journ. Biol. Chem., 85, 759. Lands and Woodard: Calcium Chloride 69 Locke. 1894. Centralblatt Physiol., 8, 166. LoEB. 1903. Pflug. Arch., 97, 394. Lyman. 1917. Joum. Biol. Chem., 30, 1. Matz. 1925. Amer. Rev. Tuberculosis, 11. Mendel and Benedict. 1909-10. Amer. Journ. Physiol., 25, 23. Nelson and Burns. 1916. Journ. Biol. Chem., 28, 231. Nelson and Willl\ms. 1916. Journ. Biol. Chem., 28, 237. Parhon Dumitrescx) and Nissipesco. 1909. Compt. rend. Soc. Biol., 66, 792. Pettbnkofer and Voit. 1866. Zeitschr. Biol., 2, 459. Powers, Bowie and Howard. 1930. Amer. Journ. Physiol., 92, 665. Renvall. 1904. Skand. Arch. f. Physiol., 16, 114. Rey. 1895. Arch. f. exper. Path. u. Pharm., 35, 298. Richter-Qnittner. 1921. Biochem. Zeitschr., 124, 106. 1921-'22 Ibid., 126, 97. RoNA and Hextbner. 1918. Biochem. Zeit.schr., 93, 353. Rothlin. 1929. Zeitschr. f. d. ges. exp. Med., 70, 634. RuDEL. 1894. Arch. f. exper. Path. u. Pharm.. 33. 87. S,\LVES0N, Hastings and MacIntosh. 1924. Journ. Biol. Chem., 60, 327. Smith and Mexdel. 1920. Amer. Journ. Physiol., 53, 323. Stehle. 1917. Journ. Biol. Chem., 31, 461. Stewart and Perctval. 1927. Biochem. Journ., 21, 301. Taylor and Fine. 1930. Amer. Journ. Physiol., 93, 544. Tei.fer. 1922. Quart. Journ. Med., 16, 45. Tereg and Arnold. 1883. Arch. f. d. ges. Physiol., 32, 122. Voit. 1880. Zeitschr. Biol., 16, 55. Voorhoeve. 1911. Dcut. Arch. Klin. Med., 110, 461. Walsh and Ivy. 1927-28. Proc. Soc. Exper. Biol. Med., 25, 839. W.ATCHORN. 1924-'25. Quart. Journ. Med., 18, 288. Weiske. 1871. Zeitschr. Biol., 7, 179; 333. Weiske-Proskau. 1872. Ibid. 8, 239. ■\A'iDMARK axd Vahlquist. 1931. Biochem. Zeitschr., 230, 245. Yanagawa. 1916. Journ. Pharm. Exper. Therap., 9, 75. YoUNGBURG and Youngburg. 1930. Journ. Lab. Clin. Med., 16, 158. THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 8. Hydrocarbon Secretions and Internal Secretory Systems of the Carduaceae, Ambrosiaceae and Cichoriaceae* M. W. MAYBERRY Department of Botany, University of Kansas Abstrvct: a study of the hydrocarbon secretions and secretory systems of a representative group of plants from the families Carduaceae. Ambrosiaceae and Cichoriaceae is recorded. Based on the comparative anatomy of the internal secretory systems in the various species, it is indicated that the occurrence of specialized systems is typical of this large group of plants. The secretion systems of the species of Cichoriaceae being laticiferous while in the Carduaceae and Ambrosiaceae existing as definite canals. As a result of the anatomical study of secretory systems it has been possible to group the various species in the form of a key. The secretions of the canals, laticiferous sj'stems as found in the paren- chyma cells which stain red with Scarlet R, are classified according to solubility with known resin, fat and caoutchouc solvents. As a result of these and saponification tests the secretions are indicated to be resinous substances. THE secretions and secretory systems of the Carduaceae, Am- brosiaceae and Cichoriaceae (Compositae) have been the object of research of many investigators. Some of these have been im- pelled by the purely scientific aspects of the subject and others by the possibility of finding new sources of products of economic im- portance. Early mention of the secretory system of the Compositae in bo- tanical literature was made by Trecul (1862), who distinguished between the secretory canals and latex vessels by the fact that the latter were limited by their own walls while the former were inter- * This work was carried out in the Botany Department of the University of Kansas under the direction of Prof. W. C. Stevens. The writer desires to make his appreciative acknowledg- ment to Professor Stevens for his help and encouragement during the preparation of the work here presented. (71) 72 The University Science Bulletin cellular spaces without any such layer. He also mentioned the occurrence of endodermal secretion canals in the Tubuliferae repre- sented by Vernonia praealta. Muller (1866- '67) confirmed the opinion that the secretory canals were intercellular spaces. In dealing with the Compositae he says of Inula helenium that it is distinguished by large passages in the pith containing ethereal oils and the occurrence of smaller passages in the cortex. Van Tieghem (1885) has shown that secretory canals are of com- mon occurrence throughout the Compositae and he emphasized the fact that in roots they are formed in close association with the endodermis. Van Tieghem states that in the Compositae the secre- tory system has three different forms: (1) oil-bearing canals, (2) laticiferous cells uniting in a network, and (3) isolated, long resinif- erous cells; also that, aside from certain transitional forms. "Les Radices et les Labiatiflores" have only oil-bearing canals, the Ligu- liflores have only anastomosing latex tubes, while the Tubuliflores, for the most part at least, have both oil-bearing canals and isolated resiniferous cells. Van Tieghem discussed the difference between the secretory canals of the stem and root. He held that the out- standing difference between the two was, that in the stem the canals were surrounded by specialized cells, while in the case of the root they were considered to be merely intercellular spaces formed in close association with the endodermis. He has recorded Ambi'osia trijida as having secretion canals at the external border of each vascular bundle at the union of the pericycle with the phloem. These canals are considered by him to belong to the phloem. Triebcl (1885) gave a description of the development of the oil canals in the roots of a few members of the Compositae. He found that the formation of the canals was always preceded by the tan- gential division of the endodermal cells opposite the phloem, the canals beginning as minute intercellular spaces at the junction of the radial and newly formed tangential walls. These canals became filled with drops of almost colorless oil at a very early stage. The cells surrounding the oil canals had thinner walls and were much shorter than those of the cortex proper, they w^ere meristematic in appearance, being filled with dense protoplasm. He considered that these cells probably played an important part in the secretion of the oil in the canals. The oil was termed an ethereal oil, a conclusion Mayberry: Hydrocarbon Secretions 73 based on the following facts: (1) that most of the plants examined had a characteristic smell, (2) that the oil was soluble in alcohol, and (3) that it stained with alkanet. Col (1899-1904) studied the secretory apparatus mainly from the point of view of the classification of the group. He brings to light the following main facts: (1) a survey of the secreting apparatus of the different tribes of the Compositae shows that according to the phyllogenetic progression one finds a gradual replacement of secre- tion canals by a laticiferous system; however, the transition is not an equal gradation in the roots, stems and rhizomes. The transition takes place first in stems, then in rhizomes and roots. (2) Secretion sacs or pockets are simply reduced secretion canals, differing only in longitudinal extent. (3) There are really only two forms of in- ternal secreting apparatus in the Compositae, the canals and anasto- mosing letex tubes. The occurrence of sacs or pockets and of soli- tary tubes more or less branching is evidence of either advancing or regressive evolution. Col also distinguished three classes of secret- ing apparatus: (1) anastomosing latex vessels, (2) secretory canals, (3) isolated cells which secrete latex. AVork by Tschirch (190G-1934) on resins was, for the most part, concerned with the resin and resin ducts of the Coniferae. However, his observations included some on the secretory canals of Com- positae. He concluded, as a result of this work, that resin formation in the Coniferae took place in what he termed as ''resinogener Schicht," which he found lining the canals. He considered that the same layer was present in the secretory canals of the Compositae, but he called this a ''Schleimschicht" or mucilage layer, and it was this layer that was supposed to excrete the resin into the canal. He considered that the young resin canals were filled with mucilage be- fore an}' resin is secreted and that the older secretion canals were lined somewhat evenly with mucilage. In this layer numerous drop- lets of the secretion were thickly embedded and the cavity sur- rounded by the mucilage layer contained the most of the secretion. Tschirch was of the opinion that the secretion of resin droplets takes place only in the resin canals and not in the epithele cells. Moenike's (1924) chief object in his research was to clear up the formation of secretions in the Umbelliferae, Compositae and Ara- liaceae, and to prove up on Tschirch's idea of there being a resin- ogenous layer, and that the secretion does not occur in the epithele 74 The University Science Bulletin cells, but in the resin canals. He considers that there is no resin- ogenous layer in any of the above-mentioned families. Moenike did not observe a mucilage layer in the canals of Compositae. He held that the epithele cells do contain resin droplets, although extraor- dinarily minute. By chemical and solubility tests he shows that the droplets in the epithele cells and the resin in the canals are the same substance, and so concludes that the resin is secreted in the epithele cells and then transferred to the canals. Moenike does not know how the droplets get through the wall of the epithele cells when they are excreted into the resin canals. He finds droplets of the secretion adhering to the walls bordering the resin canal just where the droplets accumulate in the epithele cells. Another object of Moenike was to determine the nature of the secretions in the se- cretion canals of the roots of the Umbelliferae, Compositae and Araliaceae. Tetley (1925) in more recent work on the roots of the Compositae has discussed the formation of the canals and attempted an in- terpretation of her observations in the light of work which has been done on the endodermis. She classifies the secretion canals in the roots into two main groups: (1) endodermal, and (2) nonendoder- mal. The canals are said to be invariably formed opposite the phloem in very early stages. Microchemical studies of the secretion are discussed. She considers the saponification test which Moenike used as unfeasible, due to the minute quantity of substance secreted within the cells. However, she concludes that in developmental stages the secretion is a fatty substance with the nature of an un- saturated drying oil. Various theories of the method of deposition of the secretion in the canals are discussed, and she concludes that the fat is released during the differentiation of the phloem, from which it passes outwards across endodermis by way of the radical walls to the endodermal canals in which it is deposited. Lloyd (1932), in an investigation of caoutchouc in Partheniwn argentatum, discusses possible uses of the secretion to the plant. He feels that very little evidence has been shown for its being used in the metabolic processes of the plant or as an aid to drought re- sistance. Whitaker (1922), in the study of the anatomy of certain golden- rods, states, "Another point of interest, which is of course a common anatomical character of the Tubuliflorae, is the presence of oil canals in the pith and cortex, or in both." Mayberry: Hydrocarbon Secretions 75 Comparative Anatomy of the Internal Secretory Systems in THE Various Species Agoseris cuspidata Root. Latex tubes are scattered throughout the phloem and cortex. Stem. Latex tubes occur in the phloem and on the inside of, and lying in contact with, a slightly suberized endodermis. The suberized endodermal cells are decidedly tangentially flattened. Latex tubes are also found in the pith in association with isolated groups of phloem. Leaf. Latex tubes are found below each of the veins and in midrib. Those associated with the veins are in contact with the phloem. In the midrib the tubes are next to the group of primary hard bast beneath the phloem. Ambrosia elatior L. Root. Relatively s^mall, inconspicuous canals averaging .015 mm. in cro-ss diameter are found in the inner cortex, being more numerous radially op- posite the phloem strands. Stem. Canals averaging .015 mm. in cross diameter occur within the stem cortex lying alternate with the groups of primary hard bast (Plate III, fig. 5). Leaf. No specialized secretory system is present. Ambrosia trifida Root. Numerous canals without a differentiated epithele cell layer are found in the inner cortex and phloem which average .015 mm. in cross diameter (Plate I, fig. 2). Those of the inner cortex on the root are grouped radially opposite the phloem strands. Stem. Canals in the cortex, phloem and pith. Those of the cortex are com- paratively few, numbering from four to five as seen in cross section. The average canal is .03 mm. in cross diameter and is surrounded with approxi- mately two rows of secreting cells. Canals of the phloem are very inconspicuous, as their epithele cells are about the same size and shape as the cells of the surrounding phloem par- enchyma. The canals average .007 mm. in cross diameter and there may be from one to several canals within the phloem of each bundle. In the pith the canals are located near the protoxylem points at the inner margin of the rays and are comparatively small, averaging .009 mm. in cross diameter. They have a definite epithele cell layer composed of from five to eight cells. Leaf. Canals are found in the midrib only, one in the ground tissue and several in the phloem. The one occurring in the ground tissue is similar to those in the stem cortex. It§ position is below the central vascular bundle. Around this canal are two to three layers of secreting cells which are filled with a dark-brown substance. These canals average .02 mm. in cross diam- eter. The canals of the phloem are quite small and obscure, averaging .01 mm. in cross diameter. There are often several of these canals in the phloem of a single vascular bundle. 76 The University Science Bulletin Aniphiachyris dracunculoides Root. There is a suberized endodermis, but no specialized secretory system is present. Stem. The canals, averaging .05 mm. in cross diameter, occur within the cortex only and often lie in an indentation of the endodermis. They have a definite epithelium with some cells joined with the suberized endodermis. Leaj. One canal occurs beneath each of the larger veins of the leaf blade and one in the ground tissue of the midrib (Plate V, fig. 3), those below the veins having their epithelium cells in contact with the phloem. An average canal of the midrib is .05 mm. in cross diameter and an average canal of the mesophyll is .03 mm. in cross diameter. The single canal of the midrib has the same position in relation to the phloem as have the canals beneath the veins. Aster azureus Root. Canals, averaging .012 mm. in cross diameter, are found located radially opposite the phloem, their epithelium cells lying in contact with the suberized endodermis. Stem. There is a suberized endodermis, but no specialized secretory system. Leaf. Only one canal is found in the leaf, lying in the ground tissue on tlie lower side of the vascular bvmdle of the midrib. The average cross diameter was .003 mm. Aster ericoides Root. There is no specialized secretory system. Stem. Canals, averaging .03 mm. in cross diameter, occur in the stem cor- tex with their epithelium cells lying in contact with a suberized endodermis. Leaf. Canals are present in both midrib and mesophyll, those of the meso- phyll always occurring singly below a larger vein. Only one canal is found in the midrib, near the phloem in the ground tissue. Average canals of the Iraf are .02 mm. in breadth. Rhizone. A specialized secretory system is lacking. Aster Drummondii Root. No occurrence of a specialized secretory system is found. Stem. No specialized secretory system is in evidence. Leaf. There is no specialized secretory system. Rhizome. There are canals averaging .045 mm. in cross diameter in the cortex, lying in contact with the endodermis. Bidens frondosa Root. A specialized secretory system does not occur. Stem. Canals are found in the cortex and "pith. The cortical canals are irregularly dispersed within the inner cortex, and are comparatively numerous, but relatively small, averaging .02 mm. in cross diameter. The epithelium cells are not different in shape and size from the other cortical cells; how- ever, from one to three rows of the adjacent cortex cells contain a dark-brown substance which marks very conspicuously the position of the canals. (Plate II, fig. 1.) Mayberry: Hydrocarbon Secretions 77 The canals of the pith are veiy sparse and relatively inconspicuous, ex- ccpting for the fact that their epithelium cells also contain a dark-brown substance. The canals average .01 mm. in cross diameter and are surrounded with four or five epithele cells. They occur close to the protoxylem points. Leaj. Comparatively small canals averaging .003 mm. in cross diameter occur singl.y on the lower side of the veins and on the midrib. These are made conspicuous by the presence of a dark-brown substance found in the epi- thelium celL-;. These cells always follow the veins of a leaf, as is plainly apparent in bleached leaves, whether or not a canal is formed. Bidens involucrata Root. Several relatively inconspicuous canals averaging .015 mm. in cross diameter are found in the inner cortex. Stem. Canals occur in cortex and pith. The canals of the cortex are quite similar to those of Bidens frondosa, but occur radially opposite the rays. The canals average .01 mm. in cross diameter and are surrounded with from one to three rows of cells containing a dark-brown substance. These cells do not form a well-differentiated epithelium, but have the shape and appearance of the other cortical cells. The pith canals are not so closely associated with the protoxylem points as in Bidens frondosa, but do lie in the outer part of the pith cylinder. These have a comparatively definite epithelium of from four to seven somewhat four-sided cells. Leaf. Secreting velh with dark-brown contents are found conspicuous in the mesophyll and midrib of this species, as was the case m Bidens frondosa. The canaLs of the midrib are dispersed through the ground tissue with a somewhat definite arrangement in relation to the vascular system. Where the secreting cells surroimd a definite canal they average .005 mm. in cross diameter. Boltonia latisquama Root. There are all told five canals which average .015 mm. in cross diam- eter, occurring singly and radially opposite a phloem group, and one to three cells of the suberized endodermis compose a part of the epithelium. Stem. The only canals present in the stem are located in the cortex. These are comparatively large, averaging .045 mm. in cross diameter. The epithe- lium lies in contact with a suberized endodermis. These canals number from one to four in the cortex, alternating tangentially with several large groups of primary bast fibers. Leaf. The canals (or pockets) are relatively conspicuous and numerous in the leaf blade (Plate V, fig. 6). These average .04 mm. in cross diameter and are found singly below a vein in each case, and often are larger than the veins with which they are associated. A single canal averaging .04 mm. in cross diameter is found in the ground tissue on the lower side of the vascular bundle of the midrib.- Rhizome. Comparatnely large secretion pockets averaging .15 mm. m cross diameter are found within the cortical and pith region of the thickened rootstock. These extend longitudinally approximately .15 mm. 78 The University Science Bulletin Erigeron canadensis Root. Very inconspicuous canals, .015 mm. in cross diameter, occur in the root radially opposite the phloem within the inner cortex. Stem. Canals occur in both cortex and pith. The epithelium of the cortical canals lies against the suberized endodermis. An average canal is .04 mm. in cross diameter (Plate III, fig. I). Those canals occurring in the pith are comparatively few and are dispersed throughout the entire pith cylinder. Each canal is surrounded by several layers of secreting cells. The canals are relatively small, averaging .012 mm. in breadth. Leaf. Very conspicuous canals, averaging .09 mm. in cross diameter, occur below the veins. Where these are located the leaf is decidedly swollen, espe- cially at the margins (Plate V, fig. 8). A single canal occurs in the ground tissue below the vascular bundle of the midrib. Erigeron ramosus Root. Canals averaging .018 mm. in cross diameter are found radially op- posite the phloem groups. These occur in association with a slightly suberized endodermis, the endodermis furnishing the inside boundary of the intercellular space. Stem. Canals are found only in the cortex lying against a suberized en- dodermis. These canals are often comparatively inconspicuous from being somewhat tangentially flattened. The average canal is .02 mm. in cross di- ameter. An epithelium ^surrounds each canal, lying immediately in contact with the endodermis. These canals are usually radially opposite the groups of primary hard bast and vascular bundles. Leaj. Canals averaging .015 mm. in cross diameter are associated with the vascular bundles of the midrib and with the larger veins, a single canal occur- ring on the lower side near the phloem in each case. Eupatorium altissimum Root. Several comparatively small canals surrounded by four to five epithelium cells are seen in the inner cortex. These average .02 mm. in cross diameter. Stem. Fairly conspicuous canals, averaging .05 mm. in diameter, are found only in the cortex alternate with the primary hard-bast groups which are op- posite the vascular bundles. Leaj. There are canals averaging .02 mm. in cross diameter in both meso- phyll and midrib, laterally to the A^eins in each case. Eupatorium urticaej olium Root. There are two to three relatively small, inconspicuous canals aver- aging .015 mm. in breadth, occumng radially opposite each phloem group. Stem,. Canals occur in the cortex and pith. The canals of the cortex are found radially opposite the medullary rays and in depressions between the groups of primary hard bast. The canals are relatively inconspicuous because they and their epithelium cells are approximately the same size and shape as Mayberry: Hydrocarbon Secretions 79 the surrounding cortical cells. An average canal is .015 mm. in breadth. The innermost epithelium cells are in contact with a suberized endodermis. In the pith the canals are located indefinitely through the outer region of the pith and are similar to those of the cortex in size and shape. Leaf. Several comparatively inconspicuous canals occur in the ground tissue above and below the vascular bundle of the midrib. These have no well- differentiated epithelium layer and average .01 mm. in breadth; also, canals averaging .01 mm. in breadth are associated with the veins. Euthamia camporum Root. No specialized secretory sj^stcm is present. Slbvi. Canals are infrequent and occur only opposite the hard bast in the cortex, with the innermost epithelium cells in contact with the definitely suberized endodermis. The average canal is approximately .03 in breadth. In older stems these canals are verj^ hard to find because of the tangentially flattened epithelium cells and canals. Leaf. Relatively large and conspicuous canals, averaging .06 mm. in cross diameter, are associated with the veins on the lower side near the phloem. ' A single canal, approximatelj' .03 mm. in breadth, is embedded in the ground tissue of the midrib on the lower side. Rhizome. Canals averaging .06 mm. in breadth are found lying radially opposite the phloem groups and in contact with a suberized endodermis. Galinsoga ciliata Root. No specialized secretory system is present. Stem. Canals averaging .006 mm. in breadth are found in the cortex. These are relatively inconspicuous because the)- and their epithelium cells are ap- proximately the same size and shape as the surrounding cortical cells. Leaf. Canals averaging .008 mm. in cross diameter occur singly above the veins and midrib. Grindelia squarrosa Root. No specialized secretory system is to be seen. Stem,. Canals are found only in the cortex in slight indentations of the suberized endodermis. They are quite incon.^picuous, averaging .03 mm. in cross diameter and having no definite epithelium cell-layer. The canals are somewhat tangentially flattened against the endodermis. They occur opposite the primary hard bast which in turn is opposite the vascular bundles. Leaf. Canals averaging .04 mm. in cross diameter are located below a few of the larger veins near the phloem. In the midrib a single canal of about the same size as the others is embedded in the ground tissue below the vascular bundle. Helianthus annuus Root. Tangentially flattened canals averaging .04 mm. in breadth are found in the cortex with their epithelium lying in contact with a suberized endo- dermis. One or more are found to occur radially opposite the phloem groups. Stem. The canals are found in the cortex and pith. Those of the cortex are comparatively large, averaging .08 mm. in cross diameter, and are dispersed 80 The University Science Bulletin through the cortical parenchyma with no definite position in relation to the vascular bundles. The epithelium cells have about the same shape as the surrounding cortical cells. In the pith the canals are dispersed near its periphery and are much smaller than those of the cortex, averaging .015 mm. in breadth. They have a veiy definite epitheliimi layer composed of rectangular cells. Leaj. The canals, averaging .04 mm. in diameter, are associated with both the veins and the midrib, occurring singly above the veins, and in the midrib there are eight canals dispersed within the ground tissue of the lower side in definite relation to the five vascular bundles. Helianthus rigidus Root. Numerous comparatively inconspicuous canals averaging .015 mm. in breadth occur within the inner cortex radially opposite the phloem. These lie in contact with the endodeiTuis. Stem. Canals are found in the cortex, phloem and pith. Those of the cortex are relatively large, averaging .09 mm. in breadth, while some are as large as .2 mm. (Plate II, fig. 3). They are dispersed in the cortex, sometimes opposite the vascular bundles, and again opposite a ray. The epithelium is very pronounced, with comparatively large rectangular cells. The canals of the phloem also are larger than those in the phloem of most other species, averaging .012 mm. in breadth. Several canals may be found in each phloem group, each with a definite epithelium (Plate II, figs. 2 and 3). In the pith the canals are fairly evenly dispersed throughout the entire pith cylinder. These are also relatively large, averaging .04 mm. in cross diameter, and ha\ing a definite epithelium layer with decidedly tangentially flattened cells. Leaj. Canals are found above the veins and in the ground tissue and phloem of the midrib, those above the xeins and in the midrib ground tissue averaging .03 mm. in breadth, while in the midrib phloem and average canal is .015 mm. in breadth. In the midrib there are four canals located below the three main vascular bundles, and the phloem of each vascular bundle may have several canals (Plate V, fig. 7). Canals are found in both cortex and pith; those of the cortex are com- paratively large, averaging .06 mm. in cross diameter (Plate IV, fig. 3). The canals of the pith are dispersed throughout the pith cylinder with an average cross diameter of .02 mm. Helianthus salicifolius Root. There are several tangentially flattened canals in the inner cortex which average .04 mm. in breadth. These do not have a well-defined epithelium. Stem. There are numerous comparatively large canals in the cortex of the stem and several smaller canals in the pith. Those located in the pith occur near its periphery but have no particular position in relation to the protoxylem points. These average .015 mm. in breadth, and have from six to eight epi- thelium cells. The cortical canals average .04 mm. in cross diameter and are surrounded with from twenty to twent3'-five rectangular epithelium cells. Mayberry: Hydrocarbon Secretions 81 Leaf. Canals occvir in the midrib and above the veins, averaging .03 mm. in cross diameter. There are four canals located in the ground tissue of the midrib on the lower side of the vascular bundle (Plate V, fig. 5). These are conspicuous for their well-diflferentiated, triangular epithelium cells. Rhizome. Numerous canals forming a broken cj-linder are found within the inner cortex. The canals group themselves most noticeablj- radialK- opposite the phloem strands. These canals average .03 mm. in breadth, while canals occurring in the phloem average .05 in breadth. Helianthus strumosus Root. Numerous comparatively large tangentially flattened canals are found scattered within the inner cortex and phloem averaging .04 mm. in breadth (Plate I, fig. 4). The epithelium is not well-differentiated. Stem. Canals occur in the cortex, phloem and pith. The canals of the cortex have no definite location in respect to the vascular system. An average canal here is .06 mm. in breadth and is usually .somewhat tangentially flattened. These canals have a comparatively pronounced epithelium composed of rec- tangular cells (Plate II, fig. 5). The phloem canals are relatively small, averaging .007 in breadth. They each have a definite epithelium. The canals of the pith are also comparatively small, averaging .008 in cross diameter, but are numerous and scattered throughout the pith cylinder. Leaf. Canals averaging .02 mm. in breadth occur above the veins. In the midrib the canals averaging .04 mm. across, are found in the ground tissue. Rhizome. There are numerous canals within the cortical parenchjma and pith of the rhizome. Those of the cortex average .06 mm. in breadth, while in the pith the canals average .04 mm. in breadth. Canals averaging .015 mm. are often found in the phloem (Plate IV, fig. 4). Helianthus trachelifoiuis Root. Several canals averaging .015 mm. in breadth are found immediately opposite the phloem groups and each phloem group has from one to three canals. Stem. Canals occur in the cortex, phloem and pith of the stem. Those in the cortex have no uniform arrangement in relation to the vascular system. They usually are slightly tangentially flattened and average .04 mm. in breadth. The epithelium is not as pronounced as in other species of this genus, though each canal does have a definite layer. Canals of the phloem often occur two or three to a vascular bundle and average .007 mm. in breadth. There is a definite epithelium surrounding each canal. In the pith the canals are also comparatively small, averaging .007 mm. in cross diameter. These canals are located in considerable numbers near the protoxylem points and their epithelium cells average four or five to a canal. Leaf. In the midrib canals occur in the ground tissue and in the phloem of the vascular bundle. There are six canals within the ground tissue, averaging 0—6037 82 The University Science Bulletin .04 mm. in breadth. These have a definite arrangement in relation to the five vascular bundles. The canals in the phloem average .012 mm. in breadth. No canals are with the veins. Rhizome. Numerous relatively small canals averaging .018 mm. in breadth are found in the cortex, pith and phloem. Heliopsis scabra Root. The canals are relatively inconspicuous, appearing as minute inter- cellular spaces filled with secretion. These average .01 mm. in cross diameter and are located in the inner cortex. Stem. Canals averaging .01 mm. in cross diameter occur in cortex and pith. They lie in the cortical parenchyma between the large groups of primary hard bast and opposite a medullary ray in each case. The epithelium is composed of comparatively small cells which have in general the same shape as the sur- rounding cortical parenchyma cells. In the pith the canals are similar in shape to those of the cortex and are only slightly smaller in size, averaging .008 mm. in cross diameter. These are close to the protoxylem points and alternate with them. Leaf. There are canals below the larger veins and in the midrib. Those with the veins average .015 mm. in cross diameter. In the ground tissue of the midrib the six relatively small canals are dispersed with definite regularity in relation to the five vascular bundles. An average canal of the midrib is .02 mm. in cross diameter. Iva ciliata Root. No specialized secretory system occurs. Stem. The stem of Iva ciliata has canals occurring in the cortex only. They are just outside the groups of primaiy hard bast. They average .04 mm. in cross diameter and have a definite epithelium layer composed of rectangular cells. Leaf. The larger veins of the leaf blade have canals located above them averaging .015 mm. in breadth. A single canal averaging .015 mm. in breadth occurs above the vascular bundle of the midrib. Lactuca pulchella Root. There are numerous latex tubes occurring in radiating groups op- posite the phloem. Stem. Latex tubes are found in the cortex and phloem. Those of the cortex form a row lying in contact with each of the groups of primary hard bast (Plate III, fig. 4). Leaj. Latex tubes occur in association with each of the veins and in the ground tissue of the midrib. Lepachys columnifera Root. Several large secreting pockets are located within the inner cortex. These average .06 mm. in breadth. Mayberry: Hydrocarbon Secretions 83 Stem. The stem of Lepachys columnifera has comparatively few and small canals. These occur in the pith only, near the protoxylem points. The aver- age canal is .006 mm. in cross diameter and has relatively small epithelium cells. Leaf. Canals averaging .015 mm. in breadth are found in the mesophyll and midrib. Lepachys pinnata Root. Canals appearing as ordinary intercellular spaces, because of the similarity of the epithelium cells to the cortical cells, are found in groups of three or four radially opposite the phloem. An average breadth is .04 mm. Stem. Canals occur in both cortex and pith (Plate III, fig. 3). The canals in the cortex are not numerous and are opposite the groups of primary hard bast. They are comparatively obscure because they are tangentially flattened against the suberized endodermis. The average canal is .03 mm. in cross diameter. The epithelium layer is composed of relatively small cells. Leaf. Two canals averaging .01 mm. in breadth occur in the ground tissue of the midrib, one above and one below the vascular bundle, and canals aver- aging .008 mm. in cross diameter occur above some of the main veins. Rhizome. Tangentially flattened canals averaging .06 mm. in diameter occur in the inner cortex radially opposite each phloem group. Liatris pycnostachya Root. There is an average of three canals seen in cross section. These lie next to the endodermis and are radially opposite the phloem. An average canal is .03 mm. in breadth. Stem. Canals found in the stem occur in the cortex only. They are com- paratively inconspicuous, lying against and within an indentation of the un- dulated primary hard bast. The average canal is .015 mm. in breadth and is surrounded with from five to seven epithelium cells which are not well differ- entiated from the cortical parenchyma. Leaf. Comparatively inconspicuous canals averaging .01 mm. in breadth occur laterally to the veins, while in the midrib there is a canal in the ground tissue on each side of the vascular bundle. Corm. The corm has exceptionally large secretion pockets embedded in the parenchymatous tissue, a feature common in this genus (Plate IV, fig. 5). The pockets, averaging .9 mm. in cross diameter, are lined with several rows of tangentially flattened cells. Liatris punctata Root. The canals are similar in position and size to those of Liatris pycnostachya. Stem. The canals of this species, averaging .02 mm. in breadth, occur in the cortex only, lying against the primary hard bast, and having about the same location and size as in the species aspera and pycnostachya. Leaf. The location and size of the canals of this species are similar to those of Liatris pycnostachya. Corm. Secretion pockets occur like those in Liatris pycnostachya. 84 The University Science Bulletin Liatris aspera Root. The canals have comparatively no difference in size and location from those in Liatris pycnostachya and punctata. Stem. Canals are found in the cortex only and average .02 mm. in cross diameter. These have no well-differentiated epithelium layer. Leaf. The canals in position and size resemble those of Liatris pycnostachya and punctata. Corm. Secretion pockets occur like those in Liatris pycnostachya and punctata. Mesadenia atri-plicijolia Root. Relatively large canals, averaging .12 mm. in cross diameter, one radially opposite each phloem group, are seen in cross section (Plate I, fig. 6). These are next to a suberized endodermis. Stem. Canals with definite epithelium occur in both cortex and pith. The canals of the cortex are comparatively numerous and occur radially opposite a vascular bundle and close to the primary hard bast (Plate II, fig. 6). An average canal is .04 mm. in cross diameter. Canals occurring at the peripheiy of the pith cylinder are like those of the cortex in size and shape. Leaj. Canals occur in both the mesophyll and the midrib, averaging .03 mm. in cross diameter. In the midrib there are usually from two to four canals at the margins of the phloem and in the mesophyll canals occur singly above and below the veins. Rhizome (tuberous). Comparatively large secretion pockets, averaging .08 mm. in cross diameter, are found within the cortex of the tuberous under- ground stem. Pyrrhopappus carolinianus Root. Numerous latex tubes are found in the cortex and phloem. Stem. Latex tubes are in the cortex, phloem and pith. Those of the cortex are in contact with an endodermis, occurring more abundantly radially opposite the vascular bundles (Plate III, fig. 2). Those in the pith are in association with isolated groups of phloem. Leaf. The latex tubes are beneath the veins in the midrib. Senecio plattensis Root. There is no specialized secretory system. Stem. No specialized secretory system is present. Leaf. No specialized secretory system is present. Silphium speciosurn Root. The canals of young roots, averaging .15 mm. in breadth, form a broken cylinder around the stele, noticeably grouping opposite the phloem. Stem. Canals occur in the cortical parenchyma opposite the medullary rays and in the pith (Plate III, fig. 6). In the cortex they are relatively large and conspicuous, averaging .08 mm. in breadth. Comparatively small epithelium cells form a single layer about each canal. Mayberry: Hydrocarbon Secretions 85 The canals of the pith occur in the periphery of the cyhnder. They averago .04 mm. in diameter and each is surrounded with a definite epithelium. Leaf. Comparatively minute canals averaging .006 mm. in cross diameter occur above the veins. In the midrib the canals with triangular epithehum cells are interspersed between the vascular bundles. Rhizone. Canals averaging .04 mm. in cross diameter and with a definite epithelium are found dispersed throughout the cortex, phloem and in the outer regions of the pith cylinder (Plate IV, fig. 2) . Silphium laciniatum Root. Tangentially flattened canals, averaging .018 mm. in cross diameter, are quite numerous within the inner cortex. They have no well-differentiated epithelium and appear simply as intercellular spaces in contact with the cndodermis. Stem. In this species also the canals are found in the cortex and pith. Those of the cortex are dispersed throughout the parenchyma without definite relation to the vascular system. They are comparatively numerous and large, averaging .08 mm. in diameter. Each canal has a definite epithehum of rec- tangular cells. In the pith the canals are located at the periphery of the cylinder and are much larger than the average of other species, being .08 mm. in breadth. There is a definite epithelium composed of comparatively large cells surrounding each canal. Leaf. Outside the midrib, canals averaging .015 mm. in cross diameter are usually found one above and one below the larger veins (Plate V, fig. 4). Several canals averaging .03 mm. in diameter are located in the ground tissue surrounding the vascular system of the midrib (Plate V, fig. 2). Rhizome. Numerous large tangentially-flattened secretion canals occur in the inner cortex averaging .06 mm. in cross diameter. Solidago canadensis gilvocanescens Root. Quite conspicuous canals with an average diameter of .06 mm. occur singly radially opposite each of the phloem groups and in contact with the suberized endodermis. Stem.. There are canals in the cortex and pith of this species. The cortical canals lie against the suberized endodermis and are tangentially flattened. The epithelium cells differ from the other cortical cells only by their relatively small size. An average canal is .04 mm. in cross diameter. The canals in the pith are near the protoxylem points. They average .05 mm. in diameter and have a well-differentiated epithelium. Leaf. All canals are comparatively large, averaging .04 mm. in diameter. They occur more often below than above the veins and those in the ground tissue of the midrib are located above and below the vascular bundle (Plate V, fig. 1). Rhizome. Numerous large canals are found in the cortex and pith (Plate IV, fig. 1). The canals in the cortex he next to a suberized endodermis and average .07 mm. in breadth. Those of the pith cylinder lie near the xylem and average .09 mm. in diameter. 86 The University Science Bulletin Solidago Lindheimeriana Root. The canals, ten in number, averaging .06 mm. in diameter, lie within the inner cortex and in association with a slightly suberized endodermis. The epithehum cells are tangentially flattened and relatively small. Each canal occurs radially opposite a phloem group. Stem. The canals, found in the cortex only, are very inconspicuous and comparatively few. They are decidedly tangentially flattened against the suberized endodermis bounding the primary hard bast. The epithelium cells are comparatively small and tangentially flattened. An average canal is ap- proximately .03 mm. in diameter radially and .06 mm. tangentially. Leaf. The canals are comparatively inconspicuous, averaging .015 mm. in diameter. They occur singly below the veins and in the ground tissue of the midrib below the vascular bundle. Rhizome. Numerous large canals averaging .1 mm. in breadth are found lo- cated within the inner cortex. The cells composing the epithele layer are quite tangentially flattened. Solidago rigidiuscula Root. A single canal lying in contact with a suberized endodermis and averaging .08 mm. in diameter is found opposite each phloem group (Plate I, fig. 1). Stem. This species of Solidago also has canals in the cortex only, located against a suberized endodermis and radially opposite the vascular bundles. The average canal is .06 mm. in diameter. Also, there are canals associated with the suberized endodermis which surrounds each leaf trace as it traverses the cortex (Plate II, fig. 4). The epithelium is not well differentiated. Leaj. A single canal averaging .06 mm. in diameter occurs beneath each of the three vascular bundles of the midrib. Beneath the larger veins is a canal averaging .03 mm, in diameter (Plate V, fig. 9). Solidago rigida Root. The canals have the same position as in the other species of Solidago but are smaller, averaging .04 mm. in breadth. Stem. The canals of this species are very conspicuous because of their size, an average canal being .05 mm. in diameter. They are found in the cortex only, radially opposite the vascular bundles, with the epithelium in contact with the suberized endodermis. Leaf. Canals averaging .05 mm. in diameter occur in the mesophyll and midrib. Those of the mesophyll occur singly below the larger veins. In the midrib a single canal is found in the ground tissue below each of the several vascular bundles. Vernonia fasciculata Root. The root has both a latex system and canals. It is typical of the above ground parts to have latex vessels only, but in the root a single canal, averaging .03 mm. in diameter, is found opposite each phloem group. The latex vessels are quite numerous in the cortex and phloem. Stem. Latex vessels occur in the cortex and phloem, but there are no canals. Mayberry: Hydrocarbon Secretions 87 Leaf. Latex tubes occur in the ground tissue of the midrib and in associa- tion with the veins. Rhizome. One to three canals, averaging .045 mm. in breadth, occur within the inner cortex radially opposite each of the primary phloem groups. Scat- tered throughout the phloem and cortex there are comparatively numerous latex tubes. DISCUSSION The Carduaceae, Ambrosiaceae and Cichoriaceae have been stud- ied by a number of investigators from both anatomical and micro- chemical viewpoints. Those principally interested in the anatomical investigation of the three families, grouped as one family, the Com- positae, were Trecul (1862), Vuillemin (1884), Van Tieghem (1885), and Col (1899). These investigators recognized certain character- istic features of the secretory systems of the Compositae. It was early observed by them that resin canals were widely distributed in the Carduaceae and Ambrosiaceae, while latex systems dis- tinguished the Cichoriaceae. Supplementing these earlier anatomi- cal studies, the research embodied in this paper presents a more complete record of a large number of species, especially in the Carduaceae. Nearly all the species of Carduaceae and Ambrosiaceae that I have studied have resin canals present in roots, stems or leaves; however, Senecio lobatus has no specialized secretory system and Aster drummondii has canals only in the underground stem. Vuil- lemin (1884), however, says that Senecio cordatus has a specialized secretory system in the stem. The roots of species of the Carduaceae present a much more uni- form arrangement of the canals than do the stems, as was recognized by Col (1899). The majority of species, indicated in Table I, have canals; however, none were observed in Amphiachyris dracuncu- loides, Aster azureus, Aster ericoides, Aster drummondii, Bidens frondosa, Euthamia camporum, Galinsoga ciliata, Gnndelia squar- rosa and Senecio plattensis. The resin canals of the roots are found occurring singly or in groups radially opposite the primary phloem. Specialized secreting cells forming an epithelium are not present with the canals of the roots. As stated by Van Tieghem (1885), we do not recognize specialized secreting cells in the roots because each secreting cell has the same form and dimensions as the nonsecreting cells and each excretes its products into the canal which it borders. The comparatively small sizes of canals of some species indicated in Table I, and the lack of a well-defined epithelium, make it difficult at times to locate canals until after treating sections with Scarlet 88 The University Science Bulletin R. The majority of species have relatively small intercellular-space canals in the younger roots, which occur in groups opposite the pri- mary phloem and against the endodermis when this is present, often forming a broken cylinder about the stele. In the genus Solidago the canals are comparatively large (Table I), one occurring radially opposite each primary phloem group (Plate I, fig. 1). The presence of canals in the roots distant from the endodermis is not common, but Helianthus strumosits has canals within the phloem and scat- tered throughout the cortex (Plate I, fig. 4). In only one species of the Carduaceae, Vernonia fasciculata, have I found both resin canals and a latex, system. Here the canals, averaging .03 mm. in cross diameter, occur singly radially opposite each of the primary phloem groups. The latex tubes in this species are dispersed throughout the cortex and some are found in the phloem. In the roots of the species of Ambrosiaceae studied by me the canals in the endodermal region are comparable to those of the Carduaceae. Ambrosia triflda has canals in the phloem of the root also. No secretion system was observed in Iva ciliata. In the Cichoriaceae a latex system is commonly found in the roots as well as in other parts. The latex tubes here are dispersed somewhat in the cortex, and mostly radially opposite the phloem, with a few in the phloem. The species of Carduaceae, excepting Senecio plattensis and Aster drummondii, have canals in some part of the stem (Table II). The canals are relatively more common in the cortex, but sometimes they are found in the phloem and pith. Those species having canals in the cortex, as indicated in Table II, may be divided into two groups according to the location of the canals. When an endodermis is present the canals characteristically lie with their epithelium against the endodermis. This characterizes one group, while in the second group the canals are found dispersed in the cortex. Col (1889) says that in the stem the secretion canals are always endo- dermal in origin. Since my finding a large group of species in which the cortical canals are in no way associated with an endodermis Col's statement needs modification. Van Tieghem (1885) makes the statement that besides the endodermal secretion canals others occur in the cortex of stems beneath the epidermis, and in the parenchyma of the leaves, but he considers these to be rare. Species mentioned by Van Tieghem as having cortical canals not in association with an endodermis were Kleninia neriifolia, Solidago limonifolia, Solidago sempervirens, Solidago laevigata and fuscata. Mayberry: Hydrocarbon Secretions 89 I find Solidago canadensis gilvocanescens, rigidiuscida (Plate II, fig. 4), Lindheimeriana, and rigida to have cortical canals in contact with an endodermis. There are comparatively great differences in sizes of cortical canals in the various species. The sizes range from .008 mm. in cross diameter in Heliopsis scabra to .05 and .09 mm. in species of Helianthus, Solidago and Silphium (Table II). Position of the canals in relation to the vascular system, whether radially opposite or alternating with the phloem, varies with the different species, but this feature is not stable enough in the various species for their identification, although Solereder (1908) refers to it as being the most useful for systematic purposes (mostly as a generic character). The occurrence of well-defined epithelium cells is not consistent in the stem, but is more frequent there than in the root. Species of Helianthus exhibit strikingly well-differentiated secreting cells surrounding canals in the cortex (Plate II, fig. 5). Solereder (1908) says that the epithelium is probably of little value as a char- acteristic anatomical character, since resin canals devoid of epithe- lium and others provided with it occasionally occur side by side in the same plant {Lasthenia glabrata, Cynaria Corsica) and that the canals without an epithelium do not always appear in all species of a genus (Cacalia, Senecio). I have observed Mesadenia (Cacalia) atriplicifolia (Plate II, fig. 6) to have canals in the cortex with an epithelium, while Senecio plattensis has no canals in the stem. Those species having canals within the phloem of the stem are not numerous in the Carduaceae. Helianthus rigidus (Plate II, fig. 2), stru7nosus and trachcUfolius were observed to have one or more canals in each phloem group, averaging .012 mm., .007 mm. and .007 mm in breadth, respectively. In the Ambrosiaceae, Ambrosia trifida has one or more canals in each phloem group of the st€m. Vuillemin (1884) considers it an exception to find canals located within the phloem of the stem. Canals in the pith are found in a majority of the species of Car- duaceae and Ambrosiaceae. These canals are smaller on an aver- age than those of the cortex, as indicated in Table II, ranging from .009 mm. in Ambrosia trifida to .08 in Silphium laciniatum. The characteristic position of the canals in the pith is at the periphery where they usually occur opposite the protoxylem points in the va- rious species (Plate III, fig. 3). Van Tieghem (1885) observed canals occurring either singly or several together in the pith of species of Ageratum, Carduus, Dahlia, Helianthus, Serratula, Soli- dago, and Spilanthes. 90 The University Science Bulletin Of the Carduaceae I have found only one member, Vernonia fas- ciculata, to have a latex system in the stem. In this species the latex tubes are scattered within the inner cortex, phloem and more abundantly in the pith. Trecul (1862) speaks of Vernonia praealta as having endodermal secretion canals, and Van Tieghem (1885) has found latex tubes in the phloem of this species. The stems of species of Cichoriaceae have secretory systems in the form of latex tubes, a feature said by Van Tieghem (1885) and Col (1899) to be typical of the group. I find that these may occur in the cortex, phloem and pith. In the cortex they are most abun- dant within the inner cortex forming a broken-undulated cylinder (Plate III, fig. 4). In Pyrrhopappus carolinianus, there are latex tubes in association with isolated phloem groups within the pith (Plate III, fig. 2). The perennial species of Carduaceae whose underground stems I have studied have either canals or secretion pockets as typical fea- tures. The rhizome may have canals located in the cortex, phloem and pith, as is true for the above-ground stem also (Plate IV, figs. 1-6). Helianthus strumosus, Helianthus trachelifolius and Silphium speciosum were of this type. The rhizomes of Boltonia latisquama, Helianthus rigidus and Solidago canadensis gilvocanescens have canals in the cortex and pith, while Aster Drummondii, Mesadenia atriplicifolia, Euthamia camporwn, Lepachys pinnata, Silphium laciniatum, Solidago Lindheimeriana and Vernonia fasciculata have canals only in the cortex. The species of Liatris I have studied have corms and the secretion systems found here consist of large pockets scattered through the parenchymatous tissues (Plate IV, fig. 5). The underground stem of Aster ericoides did not seem to have a specialized secretory system. The presence of a specialized secretion system in the leaves of the various species of Carduaceae and Ambrosiaceae is as typical of the two groups as is its presence in other parts, as indicated in Table III. Senecio plattensis, Ambrosia elatior L. and Aster drum- mondii are the only species I studied which did not have a special- ized secretion system in some part of the leaf blade. In the leaves, canals may be found, one or more in the ground tissue of the mid- rib and in association with the veins of the mesophyll (Plate V, figs. ]-9). In only three species, Ambroisia triflda, Helianthus rigidus (Plate V, fig. 7), and Helianthus trachelifolius, were one or more canals found in the phloem of the midrib. The position of the canals of the leaf is a feature of taxonomic value affording the de- MaYBERRY: H'iDROCARBON SECRETIONS 91 markation of species into groups (see page 94) . The usual position of the canals is above or below the veins. However, cases were found where the canals occur laterally to a vein. The approximate breadth range of the canals is from .003 mm. in Bidens frondosa to .09 mm. in Erigeron canadensis. Only one member of the Car- duaceae, Vemonia fasciculata, was observed to have latex tubes in the leaf. These, as was characteristic of the canals, were closely associated with the veins of the mesophyll and midrib. Latex tubes occur characteristically in the leaves of Cichoriaceae, where they are seen to follow the veins. TABLE I. — Summarizing the cross diameter in mm. of the canals of roots Actinomeris alternifolia 015 Agoseris cuspidata Latex system Ambrosia elatior L 015 Ambrosia trifida 015 Amphiachyris dracunculoides Aster azureus 012 Aster ericoides Aster drummondii Bidens frondosa Bidens involucrata 015 Boltonia latisquama 015 Erigeron canadensis 015 Erigeron raniosus 018 Eupatorium altissiimim 02 Eupatorium urticaefohum 015 Euthamia camporum Galinsoga ciliata Helianthus annuus 04 Holianthus salicifolius 04 Heliantlius rigidus 015 Hehanthus stnimosiis 05 Helianthus trachelifolius 015 Heliopsis scabra 01 Iva ciliata Lactuca pulchella Latex system Lepachys columnifera 05 Lepachys pinnata 04 Liatris pycnostachya 03 Liatris punctata 03 Liatris aspera 03 Mesadenia atriplicifolia 12 Pyrrhopappus carolinianus Latex system Senecio plattensis Silphium speciosum 015 Silphium laciniatum 0^5 Solidago canadensis gilvocanescens 06 Solidago Lindheimeriana 06 Solidago rigidiuscula 08 Solidago rigida 04 Vemonia fasciculata (Also latex system) . 03 92 The University Science Bulletin TABLE II. — Summarizing the location and diameters in mm. of the specialized secretory systems in the stem Cortex Actinomeris altemifolia Agoseris cuspidata Latex Ambrosia elatior L 015 Ambrosia trifida 025 Aster azureus Aster ericoides Aster drummondii Bidens f ondosa 02 Bidens involucrata 01 Boltonia latisquama Erigeron canadensis Erigeron ramosus Eupatorium altissimum 05 Eupatorium urticaefolium Euthamia camporum Galinsoga ciliata 006 Grindelia squarrosa Hehanthus annuus 08 Helianthus sahcifolius • - • • -06 Helianthus rigidus 09 Helianthus strumosus 06 Helianthus trachelifolius 04 Heliopsis scabra 008 Iva ciliata 04 Lactuca pulchella Latex Lepachys columnifera Lepachys pinnata Liatris pycnostachya 015 Liatris punctata 02 Liatris aspera 02 Mesadenia atriplicifolia 04 Pyrrhopappus carolinianus Lat«x Senecio plattensis Silphium speciosum 08 Silphium laciniatum 08 Solidago canadensis gilvocanescens Solidago Lindheimeriana Solidago rigidiuscula Solidago rigida Vernonia fasciculata Latex Amphiachyris dracunculoides Cortex* .015 system .03 system .03 Phloem fith in all 007 parts .045 .04 .02 .015 .03 .03 svstem in all .04 .05 .06 .15 system .05 012 007 007 all parts parts 009 01 02 012 015 015 015 04 008 007 01 006 012 04 04 08 in all parts Secretory system in the cortex with epithelium in contact with an endodermis. Mayberry: Hydrocarbon Secretions 93 TABLE III. — Summarizing the location and sizes in mm. of the specialized secretory systems of the leaf Midrib Actinomeris alternifolia 008 Agoseris cuspidata Latex Ambrosia elatior L Ambrosia trifida 02 Amphiachyris dracunculoides 05 Aster azureus 003 Aster ericoides 02 Aster dnimmondii Bidens frondosa 003 Bidens involucrata 005 Boltonia latisquama 04 Erigeron canadensis 09 Erigeron ramosus 015 Eupatorium altissimum 02 Eupatorium urticaefolium 01 Euthamia camporum 03 Galinsoga cilia ta 008 Grindelia squarrosa 04 Helianthus annuus 04 Helianthus salicifolius 03 Helianthus rigidus 03 Helianthus strumosus 04 Helianthus trachelifolius 04 Heliopsis scabra 02 I va ciliata 015 Lactuca pulchella Latex Lepachys columnifera 015 Lepachys pinna ta 01 Liatris pycnostachya 01 Liatris punctata 008 Liatris aspera 008 Mesadenia atriplicifolia 03 Pyrrhopappus carolinianus Latex Sonecio plattensis Silphium speciosum 006 Silphiuni laciniatura 03 Solidago canadensis gilvocanescens 04 Solidago Lindheimeriana 015 Solidago rigidiuscula 06 Solidago rigida 05 Vernonia fasciculata Latex Midrib (phloem) Mesophyll system system system 01 in all parts 015 012 .03 .02 .on 3 .005 .04 .09 .015 .02 .01 .06 .008 .04 .04 .03 .03 .02 .015 .015 in all parts .015 .008 .01 .01 .01 .03 in all parts .006 .015 .04 .015 .03 .05 system in all parts 94 The University Science Bulletin Key to Species Based on Location, Size and Characteristics of Epithelial Cells of the Internal Secretory System I. No specialized secretory system present. Senecio plattensis. Aster drummondii. II. Secretory system present (canals) in roots, steins, and leaves. A. In the stem the canals are present in cortex, phloem and pith. 1. Canals of the leaf in the mesophyll and midrib. a. Canals in the phloem of the midrib. Helianthus rigidus. b. No canals in the phloem of the midrib. Helianthus strumosus. 2. Canals of the leaf in the midrib only. a. One canal in the ground tissue below the midrib vascular bundle. Ambrosia trifida. b. Six canals in the ground tissue around the midrib vascular bundles. Helianthus trachelifolius. B. In the stem the canals are present in the cortex and pith only. 1. Canals in the stem cortex with epithelium in contact with an endodermis. a. Canals of the leaf with cavities not over .015 mm. in cross diameter. (1) Midrib with two canals; one above and the other below the central vascular bundle. Lepachys pinnata. (2) Midrib with several canals; occurring above and below the centra! vascular bundle. Eupatorium urticaefolium. b. Canals of the leaf with cavities ranging from .04 mm. to .09 mm. in cross diameter. (1) Canals of the mesophyll sometimes above, sometimes below the veins. Solidago canadensis gilvocenescens. (2) Canals of the mesophyll always below the veins. Erigeron canadensis. 2. Canals in the stem cortex with epithelium cells apart from the endodermis. a. Canals of the stem cortex averaging .08 mm. or over in cross diameter. (1) Canals of the leaf midrib averaging .03 mm. in cross diameter. Silphium speciosum. (2) Canals of the leaf midrib averaging .04 mm. in cross diameter. (a) Canals of the mesophyll occurring above and below the veins. Silphium laciniatum. (b) Canals of the mesophyll occurring above the veins only. Hellianthus annuus. b. Canals of the stem cortex averaging .06 mm. or less in cross diameter. (1) Epithelium cells of the canals well defined; composed of rectangular cells. Helianthus salicifolius. (2) Epithelium cells erf the canals not well defined nor rectangular. (a) Epithelium cells of the canals containing a dark brown mass. Bidens involucrata. (b) Epithelium cells not as above. 1. Canals of the stem cortex average .008 mm. in cross diameter. Heliopsis scabra. 2. Canals of the stem cortex average .04 mm. in cross diameter. Mesadenia atriplicifolia. C. In the stems the canals are present in the cortex only. 1. Canals of the stem not in contact with an endodermis. a. Underground stem in the form of a corm with large secretion pockets. Liatris aspera. Liatris pycnostachya. Liatris punctata. Mayberry: Hydrocarbon Secretions 95 b. Underground stem not in the form of a corm. Eupatorium altissimum. Actinomeris altemifolia. 2. Canals of the stem cortex with epithelium in contact with an endodermis, a. Canals of the stem averaging .05 mm. or over in cross diameter. Solidago Lindheimeriana. Solidago rigidiuscula. Solidago rigida. Boltonia latisquama. b. Canals of the stem averaging .04 nmi. or less in cross diameter. Erigeron ramosus. D. In the stem the canals are prespnt in the pith only. Lepachys columnifera. III. Canals present in stems and leaves only. A. Canals in stem cortex with epithelium cells in contact with an endodermis. Aster ericoides. Amphiachyris dracunculoides. Euthamia camporum. Grindelia squarrosa. B. Canals in the stem cortex not having epithelium cells in contact with an endodermis. 1. Canals in both cortex and pith. Bidens frondosa. 2. Canals in cortex only. Galinsoga ciliata. Iva ciliata. IV. Canals present in the roots and stems only. Ambrosia eliatior L. V. Canals present in the roots and leaves only. Aster azureus. VI. Canals in the roots only ; latex sy.stem in other parts as well as in the roots. Vemonia fasciculata. VII. Only a latex system present. Agoseris cuspidata. Lactuca pulchella. Pyrrhopappus carolinianus. THE OCCURRENCE AND IDENTIFICATION OF SECRETIONS STAINED RED WITH SCARLET R Secretions which stain red with Scarlet R are found in abundance in the Carduaceae, Ambrosiaceae and Cichoriaceae. Sections taken from various parts of the plant show, as a rule, numerous globules of these secretions dispersed in the parenchymatous tissues, and in masses and specialized secretory systems. In many species which have large-cavitied canals the secretion exudes easily from the fresh- cut surfaces. The tissue of the plant which contains the greatest number of dispersed globules is the mesophyll of the leaf. Cross sections of leaves of such species as Solidago canadensis gilvocanescens (Plate V, fig. 1), Amphiachyris dracunculoides (Plate V, fig. 3), Mesadenia atriplicifolia and Boltonia latisquama treated with Scarlet R pre- sent a striking appearance due to the great quantity of dispersed globules. 96 The University Science Bulletin The quantity of secretion in the mesophyll varies among the species observed, but in each some secretion was present. The fol- lowing table gives a comparison of the species based upon the rela- tive amounts of dispersed globules within the mesophyll of the leaf: TABLE IV. — The symbol (t) indicates comparative amounts of secretion in the leaves Agoseris cuspidata .' tt Ambrosia elatior L t Ambrosia trifida t Amphiachyris dracunculoides tttt Aster azureus ttt Aster ericoides ttt Aster drummondii ttt Bidens frondosa t Bidens involucrata t Boltonia latisquama tttt Erigeron canadensis tt Erigeron ramosus tt Eupatorium altissimum tt Eupatorium urticaefolium tt Euthamia camporum tttt Galinsoga ciliata t Grindelia squarrosa tttt Helianthus annuus ttt Helianthus salicifolius ttt Helianthus rigidus ttt Helianthus strumosus ttt Helianthus trachelifolius ttt Heliopsis scabra ttt Iva ciliata ttt Lactuca pulchella tt Lepachys columnifera tt Lepachys pinnata tt Liatris pycnostachya tttt Liatris punctata tttt Liatris aspera tttt Mesadenia atriplicifolia tttt Pyrrhopappus carolinianus tt Senecio plattensis tt Silphium speciosum tttt Silphium laciniatum tttt Solidago canadensis gilvocantscens ttttt Solidago Lindheimeriana tttt Solidago rigida tttt Vernonia fasciculata ttt Actinomeris alternifolia ttt The secretions of the above- and below-ground stems are prin- cij^ally in the canals, but small globules are found dispersed in all of the living tissues. If the stem has chlorenchyma the globules are abundant as in the leaf. The frequency and size of the canals in the stems of the different species indicate the relative amounts of secretion. Mayberry: Hydrocarbon Secretioxs 97 Roots have the least amount of the secretion of any of the plant i:)arts. Certain species are outstanding for their large canals and abundant secretions, such as species of Solidago and Helianthus. Evidence of the nature of the secreted materials within the va- rious parts of the plants has been sought for by various microchemi- cal tests. The microchemical tests have been found altogether satis- factory on account of the relatively small size of the globules present. Chemically different substances have been demonstrated: fats, ethereal oils, i-esins and caoutchouc, all staining red with Scarlet R. A fat is an ester of glycerol with one or more of the higher fatty acids. Essential, or ethereal, oils consist of a mixture of hydro- carbons, mostly terpenes, together witli alcohols such as terpeneol and camj)hors. The chemistry of resins is not nearly as well known as that of fats and ethereal oils. Resin, according to Tschirch (1934) , is neither a chemical nor a physical term applied to any specific substance, hut is one of conunon usage, as is the term tannin, be- ing applied to a group of similar substances. Wcisncr (1!)27) con- siders resins as an excreted product resulting from the plant's me- tabolism. Chemically, resins ai'e considered very complex sub- stances formed by a combination of linkages of various kinds, the majority of which contain phenol groups, aliphatic acids, aldehydes and terpenes. The mici-ochemical tests have been carried on only with fresh materials. Aftei' using Scarlet R for the differentiation of the con- tents occurring in the canals and other parts, relative solubility tests were employed. These alone may not be considered sufficient to distinguish between fats, resins, ethereal oils and caoutchouc, but do serve for a major ])art in making this distinction possible. The soh'ents used were alcholiol, acetone, chloroform, acetic acid, carbon disulphide and benzene. Any of these solvents may dissolve va- rious resins. Alcohol is a solvent for but very few fats, while chloro- fonn and carbon disulphide are well-known fat solvents. Caoutchouc is insoluble in acetone but soluble in benzene. Wiesner (1927) speaks of "Harze" as being distinguished from caoutchouc and gut- tapercha through the fact that resin and resin-like materials are soluble in alcohol and acetone while the other two are insoluble. For further distinction between fats and resins I have used the sa- ponification test for fats. ^Nloenike (1924) considered this to be the most critical tests for fats. For this test I used a concentrated so- 7—6037 9H The University Science Bulletin liition of KOH in methyl alcohol which has been found to produce myelin bodies within five to ten minutes with any of the more com- mon i^lant oils, such as castor bean seed oil, cocoanut oil, palm oil and others. This reagent taken from Tunmann (1!)13) is superior to other saponification tests in the speed of the reaction. This test has been carried on witli sections first treated wdth Scarlet R to in- crease the visibility of the secretions and also with exuded "juice" carrying dispersed globules wliieh were stained red with Scarlet R. Ethereal oils have been tested for, in addition to the solubility test of acetic acid, by using microdistillation. These tests, in conjunction with the tabulated solubility tests, point to the fact that the secre- tions are resinous in their chemical nature. The presence of caout- chouc, which has been found to occur in a number of the examined plants, was made certain by observing the solubility. That is, when the substance was stained red with Scarlet R and was non- saponifiable, but was insoluble in acetone, but soluble in benzene, chloroform, alcohol or carbon disulphide, it was considered to be caoutchouc. A complete list of the plants with the solubility of their secretion follows: Mayberry: Hydrocarbon Secretions 99 TABLE V. — Showing solubility of secretions which stain red with scarlet R Agoseris cuspidata: (globules) (latex) Ambrosia trifida Amphiachyris dracunculoides Aster azureus Aster drummondii Aster ericoides Bidens frondosa Bidens involucrata Boltouia latisquama Erigeron ramosus Erigeron canadensis Eupatorium allissimum Ejpatoriuni urticacfolium Euthamia ramporum Grindelia squarrosa Helianthus annuus Heliantlms salicifolius Helianllius .strumosus Heliantlius trachelifolius Heliopsis scabra Iva ciliata Lactuca pulchella (globules) Laetucca pulchella (latex) Lepachys pinuaf a Liatris pycnostachya Liatris aspera Mesadenia atriplicifolia Pyrrhopappiis caroliuianus (globules) . . Pyrrlioi):ippus carolinianus (latex) Senecio plattensis Silphium speciosum Silphium laciniatum Solidago canadensis gilvocaneseens . . . . Solidago Lindheimeriana Solidago rigidiuscula Vernonia fasciculata Acetone. sol. insol. sol. insol. insol. insol. insol. insol. insol. insol. insol. insol. sol. sol. sol. sol. sol. insol. sol. sol. sol. insol. sol. insol. insol. sol. sol. insol. insol. sol. sol. sol. sol. insol. insol. insol. sol. Benzene. sol. sol. insol. sol. sol. sol. sol. insol. insol. sol. sol. insol. sol. sol. sol. insol. insol. sol. sol. insol. sol. insol. insol. sol, sol. insol. insol. sol. sol. insol. insol. insol. insol. sol. sol. sol. sol. Chloro- form. sol, sol. sol. insol. insol. insol. insol. insol. insol. insol. sol. sol. p. sol. sol. sol. sol. sol. insol. p. sol. sol. sol. sol. sol. sol. p. sol. p. sol. sol, insol. sol. sol. sol. insol. sol. p. sol. p. sol. p. sol. insol. Alcohol. sol. insol. insol. insol. insol. sol. insol. sol. sol. insol. insol. insol. sol. sol. sol. sol, sol, insol, sol. sol. sol, insol. sol. insol. insol. insol. sol. insol. sol. sol. insol. insol. sol. insol. insol. insol. sol. Carbon disul- phide. sol, sol. sol. sol. insol. sol. insol, insol. insol. sol, sol. sol. p. sol. sol. sol. sol. sol. sol. sol. sol. sol. sol. sol. sol. sol. insol. sol. sol. sol. sol. p. sol. sol. insol. sol. sol. sol. insol. Acetic acid. sol. insol. insol. insol. insol. insol. insol. insol. insol, insol, insol, insol. insol, insol. insol. insol. insol. insol. insol. so'. insol. insol. sol. insol. insol. insol. sol. insol. insol. sol. insol. insol. insol. insol. insol. insol. sol. ICO The University Science Bulletin TABLE ^T. — Chemical nature of secretion based upon solul)ility tests Agoseris cuspidata (globules) Resinous (latex) Caoutchouc Ambrosia trifida Resinous Amphiachyris dracunculoides Caoutchouc Aster azureus Caoutchouc Aster drummondii Caoutchouc Aster ericoides Caoutchouc Bidens frondosa Resinous Bidens involucrata Resinous Boltonia latisquama Caoutchouc Erigeron canadensis Resinous Erigeron ramosus Caoutchouc Eupatorium altissimuui Resinous Eupatorium urticaefolium Resinous Euthamia camporum Resinous Grindelia squarrosa Resinous Helianthus annuus Resinous Helianthus salicifolius Caoutchouc Helianthus struniosus Resinous Helianthus trachelifolius Resinous Heliopsis scabra Resinous [va ciliata Resinous Lactuca pulchella (glol)ule<) Resinous (latex) Caoutchouc Lepachys pinnata Caoutchouc Liatris aspera Resinous Liatris pycnostacl:ya Resinous Mesadenia atriplicifolia Caoutchouc Pyrrhopappus carolinianus Resinous Sen"cio plattensis Resinous Silphium speciosum Resinous Silphiuni laciniatum Resinous Solidago canadensis gilvocan;scens Caoutchouc Solidago rigiduscula Caoutchouc Solidago rigida Caoutchouc Vernonia faseiculata Resinous In following the development of a number of species in their pri- mary permanent tissue stages it has been observed that the ma- jority of the young plants do not contain the dispersed globules in their parenchymatous tissues typical of the more mature plants. However, the canals did contain secretion during the formation of the primary meristematic tissues. Considerable difference in time of formation of the dispersed glob- ules so typical of the mature leaf mesophyll has been noted for several species. For this purpose plants of Silphium laciniatum and speciosum, Chrysanthemum leucanthemum. and Helianthus strumo- sus were grown in the greenhouse. The plant of Chrysanthemum leucanthemum. when brought in from the field had a large group of basal leaves, but these contained no globules. Formation of glob- ules occurred soon after the plant had produced its first flowers. The plant of Helianthus strwnosus grew to a height of five feet and Mayberry: Hydrocarbon Secretions 101 was ready to produce flowers when globules began to appear in the leaf mesophyll. Silphium laciniatinn produced a number of large basal leaves without liaving the dispersed globules until after three weeks. Silphium speciosum grew approximately two feet tall before globules were observed. In each of the latter two cases globule formation occurred considerably ahead of flower formation. Similar observations were made of these species when they grew in the field which gave comparable results. Possible functions of these secretions in the plant are not evident. To consider tliat they actually serve as a reserve substance later used in the plant's metabolism might seem justified by the large quantities formed. However, dead parts, especially the leaves of certain species, when examined microchemically exhibit an abun- dance of the secretion remaining. Species showing this were Solidago canadensis gilvocanescens, Boltona latisquama, Helianthus salici- folius, Solidago rigida, Euthamia camporum, Mesadenia atriplici- jolia, Liatris aspcra and pycnostachya. Lloyd (19321, in discuss- ing possible uses of caoutchouc to the plant, says that whether the hydrocarbon in (juestion is, in the case of the guayule, actually a re- serve substance and is actually made use of in metabolism i-equires proof which has not yet appeared. That the presence of the glob- ules in the leaves might serve in drought resistance has been sug- gested by Transeau (1904). Lloyd (1932) says that the view which has been adxanced from time to time, that caoutchouc in the gua- yule confers on the plant some ability to withstand drought, has no con\'inciiii: e\idence. In an attemi)t to dctciinine if the globules would disappear when the plant was starved, indicating their being used in metabolism, young and old plants were brought in and ]:)laced in the dark. The young plants were taken from the field just after the first evident globule formation. The globules remained in the older plants but disai)peared from the younger j^lants. Plants showing this were ]\Ie,'pareil Secreteur des Composees. Bull, de la Soc. bot. 14. Whitaker, Edith S. 1922. Anatomy of Certain Goldenrods. Botanical Gaz., 65:251-260. 15. Wiesner, Julius \on. 1927. Die Roh?toffe des Pflanzenreichs. Vol. 1-2 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 9. The Genus Taphrina. I : An Annotated Bibliography A. J. MIX Department of Botany, University of Kansas Abstract: A critical review is presented of the important literature on the genus Taphrina (including Ascomj^ces, Exoascus, Magnusiella)-, from the time of Fries (1815) to the present. Many references containing'merely information as to distribution of various species have been omitted, as well as some papers of purely practical nature on disease control. NO monographic treatment of the genus Taphrina is avaihible to the average worker hiter than that of Giesenhagen in 1901. Many species have been described since then. Jaczewski published, in 1926, a key to known species with descriptions. His paper is in Russian and is not easily obtainable. Otherwise it would serve well as a means of determining species. A complete list of known species is also given by Laubert in the latest edition of Sorauer's Handbuch der Pflanzenkrankheitcn. This list, however, gives descriptions of only the most important species, and with rare ones the investigator is left with only the name of the parasite and the host. It is proposed to remedy this lack by the publication of a descrip- tive list of valid species of Taphrina (including Ascomyces, Exoas- cus, Taphrina, and Magnusiella). Preliminary to this it seems desirable to present a review of the important literature on the genus. This literature is considerable, and much of it is contained in papers not easily available. The re- view here presented is intended to summarize all important findings of previous investigators and to enable anyone interested in the literature of Taphrina to ascertain readily where the particular in- formation he desires may be found. (113) 8—6037 114 The University Science Bulletin The arrangement of this review in the form of an annotated bibli- ography is considered convenient for consultation, as v^ell as valuable in furnishing a strictly chronological treatment of the literature. Many papers have been consulted whose titles are not here pre- sented. Notes of the occurrence of particular species in new local- ities have, for the most part, not been included, and of the large body of literature of peach-leaf curl only those papers have been cited which seem significant from a mycological point of view. An attempt has been made to follow the best modern usage in the citation of authors' names in connection with the names of host species. In the original articles authors' names are frequently omitted. All the papers listed have been consulted in the original, and it is believed that the abstracts of their contents given below are essen- tially correct. 1. Fries, E. Observationes Mycologicae. 1: 217. 1815. Describes the genus Taphria, and the species Taphria populina aurea on leaves of Populus nigra L. 2. Fries, E. Observationes Mycologicae. 2: 378. 1818. Again describes the above-named species and figures it (PI. VIII, fig. 3). 3. Fries, E. Systema orbis terrarum vegetabilis. Pars I. Plantae Hom- onemeae. Lund. 1825. P. 317. Changes the genus name to Taphrina since Taphria has already been era- ployed as the genus name of an insect. 4. Fries, E. Systema mycologicum. 3: 520. 1832. Changes name of Taphrina aurea, on leaves of Populus nigra, to T. populina. Also fists (p. 30, No. 520) Taphrina alnea Schmidt on leaves of Alnus glutinosa Medic. The latter fungus, not being mentioned in later literature, was prob- ably an Erineum. 5. Zollinger, H. Observationes phytographicae, praecipue genera et species nova nonnulla respicientes. Natur-et Geneeskundig Archief voor Neerlands Indie 1: 372-405; 2: 1-19, 200-273, 563-587; 3: 51-92. Abs. in Flora 5: 300. 1847. Describes a new species Taphrina cissi Zoll. on Cissus varius (?). This species seems uncertain and has not been recognized by most of the later writers. 6. Dbsm.\zieres, J. B. H. J. Seizieme notice sur les plantes cryptogames recemment decouvertes en France. Ann. d. Sci. Nat. 3^ ser. Botanique. 10- 342-361. 1848. Mix: The Genus Taphrina I 115 Describes the new genus Ascomyces Mont, and Desmaz., characterized by ascus-hke sporangia on the surface of the host leaf. Also the new species Ascomyces caerulescens Mont, and Desmaz. on leaves of Qucrcus coccifera L. The new genus is not clearly differentiated from the existing genus, Taphrina. In the species description, however, it is stated that the sporangia constitute the whole fungus, i.e.: mycelium is absent. The spelling of the species name "caerulescens" is to be noted. This spell- ing was followed by Robinson (55) and others. Some later authors have used the spelling "coerulescens." In this paper the original spelling is restored. 7. Fries, E. Summa vegetabilium Scandinaviae. Sectio Posterior. Upsala. 1849. On page 518 gives a description of the genus Taphrina, naming one species, Taphrina populina Fr. as occurring in Sweden. 8. Berkeley, M. J. Jour. Roy. Hort. Soc. London 9: 48. 1854. Describes a new species, Ascomyces bullatus Berk, on pear leaves. 9. Be:rkeley, M. J. Introduction to Cryptogamic Botany. London. 1857. Figures (p. 284, fig. 66c) without description, Ascomyces bullatiis Berk, and describes a deformation of peach leaves as due to another species of this genus. 10. Berkeiley, M. J. Outlines of British Fungology. 1860. Describes (p. 376) briefly the genus Ascomyces and lists four species occur- ring in England: 1. Ascornyces bullatus Berk, on pear leaves, 2. A. deformans Berk, on peach leaves, "causing one form of blister," 3. A. trientalis Berk, on leaves of Tilia europaea L., 4. A. juglandis Berk, on walnut leaves. The two last-named forms did not prove subsequently to be members of the Taphrinaceae. A. juglandis is definitely excluded by Giesenhagen (94). Figures (Plate 1, fig. 9) Ascosporiwn deformans (sic). 11. FucKEL, L. Enumeratio fungorum Nassoviae. Jahrb. d. Ver. f. Naturk. in Herzogt. Nassau 15: 1-123. 1860. Describes (p. 29, No. 189) the genus Exoascus, and the species E. pruni Fkl. on fruits of Prunus dom,estica L. and Prunus spinosa L. Also figures (fig. 26) this species. Lists (p. 30, No. 200) Taphrina alnea Schmidt on leaves of Alnus glulinosa Medic. 12. Westendorp, G. D. Sur quelques cryptogames nouvelles ou inedites pour la flore beige, septieme notice. Bull. d. 1. Acad. Roy. d. Sci. d. Lettr. et. d. Beaux-arts d. belgique. 2e ser. ii; 644-660. 1861. Describes (p.. 655) Ascomyces tosqidnetii West, on leaves of Alnus glutinosa Medic. 13. B.ARY, A. de. Beitrage zur Morphologic und Physiologie der Pilze. II. Exoascus pruni und die taschen oder narren der pflaumenbaume. Abhandl. d. Senkenberg. Naturforsch. Ges. 5: 169-191. 1865. An account of Exoascus pruni Fkl. on fruits of Prunus domestica L., P. spinosa L., and P. padus L. In spite of smaller asci of the form on P. padus 116 The University Science Bulletin refers it to the same species. Gives a full account of the pathological morphol- ogy and histology of the diseased fruit. Describes ejection of spores, and bud- ding of ejected spores in water, sugar solution, and yeast decoction. The "sprossen" derived from ascospores by budding do not cause alcoholic fermen- tation. The "sprossen" are believed to give rise (in some manner) to new infections. 14. TuLASNE, L. R. Super Friesiano Taphrinarum genere, et Acyptolospore Mazeriana, accedente Ustilaginis marinae. Ann. d. Sci. Nat. 5e Ser. Botanique. 6: 122-136. 1866. Revises Fries' description of the genus Taphrina and includes the following species (the author-citation following each species name is that which would result from this revision) : I. Taphrinae polysporae 1. Taphrina aurea Fr. 2. Taphrina caerulescens (Mont, et Desm.) Tul. II. Taphrinae octospoi-ae or Exoasci 3. Taphrina bullata (Berk.) Tul. 4. Taphrina deformans (Berk.) Tul. 5. Taphrina pnmi (Fkl.) Tul. 6. Taphrina alnitorqua (Westend.) Tul. Ascomyces and Exoascus thus become synonyms of Taphrina, although no statement is made that forms previously described under Ascomyces are known to possess mycelium. A quite unwarranted procedure is the change of the specific name of the fungus described by Westendorp as Ascomyces tosquinetii (Westend.) Tul. It is to be noted that the species name tosquinetii is later restored by Sadebeck (73). 15. FucKEL, L. Symbolae Mycologicae. Beitrage zur kenntnis der rheini- schen pilze. Jahrb. d. Nassau. Ver. fur Naturk. 23 & 24: 1-459. 1869 and 1870. Refuses (p. 252) to accept the genus Taphrina as revised by Tulasne, and recognizes two genera, Ascomyces with many-spored asci, and Exoascus with eight-spored asci. Fuckel here is guilty of two errors : First in refusing to concede the priority of Taphrina over Exoascus; second, in ignoring the character on which Desmazieres erected the genus Ascomyces i. e.: the absence of mycelium. Fuckel's species are as follows : 1. Exoascus pruni Fkl. on fruits of Prunus domestica L., P. spinosa L., and P. padus L. 2. E. deformans (Berk.) Fkl. a. forma pcrsicac on leaves of Persica vulgaris Mill. {Prunus persica [L.] Stokes.) b. forma cerasi on leaves of Cerasus aviiim {Prumis avixim L.) 3. E. alni (de By.) Fkl. on leaves of Alnu,s glutinosa Medic. This last-named species Fuckel describes as new, calling it Exoascus alni de Bary in litt, but recognizes its previous description by Westendorp since he gives as synonyms Taphrina tosquinetii Westend. and T. alnitorqua Tul. Ac- cording to modern standards of nomemclature this fungus must be called Taphrina tosquinetii (Westend.) Tul. Mix: The Genus Taphrina I 117 16. KiJHN, J. Exoascv^ alnitorqua var. alni-incanae J. Kiihn on fruits of Alnics incana Willd. In Rabenhorst, L. G. Winter, and 0. Pazschke. Fungi Europaei. Cent. 22, No. 1616. Kiihn here distinguishes for the first time the fungus on carpels of alder which later became recognized as a separate species, T. alni-incanae (Kiihn) Magn. 17. FucKEL, L. Symbolae Mycologicae. Beitrage zur kenntnis der rheini- schen pilze. Zweiter Nachtrag. Jahrb. d. Nassau. Ver. f. Naturk. 27 and 28: 1-99. 1873 and 1874. Under the genus Exoascus describes a new variety, E. hidlatus (Berk) Fkl. var. crataegi Fkl. on leaves of Crataegus oxyacantha L., and two new species, E. ulmi Fkl. on under sides of leaves of Ulmus campesiris L. and E. betulae Fkl. on leaves of Betula alba. L. 18. Thumen, F. a'on. Eine neue Protomyces species. Hedwigia 13: 97-98. 1874. This note concerning another fungus mentions the collection of a form on leaves of Populus pymmidalis (P. alba L. var. pyramidalis Bunge) and P. nigra L. closely resembling Exoascus alni de By. and here named without de- scription E. populi Thm. 19. Magnus, P. Kurze notiz iiber Protomyces pachydermus. Hedwigia 13: 113-114. 1874. States that the fungus called Exoascus populi by von Thumen is Taphrina aurea Fr. 20. ThIjmen, F. von. Nochmals Protomyces pachydermus. Hedwigia 13: 149. 1874. Objects that Magnus has not seen his Exoascus populi nor a description of it, and that even if it proves identical with Taphrina aurea Fr. the genus name Taphrina has not been recognized by most mycologists and is unsuitable for a fungus, having been previously used for "Pili degenerati." Prefers to recognize the genus Exoascus for eight-spored forms and Ascomyces for many-spored forms, thus following Fuckel. 21. Magnus, P. Ascomyces Tosquinetii Westendorp. Hedwigia 13: 135- 136. 1874. Decides that this fungus has no mycelium, each ascus, a plant by itself, arising in and growing out from an epidermal cell of the host. The genus Ascomyces is quite distinct from Exoascus in which the asci arise from a sub- cuticular hymenium. Taphrina aurea also lacks mycelium, but the roots of the asci are between the epidermal cells. Another Taphrina is mentioned as oc- curring on fruits of Pojmlus tremula L. and P. alba L. In this fungus the bases of the asci are inserted even farther between the epidermal cells. The form on fruits of Populus is perhaps the species later described (56) as T. rhizophora Johans. The genus Taphrina is thus distinguished from the other two genera by the origin of the asci and by the fact that in Taphrina the asci are many-spored. 22. Magnus, P. Eine bemerkung zu Exoascus populi Thm. Hedwigia 14: 1-3. 1875. 118 The University Science Bulletin Having seen a specimen of von Thiimen's Exoascus populi decides that it is Taphrina aurea Fr. States that it is uncertain whether the genus Ascomyces (as erected by Desmazieres and Montagne) will eventually prove to be dis- tinct from Taphrina. If it does not, the alder fungus, A. tosquinetii, represents a true genus type, and in that event Magnus proposes the genus name Endo- ascus. 23. Magnus, P. Zur Naturgeschichte der Taphrina aurea Pers. Hedwigia U: 97-96. 1875. States that asci of T. aurea arise from a richly branched mycelium which grows between epidermal cells. Nearly every cell of this mycelium bears an ascus, leaving only rarely sterile cells between asci. 24. SoROKiNB, N. Quelques mots sur I'Ascomyces polysporus. Ann. d. Sci. Nat. 6e ser. Botanique. '4: 72. 1876. Distinguishes three genera, Exoascus with intercellular mycelium, Taphrina with subcuticular mycelium, Ascomyces with no mycelium. Describes a new species, Ascomyces polysporus Sorok. on leaves of Acer tataricum L. 25. Berkeley, M. J., and C. E. Broome. Notices of British Fungi. Ann. and Mag. Nat. Hist. 17: 129-145. 1876. Mentions (p. 144), with meager description, a new species, Ascomyces alni Berk, and Br., deforming the female catkins of alder. Nothing is given which would distinguish this fungus from the fungus pre- viously collected by Kuhn (16) and ascribed to "Exoascus alnitorqua Tul." As Tulasne's Taphrina alnitorqua is a renaming of Ascomyces tosquinetii West- end., both Kiihn's and Berkeley's fungi belong to T. tosquineti (Westend.) Tul. The first real separation of the form on catkins from that on leaves was made by Sadebeck (61). It was renamed by Magnus (70) Taphrina alni4ncanae (Kiihn) Magn. Berkeley also mentions the occurrence on sloe of "A. pruni (Fkl.) Berk, and Br." 26. CooKE, M. C. Ravenel's American Fungi. Grevillea 6: 129-146. 1878. Describes as new Ascomyces quercus Cke. on leaves of Quercus cinerea Michx., from South Carolina. There is nothing to distinguish this fungus from T. caerulescens and it was later so called by Robinson (55) and by Farlow (59). 27. Farlow, W. G. List of fungi found in the vicinity of Boston. Bussey Institute Bull. 2: 224-252. 1878. Mentions the occun-ence near Boston of Taphrina aurea Fr. on catkins of Populus grandidentata Michx., and T. alnitorqua Tul. on catkins of Alnus. 28. Rathay, E. Ueber die von Exoascusarten hervorgerufenen degenera- tionen der laubtriebe einiger Amygdaleen. Sitzungsber. d. Math. Naturw. CI. d. K. Acad. d. Wiss. 77: 67-82. 1878. Gives an account of careful studies which show that the mycelium of Taphrina pruni is not perennial. Describes a new form on almond very similar to T. deformans. Does not distinguish it from the latter. Mix: The Genus Taphrina I 119 29. Cooke, M. C, and H. W. Harkness. Calif omian fungi. Grevillea 9: 6-9. 1880. Gives a meager description of a new fungus, Ascomyces julgens Cke. and Hark, on leaves of Arctostaphylos jmngens HBK. The asci are reported to be dissolved and there is nothing to indicate that the fungus belongs to the Taphrinales. It was later pronounced by Harkness (California Acad. Sci. Bull, i : 256-268, 1886) to be an "aphidian gall," and was excluded from the genus by Giesenhagen (94). 30. Fr.^nk, a. Die krankheiten der pflanzen. Breslau. 1880. pp. 521-524. Gives descriptions of the following: Ascomyces tosquinetii Westend. {Taphrina alnitorqua Tul., Exaoscus alni de B^^), on Almis glutinosa Medic, A. bullatus on Crataegus oxyacantha L., Taphrina aurea Fr. on Populus nigra L., Exoasctbs pmni Fkl. on Primus domestica L., P. spinosa L., and P. padiis L., E. deformans Fkl. on peach. T. aurea is described as having asci with stalk cells. Characterizes three genera as follows: 1. Ascomyces. No mycelium, each plant consisting of an ascus in the host epidermal cell, eight spores in an ascus, increasing by budding. 2. Taphrina. Again each plant is an ascus, but possesses a simple my- celium, developing between the epidermal cells of the host. Each ascus has numerous spores. 3. Exoascus. Mycelium well developed, intercellular. Each ascus has 6 to 8 spores. 31. R.^THAY, E. Vorliiufige mittheilung iiber die hexenbesen der kirsch- baume und iiber Exoascus Wiesneri Rathay. Oesterreich Bot. Zeitschr. 30: 225. 1880. The fungus causing witches' broom of cherry previously known as Exoascus deformans var. cerasi Fkl. is specifically distinct and is therefore renamed Exoascus wiesneri Rathay. Occurs on Prunus avium L., P. cerasv^ L., and P. chamaecerasus Jacq. (P. fruticosa Pall.) Its mycelium is perennial in the diseased branches. 32. Thumen, F. von. Diagnosen zu Thiimens Mycotheca Universalis Centur. XIII bis XV. Flora 63: 312-322, 323-332. 1880. No. 1366 (issued 1879) is Exoascus alni de B3\ var. strobilimis Thm. on catkins of Alnus. 33. Thumen, F. vox. Zwei neue blattbewohnende Ascomyceten der flora von Wien. Verhandl. d. k. k. Zool.-Bot. Ges. in Wien. 29: 523-524. 1880. Describes as new Ascomyces alutaceus Thm. on leaves of Qucrcus pubescens Willd. This fungus is apparently identical with T. caerulescens and was re- duced to sj'nonymy by Giesenhagen (94). 34. S.\ccARjDO, P. A. Fungi novi ex herbario professoris doct. P. Magnus Berolinensis. Michelia 1: 117-132. 1880. Describes as a doubtful species Taphrina candicans Sacc. on Teucrium chamaedrys L. This fungus was apparently never recognized as a Taphrina by later authors and it was excluded from the genus by Giesenhagen (94). 35. RosTRUP, E. Mykologische Notizen. Bot. Centralbl. 5: 153-154. 1881. 120 The University Science Bulletin Names a new species, Exoascus carpini Rostr., causing witches' brooms of Car-pinus hetulus L., but gives no description beyond that of the effect on the host. 36. Sadebeck, R. Beobachtungen und untersuchungen Uber die pilz vegeta- tion in der umgegend von Hamburg. Festschrift, seitens der botanischen gesell- schaft zu Hamburg Sr. Magnificenz Herrn Burgermeister Dr. Kirchenpauer zur feier seines funfzig-jahrigen Doctor-Jubilaums gewidmet. Hamburg 1881. Notes on Ascomyces tosquinetii Westend., on a summer form of this causing white spots on leaves of Abms glutinosa Medic, on a form attacking the female catkins of the same, on a similar form on Betula alba L., and on Ascomyces buUatus Berk, on Crataegus sp. Describes budding of spores of A. tosquinetii in water and sugar solutions and production of minute amounts of alcohol by budding spores. 37. Rathay, E. Uber die hexenbesen der kirschbaume and iiber Exoascus wiesneri n. sp. Sitzungsber. d. k. Acad. d. Wi.ss. Math-naturw. Classe. 83: 267-288. 1881. Reports perennial mycelium for E. wiesneri Rathay, causing witches' brooms of Prunus avium L., P. chamaecerasus Jacq., P. cerasus L., but maintains that E. deformans and E. pruni do not possess perennial mycelium. This is one of his reasons for considering the form on cherry a distinct species. 38. RosTRUP, E. Sygdomme hos skovtraerne, foraarsagede af ikke-rustagtige snyltesvampe. II. LOvtraeer. Tidsskr. for Skovbrug. 4:113-206. 1881. Characterizes Ascomyces as lacking mycelium, and having eight-spored asci, Taphrina as also without mycelium, but having many-spored asci, Exoascus aa having intercellular mycelium with eight-spored asci. Mentions Ascomyces tosquinetii Wes-tend. as attacking the fruits as well as the leaves of Abius glutinosa Medic. States that the form has been previously named A. tos- quineti var. strobilina Thm. by von Thlimen in Mycotheca universalis. The date (1879) of von Thijmen's name for the fungus on alder fruits is later than that of Kiihn's (16) var. alni-incanae. 39. IvuTSOMiTOPULos, D. Beitrag zur kenntniss der Exoascus der kirscll^ baume. Sitzungsber. d. Phys-Med. Soc. zu Erlangen. 15: 1-11. 1SS2. Reports perennial mycelium for the fungus causing witches' broom of cherry. States that this fungus should not be called E. vnesneri since it had already been recognized as distinct by Fuckel (15) under the name E. cerasi. Fuckel made this fungus a variety: E. deformans cerasi. 40. Sade:beck, R. Ueber die entwickelungsgeschichte der pilzgattung Exo- ascus und die durch einige arten der letzteren verursachten baumkrankheiten. Bot. Centralbl. 12: 179-181. 1882. Distinguishes two species on leaves of alder (Alniis glutinosa Medic and A. incana Willd.) : one, which he calls Exoascus alni, affecting whole shoot-sys- tems in the spring but later on causing separate spots on the leaves, and another, which he leaves unnamed, causing yellow spots on the under sides of leaves, and having asci filled with yellow protoplasm like those of E. populi {Taph- rina aurea). Mix: The Genus Taphrixa I 121 Describes the formation of ascogenous cells and elongation of asci by E. alni and other species, also the presence of sterile cells (which later die) among the ascogenous cells of the former. Reports perennial mycelium as occurring in the one-year twigs in the case of E. bullatus and in older parts in the case of forms causing witches' brooms, such as E. carpini. Reports successful in- oculation of pear leaves with spores of E. bullatus and of alder leaves with E. alni. In the latter case observed penetration. Objects to Magnus' use of the name Ascomyces for the alder parasite. 41. Saccardo, p. a. Fungi gallici, Series II. Michelia 2: 39-135. 1882. Describes (p. 86, No. 727) a new species, Exoascus campestris Sacc. on leaves of Ulmus campestris L. This fungus is synonj^mous with Taphrina ulmi (Fkl.) Sadeb. according to Sadebeck (80) and Giesenhagen (94). 42. Sacc.^rdo, p. a. Fungi Dalmatici Pauci. Michelia 2: 150-153. 1882. Describes Ascomyces alutaceus Thm. in litt. on leaves of Quercus robor L. This fungus had previously been described by vonThiimen (33). 43. RosTRrp, E. Fortsatte unders0gelser over snyltesvampes angreb paa skovtraerne. Tidsskr. for Skovbrug. 6: 199-300. 1883. Describes a new species, Taphrina betulina Rostr. on Betula alba L., and B. odorata Bechst. 44. Farlow, W. G. Notes on some species in the third and eleventh cen- turies of Ellis' North American Fungi. Proc. Amer. Acad. Arts and Sci. 18: 65-85. 1883. Mentions Ascomyces tosquinelii and Taphrina aurea. Describes a new species, Taphrina flava Farlow on Betula alba L. Following Magnus dis- tinguishes Taphrina as having subcuticular mycelium and polysporic asci with rootlike extremities between epidermal cells, and Exoascus with intercellular mycelium and 8-spored asci. T. flava, however, has some subepidermal mycelium. Mentions, also, forms on Pmnus serotina Ehrh. (which he thinks may be E. pruni) on Potcntilla canadensis L., and on Rhus copallina L. (This, he believes, may be a variety of E. deformans.) The fungus here called by Far- low Taphrina aurea occurred on carpels and was later pronounced by Sadebeck (73) to be T. johansonii. 45. Peck, C. H. Report of the Botanist. New York State Mus. Nat. Hist. Ann. Rept. 33: 17-72. 1883. Reports occurrence of "Exoascus pruni" on fruits of Prunm pumila L. and P. americana Marsh. This is apparently the first report of the fungus later (80) described as Exoascus communis Sadeb. 46. Trele-ase, W. Prehminary list of the parasitic fungi of Wisconsin. Trans. Wisconsin Acad. Sci. 6: 106-144. 1884. Records the occurrence of Exoascus pruni Fkl. on fruits and of Ascomyces caerulescens Mont, and Desm. on Quescus coccinea Muensch. and Q. rubra L. These are new hosts for T. caerulescens. 47. Sadebeck, R. Untersuchungen liber die pilzgattung Exoascus und die durch dieselbe um Hamburg hervorgerufenen baumkrankheiten. Jahrb. d. Hamburg. Wissensch. Anst. 1: 93-124. 1884. 122 The University Science Bulletin This paper contains a monographic account of the species known to date, including new species. Only the genus Exoascus is recognized, but in sub- dividing the genus certain characters are used which were later made the basis (80) of separating Exoascus and Taphrina. The following is a condensed summary of Sadebeck's classification. A. Mycelium perennial within shoots, fertile hyphae completely used up in the formation of asci, stalk cells present. 1. Exoascus priini Fkl. 2. E. bullatus (Berk. & Br.) Fkl. 3. E. insititiae Sadeb. (A new species.) 4. E. deformans (Berk.) Fkl. B. Mycelium perennial beneath cuticle and confined to that location, a. Fertile hyphae completely used up in formation of asci. a.a. Ascus with stalk cell. 5. E. alnitorquus (Tul.) Sadeb. 6. E. turgidus Sadeb. (A new species.) 7. E. flavus Sadeb. (A new .species.) 8. E. betidae Fkl. b.b. Ascus lacking a stalk cell. 9. E. aureus (Pers.) Sadeb. 10. E. coerulescens (Desm. and Mont.) Sadeb. 11. E. carpini Rostr. b. Not all of the fertile hyphae used in the formation of asci, the latter consequently more or less scattered. 12. E. epiphyllus Sadeb. (A new species.) 13. E. ulmi Fkl. Of the new species described E. alnitorquus'^ on female catkins of Alnus glutinosa Medic, and A. incana Willd., and on twigs and leaves of A. glutinosa X incana Krause {A hyhrida A. Br.), is split off from the species previously known as Asconiyces tosquinetii Westend. {Exoascus alni de By.) ; E. flavus (also considered to have been included in the foregoing) causes yellow spots on leaves of Alnus glutinosa, and its asci have yellow contents; E. epiphyllus swells and curls the leaves of Almis incana; E. insititiae, causing witches' broom on Prunus insititia L., is perhaps a form previously considered to belong to E. pruni; and E. turgidus seems to be a renaming of T. betulina Rostrup. In this paper Sadebeck also gives a circumstantial account of the occurrence and development of perennial mycelium in E. alnitorquus {T. tosquinetii) , the cultivation of this form in artificial media, successful inoculations and observa- tion of penetration (the germ tube entering the radial wall between two epi- dermal cells) . He also figures nuclear division in a young ascus. E. idmi is also stated to have perennial mycelium, a statement which Sade- beck (80) later retracts. 48. FiscH, C. Ueber die pilzgattung Ascomyces. Bot. Zeit. 4^: 34-39, 49- 59. 1885. 1. This is essentially a new species since T. alnitorqua Tul. and E. alni de By. had not been previously recognized as distinct from T. tosquinetii. Mix: The Genus Taphrina I 123 Using the genus name Ascomyces in the sense of Magnus (21), i.e.: for forms lacking mycelium, Fisch renames Ascottiyccs tosquinetii Westend. as A. endoqenus Fisch. This is done because the name A. tosquinetii has been ap- plied to several species, Sadebeck having described three species from alder. A. endogeniis is stated to have no mycelium, the ascus being developed within an epidermal cell of the host. Infection was obtained when spores were sown on leaves in plum decoction. Penetration is figured and shows what was long interpreted as production of a secondary spore or appressorium, but may possibly be a case of copulation of conidia such as was reported much later by Wieben (176) for Taphrina epi- phylla. Fisch 's fungus, however, must have been either T. tosquinetii or T. sadeheckii since the host was Alnus glutinosa. Copulation of conidia has not been reported for these species. Fisch also reports having produced infection with Exoascus epiphyllus (T. epiphylla) and having seen penetration. A. endogenwi and E. epiphyllus are said to occur on the same host, but never on the same shoot. 49. Fisch, E. Ueber Exoascus aceris Linh. Bot. Centralbl. 22: 126-127. 1885. Besides the fungus described by Sorokine (24) as A.icomyces polysporus o'n Acer tataricum L. there is another on the same host, distributed by Linhart in Fungi Hungarici. This is described as Exoascus aceris Linh. It belongs to those forms in which the subcuticular mycelium all divides up into ascus- producing cells, leaving no sterile members between. These two fungi are considered to be the same species, T. polyspora (Sorok.) Johans. by Johanson (52), Sadebeck (73), and Giesenhagen (94). 50. RosTRUP, E. Om nogle of Snyltesvampe foraarsagede hos blomster planter. Bot. Tidsskr. U: 230-243. 1885. Names, without description, a new species, Taphrina umbclliferarum Rostr. on Hcracleum s-phondyliiun L., and Peucedanum palustre Much. This fungus was subsequently removed from the Taphrinaceae by Juel (114), and placed in the genus Taphridium of the Protomycetaceae. Also describes a new species, Taphrina tormentillae Rostr. on Tormentilla erecta (PotentiUa silvcstris Neck.). This fungus was mentioned, though not described, by Farlow (44). For this reason it was named by Johanson (52) Taphrina potentillae (Farl.) Johans. 51. Peck, C. H. Report of the Botanist. New York State Mus. Nat. Hist. Ann. Rept. 89: 30-73. 1886. Describes a new species, Ascomyces extensvs Pk., on leaves of Quercus mac- rocarpa Michx. This fungus was later referred by Farlow (590 to Taphrina caerulcscens (Mont, and Dsm.) Tul. 52. Johanson, C. J. Om svampslagtet Taphrina och dithorande svenska arter. Ofvers of Kongl. Svensk. Vetensk. Akad. Forhandl. 1SS5: 29-i7. 1886. On the basis of priority unites all species of Taphrina, Exoascus, and As- comyces into one genus, Taphrina. Describes as new Taphrina nana Johans. on Bctula nana L., T. sadeheckii Johans. on Alnv^ glutinosa Medic, T. sade- heckii var. borealis Johans. on Alnus incana Willd. in lower alpine regions, T. 124 The University Science Bulletin carnea Johans. on Betula odorata Bechst., B. nana L., and B. intermedia Thomas. Notes occurrence in Sweden and Denmark of T. potentillae (Far- low) Johans. on PotentiUa tormentilla L., and P. geoides L. Gives a descrip- tion of Taphrina polyspora Sorok. as it occurs on Acer tataricum L. in Sweden, and states that its asci agree in form and size with those of Exoascus aceris Linh. T. sadebeckii Johans. is a renaming of Exoascus flavus Sadeb., the latter species name having been preempted by Farlow (44). 53. Sadebeck, R. Ueber die im ascus der Exoasceen stattfindende en- twickehing der inhaltsmassen. Sitzungsber. d. Ges. f. Bot. in Hamburg. Bot. Centralbl. 25: 123-125. 1886. Describes ascus formation in Exoascus flavus Sadeb. {Taphrina sadebeckii Johans.) and E. alnitorquus (Tul.) Sadeb. A globose uninucleate ascogenous cell elongates to a cylinder, its nucleus divides by mitosis, a septum between these two nuclei cuts off the stalk cell, the ascus-nucleus then divides rapidly to form 2, 4, and 8 nuclei, around these the spores form. 54. Meehan, T. Formation of crow's nest branches in the cherry tree. Acad. Nat. Sci. Philadelphia Proc. 1886: 273-274. 1887. Reports witches' brooms on "wild cherry" (species not named) due to a fungus identified by Farlow as Exoascus Wilsneri (sic). 55. Robinson, B. L. Notes on the genus Taphrina. Ann. Bot. 1: 163-176. 1887. Agrees with Sadebeck (47) that a single genus should be recognized, but with Johanson (52) that it should be called Taphrina. Gives the following list of species known to occur in America: Taphrina pruni (Fkl.) Tub, T. deformans (Berk.) Tub, T. piirpurascens Robinson, T. potentillae (Farl.) Johans., T. flava Farl., T. alnitorqua Tub, T. caerulescens (Mont, and Desm.) Tul. T. pruni is known to occur commonly on Primus domestica L. A closely similar and probably identical form occurs on fniits of P. maritima Wang., P. virginiana L., and P. serotina Ehrh. A form resembling T. deformans has been observed on leaves of cheriy (see 54) and on leaves and twigs of P. serotina. The new species, T. pwpurascejis, occurs on leaves of Rhus copallina L. T. flava occurs on Betula alba L. var. populijolia Spach., and B. papyracea Ait. T. alnitorqv/i occurs on alder catkins, but has not been found on leaves. T. aurea affects catkins of Populus gi'andidentata Michx. Ascomyces quercus Cke. (26) is synonymous with T. caerulescens. 56. JoH.ANSEN, C. L. Studier ofver svampslagtet Taphrina. Bihang till Kongl. Svensk. Vetensk. Akad. Handlingar 13: 3-28. 1887. This paper reports careful investigations which show that T. carnea Johans. and T. sadebeckii Johans. do not possess perennial mycelium. T. betulae Fkl., though not investigated, is probably similar in this respect. These forms are in contrast to T. bctidina Rostr., T. alnitorqua Tub, and T. borealis Johans., whose mycelium is perennial in the buds of the host. It is suggested that in both perennating and non-perennating forms new infections arise from over-winter- ing bud conidia. New species described are: Taphrina alpina Johans. on Betula nana L., T. bacteriosperma Johans. on the same host, and T. rhizophora Mix: The Genus Taphrina I 125 Johans. on Populus alba L., and P.'tremula L. Descriptions are also given of T. filicina Rostr. on Polystichum sjnnulosum (Aspidium spinulosum) (O. F. Miill) Sw., of T. aurea Fries, and T. rhizophora Johans. This is the first care- ful description of T. aurea. (Sadebeck's [47] description was based on the form occurring on the fruits and this, as will be seen, is T. rhizophora.) It is pointed out that the ascus of this fungus is provided with a stalk cell, a fact not previously recognized except by Frank (30). The new species, T. rhizo- phora, is distinguished from the preceding by lack of a stalk cell and by a narrowed basal portion of the ascus extending inward between epidermal cells of the host. T. aurea occurs on leaves of Populus nigra L. and P. pyramidalis (P. alba L. var. pyramidalis Bunge.), while T. rhizophora deforms fruits of P. alba L. and P. tremula L. The latter species occurs in North America on fruits of P. tremuloides Michx. (Ellis and Everhart, North America Fungi, No. 1885). T. sadebcckii Jolians. var. boreali-s Johans. is raised to specific rank, becoming T. borealis Johans. A table is presented showing the world distribu- tion of the 21 Swedish species, together with a discussion of this distribution and of the antiquity of certain species. T. bacteriosperma is thought to be very old. The species on P. tremuloides was later referred by Patterson (92) and Sadebeck (93) to Taphrina johansonii Sadeb. 57. Knowles, E. L. The "curl" of peach leaves: a study of the abnormal structure induced by Exoascus deformans. Bot. Gaz. 12: 261-218. 1887. A description of the histological changes induced in the peach leaf by Taphrina deformans. 58. Peck, C. H. Report of the Botanist. New York State Mus. Nat. Hist. Ann. Rept. 40: 39-77. 1887. Describes two new species, Ascom,yces letijer Pk. on leaves of Acer spicatum Lam., and A. rubrobrunneus Pk. on leaves of Quercus rubra L. 59. Farlow, W. G. a provisional host-index of the fungi of the United States. Cambridge, 1888. Gives a list with synonyms of all American species of Taphrina (Ascomyces, Exoascus) so far knowoi. Taphrina alnitorqua auct. Amer. on catkins of AlmLS incana Willd. and A. rubra Bong, is referred to T. alni-incanae (Klihn) Mag- nus, and T. aurea auct. Amer. on fruits of Popuhcs jremontii Watson, P. grandidentata Michx., P. pyramidalis Roz., and P. tremuloides Michx. to T. rhizophora Johans. Ascomyces quercus Cke. on Quercus alba L., Q. cinerea Michx., Q. coccinea Wang., Q. douglasii Hook and Arn., Q. laurifolia Michx., and Ascomyces extensv^ Pk., on Q. macrocarpa Michx. are referred to Taphrina caerulescens (Mont, and Desm.) Tul. Ascomyces rubrobrunneus Pk. on Quercus rubra L. is recognized as a distinct species. 60. Massalongo, C. Uber eine neue species von Taphrina. Bot. Centralbl. 34: 389-390. 1888. Describes a new species, Taphrina ostryae Massal., on leaves of Ostrya carpinifolia Scop. 61. Sadebeck, R. Neue untersuchungen liber einige krankheitsformen von Alnus incana und glutinosa. Sitzungsber. d. Ges. f. Bot. zu Hamburg. 4-' 90. 1888. 126 The University Science Bulletin States that infection experiments have shown that E. epiphyllus Sadeb. {T. epiphylla [Sadeb.] Johans.) causes witches' brooms on twigs of Alnus incana. Claims that T. borealis Johans. is not distinct from E. epiphyllus (T. epiphylla). The latter species and E. alnitorquus {T. alnitorqua Tul.) may infect the same leaf of Alnus glutinosa Medic, and their asci may be inter- mingled. Yellow spots on leaves of Alnus are caused by E. sadcbeckii {T. sndcbeckii Johans.). Another fungus deforms the pistillate aments of A. glutinosa and A. incana. This is named (without adequate description) a new species, Exoascus amentorum Sadeb. Its asci in form and size and in lack of stalk cell resemble those of Ascomyces endogenus Fisch. 62. TuBEUF, C von. Neue parasitare pilze aus dem Bayerischen walde. 3. Hexenbesen auf Alnus incana. Beitriige zur kenntniss der baumkrankeiten. Berlin. 1888. pp. 37-40. Describes and figures a witches' broom on Alnus incana Willd. as caused by Taphrina borealis Johanson. 63. Briosi, G. Elenco delle ricerche fatte al laboratorio di botanica crit- togamica di Pavia nei mesi settembre e ottobre 1889. Ministerio di Agricolt. Industre, Commercio. Bull, di Notize agrarie. 11: 2228-2231. 1889. Reports a new host, Quescus cerris L., for Taphrina caerulescens. 64. Britton, N. L. Catalogue of plants found in New Jersey. Final Report State Geologist. 2: 28-642. 1889. Mentions (p. 507) several species of Taphrina occurring in New Jersey. Of these T. aurea Fr. on leaves of Populus tremuloides Michx. is a new record for eastern North America. The same fungus had been previously reported from California by Harkness (California Acad. Sci. Bull. 1. 1896) as occurring on P. dilatata Ait., and P. jremontii Watson. 65. Massalongo, C. Nova species e genere Taphrina. Nuov. Giorn. Bot. Ital. 21: 422-423. 1889. Describes a new species, Taphrina oreoselini Massal. on Peucedanum oreosclinum Moench. This species is quite similar to T. umbelliferarum Rostr. found by Rostrup (50) on Heracleum sphondylium L., and Peucedanum palu^tre Moench. Both these species were later placed by Juel (114) in the genus Taphridium of the Protomycetaceae. 66. Mayr, H. Die waldungen von Nordamerika, ihre holzarten, deren anbaufiihigkeit und forstliche werth fur Europa im allgemeinen und Deutsch- land inbesondere. Miinchen (Rieger). 1889. Mentions the occurrence of a witches' broom on Quercus lobata Nee. Sug- gests the possibility of this being caused by "Exoascus Quercus lobatae n. sp. (?)." No fungus was seen. 67. Passerini, G. Diagnosi di funghi nuovi, Nota IV. Atti d. R. Accad. d. Lincei. Ser. 4. 6: 457-470. 1889. Describes a new species, Exoascus aemiliae Passer, on Ccltis australis L. Although the date on the title page is 1889, this volume of the proceedings must have actually been published later, since the session of the Academy at which this report was made was that of March, 1890. Giesenhagen (94) re- duces this fungus to synonymy as Taphrina celtis Sadeb. Mix: The Genus Taphrina I 127 68. Harknbss, H. W. Curled leaf. Zoe 1: 87-88. 1890. Describes a witches' broom on Aesculus californica caused by an Ascomyces which may be identical with Ascomyces deformans (sic). This fungus wa3 later named by Patterson (92) Exoascus acsculi (Ell. and Ev.) Patterson. 69. Kruch, O. Sopra un caso di deformazione (Scopazzo) dei rami dell 'Elce. Malpighia 4: 424-430. 1890. Describes the effect on the host of a newly found species of Taphrina caus- ing witches' brooms on Quercus ilex L. 70. Magnus, P. Bemerkung liber die benennung zweier auf Alnus lebender Taphrina-arten. Hedwigia 29: 23. 1890. In view of the work of Johanson (52) and Sadebeck (47) believes that all species should be placed in one genus, Taphrina. Has examined original ma- terial of Ascomyces tosquinetii Westend. and says it is identical with Taphrina alnitorqua Tul. and should be called T. tosquinetii (Westend.) P. Magnus. Calls attention to the fact that Exoascus amentorum Sadeb. was fii'st de- scribed as E. alnitorqua (Tul.) J. Kiihn, forma alni-incanae J. Kiihn in litt. (16), and later as E. alni de By. var. strobilinus Thm. (von Thiimen, My- cotheca Universalis, No. 1366. 1879). It should be called T. alni-incanae (J. Kiihn) P. Magnus. 71. RosTRUP, E. Taphrinaceae Daniae. Vidensk. Meddel. fra den Naturh. Forening. i Kijbenhavn. ISOO: 246-264. 1890. This paper gives descriptions, with a key for identification and a host index, of twenty species of Taphrina found in Denmark. New hosts are given as follows: for Taphrina pruni (Fkl.) Tul. — Prunus insititia L., for T. crataegi Sadeb. — Crataegus monogyna Jacq., for T. insititiae Sadeb. — Prunus spinosa L., for T. aurea Fr. — Populus monilifera Ait. (P. deltoides Marsh.), for T. bidlata (Berk.) Tul. — Cydonia japonica Hort. {Chaenomeles lagenaria Koidz.). The following species are listed as having mj^celium perennial in the twigs: T. pruni, T. cerasi, T. crataegi, T. deformans, T. insititiae; the following have mycelium perennial in the buds : T. epiphylla, T. ulmi, T. hullata, T. tosquin- eti, T. betulina. Two new species are described, T. githaginis Rostr. on Agrostemma githago L., and T. lutescens Rostr. on Lastraea thelypteris (Aspidium thelypteris (L.) Sw.). 72. TuBEUF, C. VON. Botanische excursionen mit den Studirenden der Forstwissenschaft an der Universitat Miinchen. Allgem. Forst-und Jagdzeit. 66: 25-33. 1890. The witches' brooms on white alder were first described and figured by Tubeuf in 1888 (62). The fungus was named T. sadebeckii var. borealis by Johanson. Sadebeck has since claimed that this fungus is identical with E. epiphyllus {T. epiphylla). No doubt the fungi are identical, but this will necessitate a revision of the description of E. epiphyllus, since Sadebeck has not mentioned its causing witches' brooms nor its possession of perennial mycelium. (See 61.) 73. Sadebeck, R. Kritische untersuchungen liber die durch Taphrina-arten hervorgebrachten baumkrankheiten. Jahrb. d. Hamburg. Wissensch. Anst. 8: 61-95. 1890. 128 The University Science Bulletin In this paper Sadebeck disagrees with Johanson (56) and claims that Taphriva cornea, T. hetulae, and T. sadcbeckii possess perennial mj^celium. Pointing out that strict adherence to priority would call for the use of the name Taphria, (since Fries in 1815 described the poplar fungus as Taphria populina, aurea, changing the genus name to Taphrina in 1825), he accepts the name Taphrina for the united genus containing all forms described as Taphrina, Exoascus, and Ascomyces. Agrees with Johanson that the species Taphrina aurea Fr. should be re- stricted to the form on leaves of Popiilus nigra, but does not agree that this fungus has a well-defined stalk cell. The form on cai-pels of Popnlus alba is T. rhizophora Johans., that on fruits of P. tremula is named as T. johansonii Sadeb. with the description given earlier (47) for Exoascus aureus (T. aurea Fr.). Does not know where the American form on P. tremuloides Michx. belongs. Describes successful inoculation experiments on Alnus incana with asco- spores (unsuccessful with conidia) of T. epiphylla and T. borealis, concluding that the two species are the same. Gives a revised description of T. epiphylla and describes a new variety of the species, T. epiphylla var. rnaculans Sadeb. causing spots on leaves of Alnus glutinosa. Infection experiments with this fungus did not succeed. The fungus which deforms fruits of Alnus incana and A. glutinosa has been variously known as Exoascus alnitorquu^ Tul. forma alni-incanae Kiihn. (16), Ascomyces alni Berk. & Br. (25), E. alni de By. (15), and E. alni de By. var. strobilinu^ Thm. (70). It was earlier (61) raised to specific rank as E. amen- torum Sadeb. Magnus (70) changed the name on basis of priority to T. alni- incanae (J. Kiihn) Magnus. It is now accepted as T. alni-incanae (J. Kiihn) Sadeb., since Magnus and other early authors made no study of the characters of the fungus. An amended description of T. tosquinetii (Westend.) Magnus is given to exclude the preceding fungus (on fruits). T. tosquineti affects leaves and twigs of Alnus glutinosa. The name T. Sadebcckii Johans. for the fungus previously described (47) as E. flavus Sadeb. is accepted. This fungus causes yellow spots on leaves of Alnus glutinosa. Description is given of a new species, Taphrina celtis Sadeb. on leaves of Celtis australis L. The fungus on Crataegus oxyacantha L., formerly (47) included in Taphrina bvllata (Berk.) Tul., is sepai-ated as a new species, T. crataegi Sadeb. Spores of this form caused no infection on pear leaves, while previously (40) spores from pear to pear had caused infection. The diagnosis here given for T. crataegi is that formerly (47) used for T. bulkita. Taphrina bullata as it occurs on pear is now redescribed. Dimensions of asci are different from those given by Tulasne (14), since he employed only dried material. Due to the shrinkage of asci in this species fresh material or material preserved in alcohol is necessary. A new species, T. minor Sadeb. occurring on Prunus chamaecerasus Jacq., is described. Taphrina deformans (Berk.) Fkl. is restricted to the form on peach and perhaps the form on Prunus armcniaca L. A description is given. The fimgus causing witches' brooms on Prunus avium L. and P. ccrasus L., formerly known (15) as a variety of T. deformans, is raised to specific rank, becoming T. cerasi (Fkl.) Sadeb. Mix: The Genus Taphrina I 129 Taphrina insititlae Sadeb., heretofore known as causing twig and leaf de- formations on Prunus insititia L., is now recognized as affecting in the same manner P. domestica L. Taphrina pruni (Fkl.) Tul. is described and stated to occur on fruits of the following: Prunus domestica L., P. padus L., and P. virginiana L. A new species, T. jarlowi Sadeb. is described as occurring on fruits of Prunus serotina Ehrh. in North America. Two lists are given of the 32 known species of Taphrina, one on the basis of hosts affected, the other on the basis of possible relationships in the genus. In this second list the species are arranged in the following groups: I. Ascogenous layer subcuticular. A. Possessing perennial mycelium. B. Lacking perennial mycelium. II. Ascogenous cells intercellular, between epidermal cells or deeper in host. 74. Brefeld, 0. Untersuchungen aus dem gesamtgebiete der Mycologie. IX. Die Hemiasci und die Ascomyceten. MUnster. 1891. Reports culturing in nutrient solutions the budconidia of Taphrina rhizo- phora Johans. (obtained from fruits of Populxis tremula). States that the dried conidia retained their ability to germinate for many months. Similar results were obtained with Exoascus dejormans Berk. Suggests that the genus be divided into Taphrina, with 4 spores in the ascus, and Exoascus with 8. This is a curious suggestion, since all previous work had shown that species of Taphrina (or Exoascus) bore normally 8 spores in an ascus, unless by budding of ascospores the ascus became filled with conidia and hence were many-spored. 75. Vtjillemin, P. L'Exoascus kruchii sp. nov. Rev. Mycol. 13: 141-142. 1892. Describes a new species, Exoascus kruchii Vuill. on Quercus ilex L. The occurrence of this fungus was previously reported by Kruch (69). 76. Massalongo, C. Intorno alia Taphrina polyspora (Sor.) Johans., var. pseudoplatani. Bull. d. Soc. Bot. Ital. 1892:197-199. 1892. Describes a new variety, Taphrina polyspora (Sor.) Johans., var. pseudo- platani Massal. on Acer pscudoplatanus L. 77. GiESENHAGEN, K. Ueber hexenbesen an tropischen farnen. Flora 76:130-156. 1892. From herbarium material collected in Ceylon describes two new species, Taphrina cornu-cervi Giesenhag. on Aspidium aristatum Sw. (Polystichum aristatum Presl.) and T. laurcncia Giesenhag. on Pteris quadriaurita Retz. No spores were present in the asci of either of these fungi. Divides Taphrina into two subgenera ; Eutaphrina, with mycelium intercellular or subcuticular, and Taphrinopsis, with mycelium and asci formed within the epidermal cell of the host. In Eutaphrina are placed T. cornu-cervi and all previously de- scribed species, in Taphrinopsis T. laurcncia. 78. Wakker, J. H. Untersuchungen liber den einfluss parasitischer pilze auf ihre nahrpflanzen. Versuch einer pathologischen anatomie der pflanzen. Jahrb. f. Wissensch. Botanik 2^:529-548. 1892. 9—6037 130 The University Science Bulletin Contains a brief account of the morphological and histological changes in- duced in Primus padus by Taphrina pruni and in Abiiis glutinosa by T. alni- incanae (Exoascus alnitorquus) . 79. Pammbl, L. H. Notes on some fungi common during the season 1892 at Ames, Iowa. Agric. Sci. 7:20-27. 1893. Mentions the occurrence of Taphrina deformans on peach, "T. pruni" on Primus chicasa {P. anffustifolia Marsh.), and P. americana Marsh., and T. autrea on leaves of Populus certinensis Dieck. (P. berolinensis Dipp. [?]. Ac- cording to L. H. Bailey in Cyclopedia of Horticulture, the "Populus certinensis" grown in America may not be correctly named), and P. monilijera Ait. (P. virginiana Fourg.). SO. S.\DEBECK, R. Die parasitischen Exoasceen, eine monographic. Jahrb. d. Hamburg. Wissensch, Anst. iO:5-110. 1893. Objects to Brefeld's division of the genus into Exoascus with 8 spores, and Taphrina with 4. From a study of Taphrina ulvii, Sadebeck finds that many asci are 8-spored while others are only 4-sporcd. A similar variation m spore number was ob.?erved in T. bullata, E. epiphyllus and E. farloivi. Spore num- ber cannot be made the basis of generic distinction. Sadebeck, however, would now divide the genus into three: Exoascus, Taphrina, and Magnusiella. Exoascus has perennial mycelium, deforms shoots as well as leaves, and all of its subcuticular mycelium separates without pre- vious differentiation into ascogenous cells. Taphrina lacks perennial mycelium, causes spots on leaves only, and its subcuticular mycelium differentiates into fertile and sterile cells, the former becoming mother cells of the ascogenous cells, the latter degenerating. Magnusiella forms no ascogenous layer, the asci arising on the ends of branches of the intercellular mycelium. He excludes the genus Ascomyces, which has been characterized by lack of mycelium, believing that Westendorp in describing A. tosquinetii, and Fisch with A. endogenus, were mistaken as to lack of mycelium, and that both had Taphrina sadebeckii Johans. The account of his previous (47) investigations showing that E. tosquinetii possesses perennial mycelium is repeated. It is also stated (without evidence) that mycelium of E. epiphyllus, E. betulinus, E. turgidus, and E. carpini is perennial in the buds. (Later in the paper the earlier [47] statement that T. ulmi has perennial mycelium is corrected.) In these forms the mycelium grows from the bud-anlage into the leaves, where it develops subcuticularly. With E. tosquinetii (Westend.) Sadeb. observed germination of spores and formation of hyphae in artificial culture, but was unable to keep cultures go- ing for very long. He succeeded in obtaining infection with this fungus by inoculating buds and young seedlings. He corrects his former statement (47) that in formation of ascogenous cells by E. tosquinetii there is a differentiation into sterile and fertile hyphae. This statement and the accompanying discus- sion should have referred to T. soidebeckii Johans. By way of contrast three forms causing deformation of shoots of the host {E. pruni, E. rostrupianus, E. tosquinetii) have mycelium, which is perennial in the twigs, and grows intercellularly in the new leaves. The forms listed imder Taphrina do not have perennial mycelium, and these fungi never form hyphae in nutrient solutions. A full description is given of the manner of Mix: The Genus Taphrina I 131 formation of ascogenous cells in Exoascus and Taphrina (see above). The subcuticular habit of mycelium of Taphrina, and the disappearance of the sterile mycelium after formation of ascogenous cells results in no mycelium being left in the host to become perennial. New species described are: Exoascus communis Sadeb. on Primus americana Marsh., P. pumila L., and P. maritima Wang., and E. rostrupianus Sadeb. on Prunus sjdnosa L. A complete host index of known species, a taxonomic account of all known species, with full description of each one, and a table showing world distribu- tion of these species are presented. Twenty-one species of Exoascus, fourteen of Taphrina, and five of Magnusiella are listed. 81. ScHROETER, J. Die Pilze. In Cohn, F. Kryptogamenflora von Schlesien. Vol. 3, pp. 8-15. 1893. Makes two genera, Exoascus with'8-spored asci, and Taphria with asci be- coming many-spored by budding of the ascospores. Gives descriptions of eleven species of Exoascus, and seven species of Taphria (all of which have been previously de.scribed) with notes of their occurrence in Silesia. 82. Hennings, p. Taphrina gilgii P. Hennings et Lindau n. sp., eine neurer parasitischer pilz der Mark. Hedwigia 3;?: 156-157. 1893. Describes a new species, Taphrina gilgii P. Henn. and Lindau, on Prunus cera^is L. Distinguished from T. cerasi by its broader asci. 83. RouMEGUERE, C. Fuugi exsiccati praecipue gallici, LXIIIe Centurie, puqliee avec le concours de MM. Briard, F. Cavara, Eugene Niel, F. Fautrey, R. Ferry, L. Boudier, Lambotte, L. Quelet, C. Raoult, et L. Rolland, et les Reliquiae de Balansa. Revue Mycologique 16:15-25. 1893. In this list of exsiccati No. 6228 is described as a new species, Exoascus marginatus Lamb, and Fautr., occurring on Crataegus oxyacantha L. This specimen was later determined to show only injury due to Erineum mites. See Jaczewski (170.) 84. Mass.alongo. C. Nuova contribuzione alia Micologia Veronese. Mal- pighia 5:97-130. 1894. Describes a new species, Taphrina acericola Massal. on Acer campestris L. Changes T. polyspora Sorok. var. pseudoplatani Massal. to T. acericola (Massal.) var. pseudoplatani Massal. Also reports as ''new" hosts for T. caerulescens (Mont, and Desm.) Tul. Qucrcus ccrris L., Q. pedunculata (Q. robur L.), Q. pubescens Willd. 85. P.\TTERS0N, F. W. Species of Taphrina parasitic on Populus. Bot. Gaz. 19:380. 1894. The fungus occurring in America on fruits of Populus tremuloides Michx. and Populus spp. which has been called Taphrina aurea Fr. and subsequently (56) (59) T. rhizophora Johans. is really T. johansonii Sadeb. T. aurea, which occurs only on leaves, has not heretofore been observed in America. Recently a form differing but slightly from this last named species has been found in Iowa, on leaves of several species of Populus. (See 79.) 86. Atkinson, G. F. Notes on some Exoascaceae of the United States. Bull. Torr. Bot. Club .'/:372-3S0. 1894. 132 The University Science Bulletin Presents, with notes and discussion, a list of fifteen species occurring in the United States. New species are: Exoascics conjusus Atk., deforming fruit and floral envelopes of Phunus uirginiana L., E. loriffipes Atk. on leaves of Pnmus americana Marsh., E. decipiens Atk. on leaves of Pruyius americana Marsh., E. decipiens Atk. var. superficialis Atk. on half grown fruits of P. americana, E. mirabilis Atk. on shoots of Prunus angustifolia Marsh., buds of P. hortulana Bailey, and P. americana Marsh., E. mirabilis Atk. var. tortilis Atk. on fruits of P. angustifolia and P. americana, E. rhizipes Atk. on buds and fruit of Prunus triflora Roxb., E. varius Atk. on leaves of Prunus serotina L., E. cecidomophilus Atk. on galls formed by insect larvae on fruits of Prunus virg-iniaiia L., E. australis Atk. on leaves of Carpinus americana Michx. (C caroliniana Walt.). Exoascus in^ititiae Sadeb. is recorded as occurring on a new host: Prunus pennsylvanica L. Taphrina aurca Fr. is reported on leaves of Populus mo- nilijera Ait. The species on aments of Populus tremxdoides Michx. which has been called T. aurea. has been referred by Farlow (59) to T. rhizophora Johans. Material from Ithaca, New York, examined by Atkinson is said to be T. rhizophora (see 92). Meechan's specimen (54) mentioned by Robinson (55) is reported as Exoascus cerasi (Fkl.) Sadeb. on Prunus avium L. 87. Atkinson, G. F. Leaf curl and plum pockets. Cornell Univ. Agric. Exp. Sta. Bull. 73. 1894. This is an account of the prunicolous species discussed in the preceding paper. 88. CoccoNi, G. Ricerche sullo svillupo evolutive di duo specie nuove di funghi, Lagenidium papillosum ed Exoascus flavo-aureus e sul parasittismo della Phoma uncinulae sull Uncinula adunca Lev. Mem. d. R. Accad. d. Sci. d. Inst. d. Bologna. Ser. 5. 4:187-198. 1894. Describes as a new species, Exoascus flavo-aureus Cocc. on Populus pyram- idalis Roz. (P. alba L., var. pyramidalis Bunge). This fungus is distinguished from T. aurea Fr. by the variability of the basal portion of the ascus. This may be prolonged to a tapering rhizoid between the host epidermal cells. 89. Smith, W. G. Untersuchungen der Morphologie und Anatomie der durch Exoasceen verursachten Spross-und blattdeformationen. Forstlich. Na- turwiss. Zeitschr. 3:420-427, 433-465, 473-482. 1894. A study of the morphological and anatomical changes induced in their various hosts by Taphrina aurea, T. bctulae, T. carnea, T. caerulescens, T. cerasi, T. deformans, T. epiphylla, T. insitiliac, T. minor, T. polyspora, T. pruni, T. tosquinetii, T. turgida (or other species on Bctula verrucosa), and Taphrina sp. on Prunus padus. 90. Dangeard, p. a. La reproduction sexuelle des Ascomycetes. Le Bo- taniste 4e ser. 4:21-61. 1895. An account of the cytology of Taphrina dvjormans. Cells of the inter- cellular mycelium and young ascogenous cells are binucleate. Nuclear fusion occurs in the ascogenous cell. This cell then puts forth a papilla which elongates to become the ascus, the fusion nucleus migrates to the center of the ascus and divided by three successive divisions to form the nuclei for the spores. A septum forms across the base of the ascus, cutting off a basal cell. The septum is considered of no importance. Mix: The Genus Taphrina I 133 91. Ei.iASSON, A. G. Taphrina acerina n. sp. Bihang t. Kongl. Svenska Vetenskaps-Akad. Handl. 20:3: 4: 3-6. 1895. Describes a new species, Taphrina acerinn Eliass. on Acer platanoides L. 92. Patterson, F. W. A study of North American Parasitic Exoasceae. Bull. Lab. Nat. Hist. Univ. Iowa 3: 89-135. 1895. This is a detailed account of the sixteen American species known to date. Sadebeck's classification is followed and nine species are listed under Exoascus, five under Taphrina and two under Magnusiella. A new species is described, Exoasciis aesculi (Ell. and Ev.) Patterson on Aesculus calijornica Nutt. A footnote by A. B. Seymour tells of the identifi- cation by Sadebeck of a new species on Ostrya virginica, and this fungus is here described by Patterson under the name Taphrina virginica Sadeb. and Seym. (See 93.) Some species and hosts new to America are recorded, and the fungus on Populus tremuloidcs Michx., P. grandidenlata Michx., and P. frcmontii Wat- son, previously called (56, 59) Taphrina rhizophora, is said to be T. johansonii Sadeb. Taphrina extensa (Pk) Sacc, T. rubrobrunnea (Pk) Sacc, T. betulina Rostr., and T. lethifera (Pk) Sacc. are listed as "species inquirendae." As- comyces julgens Cke. and Hark, is excluded. 93. Sadebeck, R. Einige neue beobachtungen und kritische bemerkungen iiber die Exoasceen. Bericht. d. Deutsch. Bot. Ges. ;5:265-280. 1895. The fungus which causes yellow, swollen spots on the carpels of Populus tremula L. is said to be Exoascus johansonii Sadeb. This fungus also occurs in North America on Popidus tremuloides Michx. Whether the American form on Populus fremonlii Watson and P. grandidenlata Michx. belongs to this species or to E. rkizophorus Johans. cannot be decided without material. An account is given of the life history of E. johansonii and of the histology of the diseased host parts. Schroter's (81) classification of the genus on the basis of spore number is criticised, it being pointed out that even in species with normally eight-spored asci the spores may multiply by budding in the young asci. Two new species are described: Taphrina virginica Seym, and Sadeb., oc- curring on leaves of Ostrya virginica Willd. in North America; and Mag- nusiella jasciculata Lagerh. and Sadeb. on leaves of Ncphrodium sp. from South America. A revised classification of the family and list of species is given: 1. Exoascus Fuckel. A. Mycelium perennial in inner tissues of stems. 2. Taphrina Fries. A. Mycelium and hymenium subcuticular (Eutaphrina). a. Fertile hyphae completely used up in formation of asci. (Nine species.) b. Fertile hyphae not completely used up in forming asci. (Three species.) B. Mycelium and hymenium within epidermal cells (Taphrinopsis). (One species.) 3. Magnusiella Sadebeck. (Six species.) 134 The University Science Bulletin 94. GiESENHAGEN, K. Die entwickelungsreihen der parasitischen Exoasceen. Flora S/: 267-361. 1895. In this paper Giesenhagen, who accepts Sadebeck's genus Magnusiella but would unite all other forms into one genus, Taphrina, criticises attempts of preceding authors to subdivide this genus. Brefeld's (74) separation into Exoascus with four spores is not possible because the researches of Atkinson (86) and Sadebeck have shown that in several species the asci may be some- times four-spored and sometimes eight-spored. Tulasne's (14) classification, followed by Schroter (81) of eight-spored forms as Exoascus and many- spored forms as Taphrina, is also impossible, since, as shown by Sadebeck (73, 80), Johanson (56), and Rostrup (71), eight-spored forms may become many-spored by budding of the spores within the ascus. Sadebeck's points of distinction between Exoascus and Taphrina are also criticised. Giesenhagen does not consider the possession or lack of perennial mycelium a good basis of generic distinction, since it is not so used with higher plants. As to the further distinction used by Sadebeck, the presence or absence of sterile cells between the asci, he points out that in Taphrina betulae, for example, the mycelium being only subcuticular, such sterile cells as occur must be among the ascogenous cells, while in T. cerasi (Sadebeck's Exoascus), with inter- cellular mycelium, branches grow up from these sterile cells to form the sub- cuticular hymenium. What this amounts to is that in the one case both the sterile and fertile mycelium are beneath the cuticle, in the other the sterilo mycelium is deeper in the leaf and the fertile beneath the cuticle. Thesn differences are essentially differences in life history and with old material might be difficult to determine. Sadebeck's genus Magnusiella is like Exoascus except that branches grow up between epidermal cells to form asci without developing a subcuticular mycelium. A strict interpretation of the characters of Magnusiella will cause the re- moval of M. flava from that genus, since it has both a subepidermal and a sub- cuticular hymenium. This fungus thus becomes Taphrina flava Farl. Giesenhagen recognizes four types among the Taphrinaceae, based on the shape of the ascus. These are: 1. The Filicina type with slender asci, nar- rowed above and below; 2. The Betula type, with cylindric asci. more or less truncate at the apex; 3. The Pruni type, with clavate or cylindric asci, often rounded at the apex; 4. The Magnusiella type, with ovate or elliptic asci. Forty-six species may be readily distributed among these four types. Seven species: T. alni-incanae, T. johansonii, T. rhizophora, T. carpini, T. knichii, T. caerulescens, T. purpurascens, cannot be fitted into any of these types unless the base of the ascus is in these forms considered the equivalent of a stalk cell and the shape of the ascus determined from the ascus proper. It is then found that the first three of these species fit well into the Betula type and the others less perfectly into the same type. Forms, within the Betula type, with or without the stalk cell are similar and differ only in the presence or absence of a septum in the basal part of the ascus. Further examination of the host relationships of the three types of Taph- rina reveals the following: Type 1 (Filicina) is confined to ferns. Type 2 (Betula) to Amentiferae, Type 3 (Pruni) to Rosaceae. On basis of host- Mix: The Genus Taphrina I 135 relations a fourth type may be distinguished, the Aesculi type on Eucycleae. These four types are thought to be four evohitionary races. The Filicina race is considered the oldest because least related to the others. The Betula race is the largest and contains groups distinguishable by presence or absence of stalk cell and intercellular or subcuticular habit of mycelium. A discussion of geographic distribution of known species is given and a list with careful descriptions of the forty-nine species of Taphrina and two of Magnusiella. Taphrina aurea may or may not have a stalk cell. Examination of her- barium material (Ellis and Everhart No. 1887) shows that T. aesculi has a stalk cell contrary to the report of Patterson (92) that it has not. Patterson's report of no stalk cell in the form on fruits of Alnus incana, A. senulala, and A. rubra must mean, if true, that T. alni-incanae occurs in North America, as well as T. robinsoniana, or else that the latter species is, like T. aurea, varia- ble as to the po.ssession of a stalk cell. The species reduced to synonymy by Giesenhagen are : Exoascus campestris Sacc, E. aemiliae Passer., E. flavus Sadeb., E. varius Atk., Ascomyces querciis Cke., A. alutaceus Thm., E. accris Linh. Species excluded are: Ascomyces julgens Cke., Taphrina candicans Sacc, and T. juglandis Berk. One new species is described, Taphrina robinsoniana Giesenhag. on fruits of Alnus incana Willd. in North America. (Ellis, North American Fungi No. 796.) 95. S.\DEBECK, R. Uebcr das auftreten imd die verbreitung einiger pflan- zenkrankheiton in ostlichen alpengebiete, namentlich in Tirol. Forstl. Naturw. Zeitschr. 4:82-88. 1895. From observations of the occurrence of Taphrina oslryae Massal. on the lower leaves only of large trees decides that this fungus does not possess peren- nial mycelium. 96. Shik-U, M. (On witches' brooms of Prunus pseudocerasus.) Bot. Mag. (Tokyo) .9:161-164. 1895. In Japanese. Species diagnosis in English. Describes a new species, Taphria pseudocerasus Shirai, on Prunus pseudocerasus Lindl., and P. miquelliana Maxim. (P. stibhirtella Miq.) in Japan. 97. Stewart, F. C. Witches' brooms on cherry tree. New York Agric. Exp. Sta. Ann. Rept. ^4:532-533. 1896. Reports the occurrence of Taphrina cerasi (Fkl.) Sadeb. on Prunus avium L. on Long Island. The only previous report of this fungus in North America was that by Meehan (54). 98. Sturgis, W. C. a leaf curl of the plum. Connecticut Agric. Exp. Sta. Ann. Rcpt. /.9: 183-185. 1896. Describes a disease of a cultivated variety of Japan plum (Prunus triflora Roxb.) caused by a fungus which is identified as Exoascus mirabilis Atk. 99. Thom.^s, F. Ueber einige Exobasideen und Exoasceen. For.stl. Naturw. Zeitschr. 6:305-314. 1897. Describes a new species, Exoascus janus Thomas., on Betula verrucosa Ehrh. Suggests that this fungus, T. bacteriosperma Johans. and T. carnca Johans., may be biologic forms of a single species. 136 The University Science Bulletin 100. Derschau, D. von. Ueber Exoascus deformans. Ein beitrag zur ent- wickelungsgeschichte des parasiten. Landw. Jahrb. ^6:897-901. 1897. Reports the occurrence of Taphrina deformans on the flower of the peach. Gives notes on varietal resistance of peaches to T. deformans and of plums to T. pruni. Reports observing infection of detached peach leaves by T. de- formans, the mycelium entering a stoma. 101. ScHROTER, J. Protodiscineae. In Engler, A., and K. Prantl., Die Naturhchen Pflanzenfamilien. i: 1:156-161. 1897. Gives the following classification of the family Exoai?caceae : 1. Asci formed on the ends of branches of the intercellular mycelium. Magnus'iella. 2. Asci from a subcuticular hymenium. a. Asci eight (rarely four) spored. Exoascus. h. Asci becoming many-spored by budding of the spores. Taphria. 102. Peck, C. H. Report of the Botanist. New York State Mus. Nat. Hist. Ann. Rept. 51 :267-312. 1898. Describes a new species, Exoascus unilateralis Pk. on leaves of Prunus tiirginiana L. 103. DuGGAR, B. M. Peach leaf-curl and notes on the shot hole effect of peaches and plums. Cornell Univ. Agric. Exp. Sta. Bull. 164. 1899. An account of the peach leaf-curl disease. Spraying experiments indicate that infection is due to overwintering spores. 104. GiESENHAGEN, K., Ueber einige pilzgallen an farnen. Flora 55:100-109. 1899. Describes a new species, Taphrina fusca Giesenhag. on Aspidium pallidum Lk. Also describes T. filicina Rostr. on Aspidium spinulosum (0. F. MUll) Sw. 105. Pierce, N. B., Peach leaf-curl: its nature and treatment. U. S. Dept. Agric. Div. Veg. Phys. and Path. Bull. 20. 1900. The classical paper on the control of peach leaf-curl and on the life history of Taphrina deformans. 106. RiTZEMA Bos, J. Over krulloten en heksenbesems in de cacaoboomen Suriname en einige opmerkingen over heksenbesems in t'algemeen. Tijdschr. over Plantenziekt. 6:65-90. 1900. Describes a witches' broom of Theobroma cacao L. from Surinam. Found a few asci present, but could not, on account of the state of the material, describe the fungus. Nevertheless names it as a new species, Exoascus theobromae Ritzema Bos. A paper by the same author in Zeitschr. f. Pflanzenkr. 11 :26-30. 1901, is essentially a duplication of the above. 107. ScALiA, G. I funghi della Sicilia Orientale e principalmente della regione Etnea. Atti dell. Accad. Gioenia di Scienge naturale in Catania. (Ser. IV.) 13: XX: 1-55. 1900. Reports (inter alia) occurrence of Taphrina deformans on the almond, Prunus communis Fritsch. 108. GiESENHAGEN, K. Taphrina, Exoascus and Magnusiella. Bot. Zeit. 55:115-142. 1901. Mix: The Genus Taphrina 1 137 This paper reviews the earlier literature and presents an exhaustive bibliog- raphy. It also gives Giesenhagen's final views as to the classification of the Exoaseales. He points out that Sadebeck's Exoascus is not the Exoascus of Fuckel, and would contain only half of the species described by Fuckel, the others falling in Sadebeck's Taphrina. As an example of the way in which Sadebeck's classification separates related species he cites Taphrina caendescens, T. kruehii, T. aurea, T. jnhansonii, and T. rhizophora. According to Eadcbcck the second, fourth, and fifth of these would be called Exoascus. Giesenhagen finds Sadebeck's distinction with regard to sterile and fertile mycelium ver>' difficult to determine without an abundance of material and therefore impractical as a basis of separating genera. He accepts the genus Magnusiella, but recognizes but one other genus, whicli on the basis of priority must be called Taphrina. He again compares the ascus-form of the species (including the new T. vestergrenii) belonging to his Filicina-race, and reiterates his former view that ascus-form and host-relationships form the best basis for classification within the genus. He divides Taphrina into subgenera on the basis of his earlier classification, the Filicina-race becoming Taphrinopsis, the Betula-race Eutaphrina, the Pruni-race Euexoascus, and the Aesculi-race Sadebeckiella. He presents a synopsis, with chief distinguishing characters, of the forty-nine known species of Taphrina and five of Magnusiella, and an alphabetically arranged host index. In Magnusiella he includes, besides the two previously (93) given, M. fasciculata Lagerh. and Sadeb. on Nephrodium sp., M. lutescens Sadeb. on Aspidium thelypteris Roth., and M. potentillae Sadeb. on Potentilla silvestris Neck., P. canadensis L., and P. geoides M. B. Giesenhagen also describes a new species, Taphrina vestergrenii Giesenhag. on Amdiiim filix-mm Sw., and describes and figures spore-bearing asci for T. laurcncia. 109. Ikbno, S. Studien liber die sporen bildung bei Taphrina Johansoni Sad. Flora S8 :229-231. 1901. Describes a form occurring on fruits of Populus tremula L., var. villosa Wesm. whose asci are intermediate between those of T. rhizophora Johans. and T. johamonii Sadeb. On the basis of the host and the apparent lack of rhizoidal extensions of the base of the ascus decides to call this fungus T. johansonii. Gives an account for this species of nuclear fusion in the young ascus, division (not by typical mitosis) of the fusion nucleus and daughter nuclei, and of spore formation. 110. J.\AP, O. Ein kleiner beitrag zur pilzflora von Tirol. Deutsche bot. Monatschr. 19:75-76. 1901. Names, on the authority of Magnus, a new species, Exoascus viridis Sadeb. in litt. ad Magnus, occurring on Alniis viridis DC. No description is given. Information from Magnus is that the fungus had been collected some years previously by Sadebeck. 111. Jaczewski, a. (Exoasci of Caucasus.) Bull. Jard. Imp. Bot. St. Petersburg. i:7-13. 1901. 138 The University Science Bulletin In Russian. Lists (with description) thirty-three species occurring in the Caucasus. Describes a new species, Exoascus confiisus Jacz. on Acer camp- estris L. This name had been previously used by Atkinson (86) for a species occurring on Prunus virginiana L. 112. Rehm, H. Beitrage zur Pilzflora von Siidamerika. Hedwigia 4^:141- 170. 1901. Describes a new species, Taphrina randiae Rehm, on Randia sp. from Brazil. 113. Hennings, p. Fungi costaricenses I. Beiblatt zur Hedwigia 4^^101- 105. 1902. Describes a new species, Taphrina tonduziana Henn. on Pteris aculeata Sw. 114. JuEL, H. O. Taphridium Lagerh. and Juel, eine neue gattung der Protomj'cetaceae. Bihang t. K. Vetensk. Akad. Handl. L'7:3-29. 1902. Removes Taphrina umhelliferaruvi Rostr. and T. oreoselini Massal. from the Taphrinaceae, placing them in the new genus Taphridium, of the Pro- tomycetaceae. 115. SvENDSEN, C. J. En ny Taphrina paa Betula alpestris. Nyt. Mag. f. Naturvidensk. 40:363-368. 1902. Describes a new species, Taphrina willeana Svends. on BetuJa alpestris Fries. 116. VoLKART, A. Taphrina rhaetica nov. spec, und Mycosphaerella aronici (Fuck.). Bericht. d. d. Bot. Ges. £;:477-481. 1903. Describes a new species, Taphrina rhaetica Volk. on leaves of Crepis blat- tarioides Vill. This fungus cannot be placed in Magnusiella, since it possesses an hymenium. This hymenium being subepidermal is not like Taphrina. However, since Taphrina already includes one atypical species, T. laurencia Giesenhag., in which the hymenium is found within the epidermal cells of the host, the new fungus is placed in that genus. Some resemblance exists be- tween this fungus and Taphridium umhellijerarum (Rostr.) Lagerh. and Juel, but since it has unmistakable asci, and these and the ascogenous cells are not multinucleate from the beginning it cannot belong to that genus. 117. S.^DEBECK, R. Einige kritische bemerkungen iiber Exoasceen. I. Ber. d. Deutsch. Bot. Ges. 5/:539-546. 1903. Decides that both Taphrina rhaetica Volk. and Magnusiella potentillae (Farl.) Sadeb. really belong in Exoascus. They both possess a subepidermal hymenium. Although Sadebeck (80) had previou.sly stated that M. potentillae had no hymenium, he now agrees with Johanson (52) that it forms one be- neath the epidermis. A discussion is given of the variations in form of asci found within the same species. This is in refutation of Giesenhagen's (94) claim that races of Taphrina distinguished on the basis of ascus-form may be correlated with host-races. A new statement is made of the generic characters of Exoascus and Taphria, (which he now prefers on basis of priority to Taphrina) as follows: Exoascus. Mycelium falls apart oidium-like. Oidia form a hymenium, and either directly or after further division become ascogenous cells. Mycelium perennial. Mix: The Genus Taphrina I 139 Taphria (Taphrina). Mycelium does not fall apart oidium-like. Fertile hyphae form, after differentiation of material, on ends and on side branches of mycelial threads. 118. Ikeno, S. Die sporenbildung von Taphrina art en. Flora 5^:1-31 1903. Describes a new species, Taphrina kusanoi Ikeno., on leaves of Pasania cuspidata Oerst. Gives an account of the cytology of this and the following species: T. johansoni Sadeb., T. cerasi (Fkl.) Sadeb., T. pruni (Fkl.) Tul., T. deformans (Berk.) Fkl., and a new variety of the last named fungus, called T. deformans (Berk.) Fkl. var. armeniaca Ikeno. Describes nuclear fusion in ascogenous cells, division of the fusion nucleus (by mitosis in T. cerasi and T. pruni) to form the spore nuclei, formation of the spores with surviving cpiplasm, and (in some cases) septation of the ascus to form a basal cell. 119. Hennings, p. Fungi fluminensis A. cl. E. Ule collecti. Hedwigia 43: 78-95. 1904. Describes a new species, Exoascus uleanus P. Henn. on Ptens decurrens Presl., from Rio de Janeiro. 120. Sadbbeck, R. Einige kritische bemerkungen iiber Exoascus sebastianae nov. spec. Ber. d. Deutsch. Bot. Ges. 2^.- 119-133. 1904. Describes a new species, Exoascios sebastianae Sadeb. on Sebastiana brasili- ensis (Mull.) Arg. from Brazil. An atypical species. 121. Went, F. A. F. C. Krulloten en versteende vruchten van de Cacao in Suriname. Verhandel. d. Koninkl. Akad. v. Wetensch. te Amsterdam. Tweede Sectie. 10: 3-40. 1904. Reports a careful study of the witches' broom disease of cacao previously described by Ritzema Bos (106). Mycehum was found but no fruiting struc- tures, and it is left uncertain whether this disease may be due to a species of Taphrina. 122. Heinricher, E. E.xoascus cerasi (Fuckel) Sadeb. als giinstiger rapras- entat hexenbesen bildender pilze fiir pflanzenbiologische gruppen. Naturw. Zeitschr. Forst. und Landw. 3: 344-348. 1905. Reports successful grafting of witches' brooms {Taphrina cerasi (Fkl.) Tul.) into healthy cheny trees, also the failure of Peyritsch to produce infection with spores. 123. Ja.\p, O. Beitriige zur Pilzflora von Mecklenberg. Ann. Mycol. 3: 391-401. 1905. Lists several species. A new host. Alnits glutinosa X incana Krause {A. hybrida A. Br.) is reported for Taphrina alni-incanae (Kiihn) Magn. 124. KusANO, S. New species of Exoascaceae. Bot. Mag. (Tokyo). 19: 1-5. 1905. Describes three new Japanese species: Taphrina trundcola Kus. on Prunus incisa Thunb., T. piri Kus. on Pyrus miyabe Sargent, and T. japonica Kus. on Alniis japonica S and Z. 125. KusANO, S. Einige neue Taphrina-Arten aus Japan. Ann. Mycol. 3.- 30-31. 1905. Repeats his descriptions of the three new species mentioned above. 140 The University Science Bulletin 126. Nbger, F. Exoascaceae. In Hennings, P., G. Lindau, P. Lindner and F. Neger. Kryptogamenflora der Mark Brandenburg. 7: 35-73. 1905. A comprehensive account of the family, recognizing Giesenhagen's classifi- cation, and including a taxonomic treatment of the twenty-two species found in the province of Brandenburg. 127. YosHiNO, K. A list of the fungi found in the province of Higo. Bot. Mag. (Tokyo) 19: 199-222. 1905. (In Japanese.) Lists six species of Taphrina found in Japan. Of these two new species are named: T. hiratsukae Nishida on Onoclea sp., and T. mume Nishida on Prunus arrneniaca L. var. ansu Maxim. The descriptions of the above species are to be found in the later paper by Nishida (140). 128. BuBAK, F. Zweiter beitrag zur pilzflora von Montenegro. Bull, del Herbier Boissier Ser. 2, 6: 393-408, 473-488. 1906. Describes a new species, Taphrina morijormis Bubak on Aspidium rigidum Sw. 129. Faber, F. von. Bericht liber die pflanzenpathologische expedition nach Kamerun. Tropenpflanzer. 11: 755-775. 1907. Names a new species, Exoasciis hxissei v. Fab., on Theobroma cacao L. 130. KusANO, S. A new species of Taphrina on Acer. Bot. Mag. (Tokyo) SI: 65-67. 1907. Describes a new species, Exoascus nikkoensis Kus. on Acer purpurascens Fr. and Sav. The ascus has a basal cell, otherwise this species seems close to A. lethifcr Pk. and T. polyspora (Sorok.) Johans. 131. Fabeir, F. C. von. Untersuchungen iiber krankheiten des Kakaos. I. Uber den hexenbesen der kakaobaume in Kamerun. Arb. aus der Kaiserl. Biolog. Anstalt fur Land-und Forstwirtsch. 6: 385-395. 1908. Describes a new species Taphrina theobromac Fab. (see 129) causing a witches' broom of Theobroma cacao L. in Kamerun. Gives an account of the morphology and anatomy of the witches' broom. Believes it distinct from the disease of cacao described by Ritzema Bos (106) as occurring in Surinam, and declares the binomial Exoascus theobromac Ritzema Bos to be a nomen nudum. 132. CoNNOLD, E. T. Plant galls of Great Britain. London, Adler. 1909, p. 117. Figures and describes a witches' broom on Corylus avcllana L. as due to ExoasciLs Avellanae Con. No evidence is presented that this deformation is due to a fungus and the binomial was later declared by Saccardo (vol. 24, 1301) to be a nomen nudum. 133. JuEL, O. Om Taphrina-Arter pa Betula. Svensk. Bot Tidskr. 3: 183- 191. 1909. Li.sts, with descriptions and some discussion of relationships, nine species of Taphrina occurring on Betula in Sweden. 134. Palm, B. Taphrina andina n. sp. Sven.sk Bot. Tidskr. 3.- 192-195. 1909. Describes a new species, Taphrina andina Palm, on leaves of Prunus sabci- jolia HBK. Mix: The Genus Taphrina I 141 135. CoKER, W. C. A new host and station for Exoascus filicinus (Rostr.) Sacc. Mycologia 2: 247. 1910. Reports occurrence of Taphrina filicina Rostr. on Dryopteris acrostichoides Kiintze {Polystichum acrostichoides (Michx.) Schott). 136. Herzfeld, S. Ueber eine neue Taphrina auf Polystichum Lonchitis. Oesterr. Bot. Zeitschr. 60: 249-254. 1910. Describes a new species, Taphrina wettsteiniana Herzf. on Polystichum lonchitis (L.) Roth. 137. M.AJRB, R. Contribution a I' etude de la flore mycologique des Alpes- Maritimes. Champignons recoltes a la session de Saint-Martin-Vesubie (1910). Bull. Soc. Botanique de France. 4 ser. 10: CLXVI-CLXXVI. 1910. Describes a new species, Taphrina viridis (Sadcb.) Maire on Alnus viridis DC. This fungus was previously named Exoascus viridis Sadeb. by Jaap (110) but no description was given. It was also distributed in Vestergren, Micro- mycetes Selectae Exsiccati No. 720. 1903, as Taphrina alnastri Lagerh. 138. Th.'Vxter, R. Not<>s on Chilean fungi. I. Bot. Gaz. 50: 435. 1910. Describes a new species, Taphrina entomospora Thaxt., on Nothojagus antarctica Oerst. from Punta Arenas, Chile. 139. Butler, E. J. The leaf spot of Turmeric (Taphrina maculans sp. nov.) Ann. Mycol. .9:36-39. 1911. Describes a new species, Taphrina maculans Butl. on Curcuma tonga L., and Zingiber casumunar Roxb. The fungus grows in the radial walls of the epidermal and hypodermal cells, but in the intercellular spaces of the meso- phyll. It is unique among the Taphrinaceae in possessing haustoria. 140. NiSHiD.\, T. A contribution to the monograph of the parasitic Ex- oascaceae of Japan. Miyabe-Festschrift. Tokyo 1911. pp. 157-212. In Jap- anese. English Summary, pp. 205-212. Gives descriptions, with notes on occurrence of twentj'-five species of Taphrina found in Japan. New species are as follows: Taphrina struthiopteri- dis Nish. on Strulhiopteris germanica Willd. (Matteuccia struthiopteridis Tod.), T. hiratsukai Nish. on Onoclea sensibilis L., T. osmundae Nish. on Osmunda regalis L.vav. japonicaW iM., T.coryli Is ish. on Corylus heterophylla Fisch. and C. rostrata Ait. var. sieboldi Maxim., T. betulicola Nish. on Betula crmanni Cham. var. nipponica Maxim., T. alni-japonicae Nish. on Alnus japonica S. and Z., T. mume Nish. on Prunus mume S. and Z., and P. armeniaca L. var. anzu Maxim. Species recorded on new hosts are as follows: T. caerulescens on Quercus dentata Thunb., Q. crispada Bl., Q. grosserata Bl., Q. glandulijera BL, Q. scrrata Thunb.; Q. glauca Thunb.; T. bullata on Pyrus sinensis Lindl.; T. cerasi on Prunus pseudocerasus Lindl. var. spontanea Maxim., and var. sieboldii Maxim., P. miqweliana Maxim.; T. pruni on Prunus tomentosa Thunb., and P. incisa Thunb.; T. farloid on Prunus ssiori Fr. Schm.; T. truncicola Kus. on Prunus maximomczii Rupr. This list of species includes MagnusieUa umbellijerarum (Rostr.) Sadeb., which is actually Taphridium umbellijerarum (Rostr.) Lagerheim and JueL The number of Taphrinas described is therefore twenty-four. 141. Sydow, p., and E. J. Buti.er. Fungi Indiae Orientalis. III. Ann. Mycol. 9: 372-421. 1911. 142 The University Science Bulletin Report occurrence of Taphrina deformans (Berk.) Fkl. on peach, E, pruni Fkl. on fruits of Prunus padus L., T. aurea Fr. on leaves of Populus ciliata Wall., T. maculans Butl., on leaves of Curcuma longa L., C. amadae, C. angustifolia Roxb., Zingiber casumunar Roxb., Zingiber zerumbet Roscoe. Describes a new species, T. rhomboidalis Syd. and Butl. on leaves of Pteris quadriaurita Retz. Apparently another species occurs on this host in Ceylon and Assam. 142. JuEL, H. 0. Beitriige zur kenntnis der gattungen Taphrina und Ex- obasidium. Svensk. Bot. Tidskr. 6: 353-372. 1912. Notes on the various species of Taphrina found on Betula in Lapland. A description is given of a new species, Taphrina lapponica Juel. on Betula odorata Bechst. Also a new variety, T. nana Johans. var. hyperborea Juel, on the same host. T. bacteriosperma Johans. was found on a new host, Betula nana L. 143. Laubert, R. Einige pfianzenpathologische beobachtungen. Zeitschr. f. Pflanzenkr. 23: 449-454. 1912. Describes a witches' broom of Prunus acida (Dum.) K. Koch, very similar to the one previously described by Sadebeck (73) as caused by Taphrina minor on Prujius chamaecerasus Jacq. (Prunus fruticosa Pall.). It is sug- gested that Sadebeck's host may have been P. acida. Morphologically the fungus on P. acida agrees well with either T. minor or T. cerasi and it seems possible that these two species are not well distinguished. 144. KuscHKE, G. Mycoflorae caucasicae novitates. Moniteur du Jard. Bot. Tiflis. 37; 23-27. 1913. Describes a new species, Exoascus cerasi-microcarpi Kuschke, on Cerasus microcarpa Boiss. (Prunus microcarpa C. A. Mey.) The article is in Russian, but a Latin diagnosis is given. 145. Clintox, G. p. Notes on plant diseases of Connecticut. Report of the Station Botanist, for 1913. Connecticut Agric. Exp. Sta. Rept. 1914 Part I: 1-30. 1914. Records a case of Taphrina dejormans attacking the fruit of the peach. 146. SvDow, H., and P. Sydow'. Diagnosen neuer philippinischer pilze. Ann. Mycol. 12: 545-576. 1914. Describes a new species, Taphrina linearis Sydow on Globba m,arantina I>. 147. ScHELLENBERG, H. C. Zur kenntnis der winterruhe in den zweigen einiger hexenbesen. Ber. d. Deutsch. Bot. Ges. 33: 118-126. 1915. Notes a shortening of the winter rest period in buds of witches' brooms caused by Taphrina cerasi and T. turgida. Believes little or no carbon assimila- tion occurs in leaves of witches' brooms. 148. Heinricheir, E. Zur frago nach der assimilatorischen leitungsfiihigkeit der hexenbesen des kirschbaumes. Ber. d. Deutsch. Bot. Ges. 33: 245-253. 1915. An account of experiments which show higher osmotic pressure and more carbon assimilation in tissues of the cherry witches' broom (Taphrina cerasi) than in healthy tissues. Mix: The Genus Taphrina I 143 149. ScHMiTz, H. Some observations on witches' brooms of cherries. Plant World. 19: 239-242. 1916. Notes higher osmotic pressure, rich food content and presence of anthocyans in tissues of witches' brooms caused by Exoascus cerasi. Similar observations were made on a witches' broom found on Prunus emarginata Walp. 150. Del-uiness, J. New or noteworthy North American fungi. Mycologia 9: 345-364. 1917. Describes a new species, Exoascus aceris Deam. and Barth. on Acer grandi- dentatum Nutt. 151. J.\AP, O. Weitere beitrage zur pilzflora der Schweiz. Ann. Mycol. 15: 97-124. 1917. Lists twelve species of Taphrina as occurring in Switzerland. Raises T. polyspora Sorok. var. -pseudo-platani Massal. to specific rank as T. pseudo- platani (Massal.) Jaap. 152. Palm, B. Svenska Taphrinaarter. Arkiv. for Botanik. 15: 1-41. 1917. Describes the following new species: Taphrina lata Palm, T. lagerheimi Palm, and T. splendens Palm on Betxda odorata Bechst., T. media Palm on Alnus glutinosa Medic. Raises T. betulae (Fkl.) Johans. var. auciumnalis Sadeb. to specific rank as T. auciumnalis (Sadeb.) Palm. Renames Exoascus confusus Jaczewski as Taphrina jaczewski Palm, since the specific name used by Jaczewski (111) had already been employed by Atkinson (86) for another fungus. Lists, with keys and descriptions, thirty-five species as occurring in Sweden. 153. Sydow, H., and P. Sydow. Beitrag zur kenntnis der pilzflora der Philippinen Inseln. Ann. Mycol. 15: 165-268. 1917. Report the occurrence in the Philippines of Taphrina maculans Butl. on Zingiber zerumbet Roscoe. 154. Swingle, D. B., and H. E. Morris. Plum pocket and leaf gall on Americana plums. Montana Agric. Exp. Sta. Bull. 123. 1918. An account of plum pockets caused by Taphrina communis on culti\'ated varieties of Prunus americana. Perennial mycelium is believed to be of rare occurrence. The ascospores are uninucleate but preliminary to conidium for- mation the nucleus divides to two or even several nuclei. 155. SiEMASZKO, W. Fungi caucasici novi vel minus cogniti. Bull. Musee du Caucase. 12: 20-28. 1919. Describes a new species, Taphrina struthiopteridis Siem. on Matteuccia struthiopteris Todaro. 156. Sydow^ H., and P. Sydow. Novae fungorum species. XVI. Ann. Mycol. IS: 154-160. 1920. Describe a new s]iecies, Taphrina mexicana Syd. on leaves of Prunus microphylla Hemsl. in Mexico. 157. JrEL, H. 0. Cylologische pilzstudien. II Zur kenntnis einiger Hemi- asceen. Nov. Act. Reg. Soc. Sci. Upsal. Ser 4, 5: 3-41. 1921 This paper contains an account of the cytology of several species of Taphrina. 144 The University Science Bulletin In T. pruni and T. rostrnpiana cells of the mycelium contain two closely paired nuclei. The young ascus has one nucleus. The first division of this nucleus may be a reduction di\ision. Two divisions follow. Spores form around the eight nuclei in a layer of cytoplasm next the ascus-walls, the in- terior of the ascus being empty. A small amount of epiplasm remains after spore formation. In T. cerasi the empty basal cell is formed by vacuolization of the lower part of the chlamydospore (which is what Juel considers the ascogenous cell) and formation of a septum. In T. bullata the young cells of the mycelium each contain several pairs of nuclei, old cells contain each one pair. In T. potentillae each ascus-bearing branch of the mycelium contains one pair of nuclei. In the widened end of this branch the nuclei fuse, the widened end elongates to an ascus and the fusion nucleus divides by three successive divisions. In places a local subcuticular layer of ascus-forming hyphae occurs. In T. alnitorqua Tul. chlamydospores are at first binucleate, later the nuclei fuse. Before division of the fusion nucleus all the protoplasm enters the ascus, leaving the chlamydospore empty. This is then cut off by a septum to form a basal cell. The first division of the fusion nucleus was observed. In T. sadebecki nuclear divisions in the ascus were not seen. Formation of the ascus and the basal cell occurs as in T. epiphylla. In T. epiphylla 3'oung vegetative cells are binucleate. The chlamydospore has a thick membrane ; it germinates by the rupture of this wall and the emergence of a thin-walled protoplast, which becomes the ascus. The empty chlamydospore is cut off from the ascus by a wall, thus becoming the basal cell. Nuclear division in the ascus was not observed. T. belulina differs from the preceding in that in ascus formation the whole wall of the chlamydospore becomes thinner and extended. An empty basal cell is cut off. Three successive nuclear divisions occur in a thin layer of protoplasm next to the ascus wall. In T. carnea the ascus forms as in the preceding. The chlamydospore empties as usual, but no wall forms and there is no separate basal cell. The first division of the ascus nucleus was observed. Ascospores bud within the ascus almost at once. In T. aurea young cells of the mj'celium have each one pair of nuclei. An apparent telophase of conjugate division was observed. Ascogenous cells are at first binucleate, then the nuclei fuse. Division of the fusion nucleus may be by amitosis. Not all of the eight nuclei may form spores. Spores bud almost at once. Basal cell forms as in other species, but it is very small and may be lacking. In all of the above species where nuclear division was observed it was mitotic, the figures being intranuclear. The genus Taphrina is considered by Juel to be closely allied to Taphridium and Protomyces of the Protomycetaceae. The ascogenous cell (as noted above) is considered a chlamydospore. The basal cell of the ascus is a feature of some Taphrinas, the forms lacking it are considered the more primitive. 158. Wedermann, E. Taphrina reichei n. sp. ein neuer mexikanischer hexenbesen. Notizbl. d. Bot. Gart. und Mus. in Berlin-Dahlem. S.- 221-222. 1922. Describes a new species, Taphrina reichei Werd. on Prunus capollin Koehne. Mix: The Genus Taphrina I 145 159. Becker, J. Ein beitrag zur ziichtung der pfirische. Gartenwelt 27: 274-275, 285-286. 1923. Discusses varietal susceptibility of the peach to curl. 160. Briton-Jones, H. R. Pear leaf blister (Taphrina bullata, Tul.). Univ. Bristol, Agric. and Hort. Res. Sta. Ann. Dept. 1923: 89-90. 1923. Reports occurrence of pear leaf blister in many localities in England in 1923. Attack by Taphrina predisposes pear leaves to attack by Venturia pinna Aderh., the cause of pear scab. Varieties of pears are listed as to susceptibil- ity to T. bullata. 161. CuNNiNGH.AM, G. H. Leaf-curl, bladder plum and cherrj^-curl, their appearance, cause and control. New Zealand Jour. Agric. 26: 85-87. 1923. Reports occurrence in New Zealand of Taphrina dejormans, T. pruni, and T. minor, with discussion of their life histories and some notes as to varietal resistance. ■ 162. Kleb.'VHN, H. Infoktionsversuche mit Taphrina tosquinetii. Ber. d. Deutsch. Bot. Ges. 41: 108-113. 1923. Reports cultivation in artificial media of Taphrina tosquinetii (Westend.) Magn., T. epiphylla Sadeb., T. sadebeckii Johans., T. aurea Fr. and Taphrina sp. from Betiila piibcficcns Ehrh. Inoculation experiments with T. tosquinetii were successful, with the. other species they failed. It is concluded that T. tosquinetii sometimes overwinters as mycelium within the host tissues. 163. Martin, E. M. Cytological studies of Taphrina coryli Nishida on Corylus americana. Trans. Wisconsin Acad. Sci., Arts, and Letters. 21: 345- 355. 1924. An account of the cytology of this species. The vegetative cells and the 5'oung ascogenous cells are binucleate. Nuclear fusion occurs in ascogenous cells. The fusion nucleus divides by reduction division, a cleavage furrow and then a septum cuts off a basal cell from the young ascus. The basal cell and the ascus each receive one of the daughter nuclei, but that in the basal cell soon degenerates. The ascus-nucleus divides by three successive divisions to form the nuclei for the spores. The bud-conidia are uninucleate. 164. Mix, A. J. Biological and cultural studies of Exoascus deformans. Phytopath. U: 217-233. 1924. This paper reports studies of the behavior of Taphrina deformans in arti- ficial culture, successful inoculations, and experiments and observations on the life historj' of this fungus. 165. Campbell, W. G. Note on an Exoascus disease on Pnaius amygdalus var. amara. Trans, and Proc. Bot. Soc. Edinburgh. 2^?: 5; 186-191. 1925. This is an account of the morphology and life history of a fungus attack- ing Prunus amygdnlus Stokes {P. communis Fritsch), var. amara Hort. and var. dulcis Hort., also of the pathological changes induced by it in the former host. Evidence is presented that mycelium over-winters in the phloem. The fungus differs from Taphrina deformans (Berk) Fkl. in the absence of a basal cell of the ascus, and in size of asci and of spores. It is considered to be a variety of that fungus. 10—6037 146 The University Science Bulletin 166. Efttmiu, p. Sur I'Exoascus deformans (Berk.) Fuck. Compt. Rend. Acad. Sci. Paris 181 : 1085-1087. 1925. This paper gives an account of the cytology of T. deformans and of the pathological changes induced in the host tissues. The findings are reported in detail in a subsequent paper (174) by the same author. 167. Martin, E. M. Cultural and morphological studies of some species of Taphrina. Phytopath. 15: 2: 67-76. 1925. Reports brief cultural studies of Taphrina johansonii T. communis, T. mha- bilis, T. deformans, and T. caerulescens. Claims successful inoculation on bur oak with T. caerulescens. 168. Martin, G. H. Diseases of forest and shade trees, ornamental aud miscellaneous plants in the United States in 1923. Plant Disease Reporter. Supplement 37. 1925. On pages 371-373 is an account, supplied by Dr. A. E. Jenkins, of the various species of Taphrina known to occur in America on Acer spp. These are: a species very similar to T. accricola Massal. on Acer saccharum Marsh, and A. nigrum Michx., an undescribed species on A. dasycarpum Ehrh. {A. saccha- rinum L.), another on A. rubrum L., and the previously recorded T. lethifer (Pk.) Sacc, and E. aceris Dearn. and Barth. Other notes on the occurrence of Taphrinas in America are found in this paper. 169. Ml\, a. J. Biological and cultural studies of Exoascus mirabilis. Phytopath. 15: 4: 214-232. 1925. Cultural studies and observations on the life history of Taphrina m,irabilis (Atk.) Giesenhag. on Prunus angustifolia Marsh. 170. Jaczewski, a. a. (Pocket key for the determination of fungi. Part first. Exoascales.) In Russian. A. A. Jaczewski Mycol. Lab. State Inst, of PJxper. Agric. Leningrad. 1926. This is the latest monographic treatment of known species of Taphrina. The genera of Sadebeck are recognized : Exoascus, with forty-seven species, and Taphrina, with fifty-three. Taphrina entomospora Thaxt. is treated as the type of a thiid genus as proposed by Saccardo {22: 765) becoming En- tomospora antarclica (Thaxt.) Sacc. The following species, excluded by previous authors, are recognized : Taphrina cis&i, T. fulgens, T. extensa, and T. flai^o-aurea. A variety of T. deformans on Prunus arnygdalus is raised to specific rank a."-- Exonftcus amygdali Jacz. [Apparently this is not the same fungus as that described by Campbell (165).] Other new species are Exoascus pruni-acidae Jacz. on Prunus acida (Dum.) Koch., E. sorbi Jacz. on Sorbics terminalis Crantz. The following new varieties are described: Exoascus pi-uni Fkl. var. padi Jacz. on Prurnis padus L., var. divaricata Jacz. on Prunus divaricata L. {Prunus inssardi Carr and Carr), var. ussuricnsis on Prunus uss-uriensis. New hosts are given as follows: for T. alni-incanae, Alnus serrulata Willd.; for T. betulina, Betula carpatica Waldst, and B. tortuosa Led.; for T. carpini, Carpinus orienialis Mill.; for T. crataegi, Crataegus sanguinea Pall.; for T. filicina, Athyrium filix-fem.ina Roth., and Nephrodium phegopteris; for T. insititiae, Prunus divaricata Led.; for T. minor, P. microcarpa C. A. Mey.; for T. rhizophora, Populu^ tremula x alba {P. bachofenii) ; for T . sadebeckii. Mix: The Genus Taphrina I 147 Alnus tinctoria Sarg. {A. hirsuta Turcz.), A. incana Willd., and A. incana var. sibirica Spach. {A. hirsuta Turcz var. sihinca Schneid.) ; for T. carnea, Betula tortuosa Led., and B. gmelini Bge. {B. fruticosa Pall.) ; for T. ulmi, Ulmus suberosa Moench {U. joliacea Gilib. var. suberosa Schneid.). 171. OvERHOLTS, L. 0. Mycological notes for 1924. Mycologia IS: 31-38. 1926. Reports a species of Taphrina (.somewhat different from T. lutescens) on Onoclea sensibilis L. 172. Wedgworth, H. H. Leaf blister of oak. Mississippi State Plant Board. Quart. Bull 6: 10-12. 1926. Reports that Taphrina caerulescens attacks nearly all native species of oaks, being most severe on Quercus nigra L., and Q. rubra L., while Q. phellos L. shows marked resistance. 173. BoYCE, J. S. Observations on forest pathology in Great Britain and Denmark. Phytopath. 17: 1-18. 1927. Reports finding, in Scotland, Taphrina aurea Fr. on Populus generosa Henry., and P. laurijolia Ledeb., as well as on P. nigra L. 174. Eftimiu, p. Contribution a 1' etude cytologique des Exoascees. Le Botaniste 18: 1-154. 1927. Species studied arc Taphrina aurea Fr., T. alni-incanae (Kiihn) Magn., T. bullata (Berk.) Tul., T. deformans (Berk.) Fkl., T. betulae (Fkl.) Tul., T. pruni (Fkl.) Tul., T. crataegi Sadeb., T. insititiae Sadeb. Cells of young mj'celium are plurinucleate or binucleate. Ascogenous cells are binucleate. Fusion of nuclei occurs just before formation of asci. Asci arise by an outgrowth of a papilla from an ascogenous cell. This outgrowth breaks the cuticle and its upper two thirds becomes the ascus, while the lower part forms the empty basal cell. (In T. alni-incanae there is no basal cell.) The ascus nucleus forms by three successive divisions the eight nuclei for the ascospores. The first of these divisions is a reduction division, there being four chromosomes in diploid and two in haploid nuclei. The ascospores germinate by budding within the ascus or after liberation. The bud receives a daughter nucleus from the mother cell, this nucleus by division initiates the binucleate condition in the mj^celium. In the host the parasites induce hyperthrophy and hyperplasia of cells, ac- companied by obliteration of intercellular spaces and thickening of cell walls. Chloroplasts become transformed into amyloplasts. At a certain stage starch disappears from the cells, being probably utilized by the fungus. The spher- ical cytosomes of healthy cells become vesicular or rod-shaped and lose chro- matin. T. aurea utilizes oil drops in the host cells. .175. R.\thburn-Gr-\vatt, A. A witches' broom of introduced Japanese cherry trees. Phytopath. 17: 19-24, 1927. This is an account of a witches" broom disease on Prunus lannesiona Wilson, and Prunus yedoensis Mat. growing in Washington, D. C. The fungus on P. yedoensis was identified as Exoascus cerasi (Fkl.) Sadeb. 176. WiEBEN, M. Die infektion, die myzeluberwinterung, und die kopula- tion bei Exoasceen. Forsch. auf. d. Geb. d. Pflanzenkr. und d. Immun. im Pflanzenr. 3: 139-176. 1927. 148 The University Science Bulletin Reports artificial culture and successful inoculation experiments with Taph- rina tosquinetii (Westend.) Magnus, T. epiphylla Sadeb., and T. deformans (Fkl.) Tul. Inoculations failed with T. aurea Fr., T. bullata (Berk, and Br.) Tul., T. crataegi Sadeb., and T. klebahni Wieben. Mycelium of T. epiphylla, T. tosquinetii, T. betulina Rostr., T. sad'ebeckii Johans., T. bullata, and T. aurea is perennial and overwinters in the buds. Copulation occurs between sexuallj'' different conidia (the eight ascospores are of two sexes) in T. epiphylla and T. klebahni. The resulting cell puts forth a long spiral tube bearing a pair of nuclei at its tip. A new species is described: Taphrina klebahni Wieben on Alnv^ incana Willd. 177. Zeller, S. M. Contribution to our knowledge of Oregon Fungi. II. Mycological notes for 1925. Mycologia 19: 130-143. 1927. Describes a new species, Exoascus pruni-subcordalae Zell. on Prunus sub- cordata Benth. This is considered nearly related to E. longpipes Atk. and E. communis Sadeb. 178. Heinricher, E. Uber einen hexenbesen auf Cydonia japonica. Ber. d. Deutsch. Bot. Ges. 46: 198-204. 1928. Describes a witches' broom on Cydonia japonica Hort. {Chaenomeles la- qenaria Koidz.), which, it is believed, may be due to a Taphrina. 179. Jankowska, K. Zewnetrzniaki polski. Mem Inst. National Polonais d" Econ. Rur a' Pulawy. 9: 182-215. 1928. Describes artificial cultivation of Taphrina sadebeckii Johans. Describes twenty-eight species occurring in Poland, four of them being reported for the first time. ISO. Laubert, R. Taphrinaccae (Exoascaceae). In Sorauer, P. Handbuch der Pflanzenkrankheiten 2: 457-499. 5th edition. Parey. Berlin. 1928. A comprehensive account of the genus, including a discussion of the im- portant diseases caused by various species. A complete list of species is given, with description of the most important species. 181. Sansone, F. Una speciale deformazione dei fruitti di mandorlo dovuta ad attaco dell' Exoascus dejormans. (Berk.), Fuck. Boll. R. Staz. di Pat Veg. N. S. 8: 291-299. 1928. Description of the attack of T. deformans on the fruit of the almond. 182. Seymour, A. B. Host index of fungi of North America. Cambridge, Mass. 1929. Lists the species of Taphrina known to occur in North America. 183. FiTZPATRiCK, R. E. The life history and parasitism of Taphrina de- jormans. Scient. Agric. H: 305-326. 1934. Reports studies on the life history of the peach leaf-curl fungus, including time of infection, method of penetration and nuclear behavior. 184. Koch, L. W. Studies on the overwintering of certain fungi parasitic and saprophytic on fruit trees. Canadian Jour, of Res. 11: 190-206. 1934. Reports isolation of Taphrina dejormans from the surfaces of dormant peach buds and successful inoculations with the cultures so obtained. Mix: The Genus Taphrina I 149 185. PoMERLEAU, R. Notes sur le Ta-phrina ulmi. Naturaliste Canadien. 1934: 305-308. Occurrence of Taphrina ulmi (Fkl.) Tul. in Quebec, presumably on Ulmus americana L. 186. Mix, A. J. The life history of Taphrina deformans. Phytopath. 25: 41-66. 1935. Reports results of several years' investigation of the peach leaf-curl disease. 187. FiTZPATRiCK, R. E. Further studies on the parasitism of Taphrina de- formans. Scient. Agric. 15: 341-344. 1935. Studies of the effect of temperature on infection of peach leaves by Taph- rina deformans and on the subsequent development of the disease. The leaf is shown to become more resistant as it matures. 188. Jenkins, A. E. An undescribed species of Taphrina on chinquapin. Mycologia SS: 31-34. 1936. Describes a new species, Taphrina castanopsidis (Ellis and Ev.) Jenkins, on Castanopsis chrysophylla DC. 189. Bataille, M. F. Monographie des Exoascacees D'Europe. Ann de la Soc. Linneenne de Lyon. 79: 121-130. This is a descriptive list, with keys, of species of Exoascus and Taphrina oc- curring in Europe. Several invalid species are included and nearly one third of the known European species omitted. THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 10. Tlie Genus Taphrina. II: A List of Valid Species A. J. MIX Department of Botany, University of Kansas Absteact: a list, with descriptions, is given of 104 species of the genus Taphrina, which seem likely to prove valid. The recognition of a single genus Taphrina to include forms described under Ascomyces, Exoascus, Taphrina, and Magnusiella, has necessitated the making of a few new combinations of scien- tific names. Preoccupation of the species name has necessitated the renaming of one species, Taphrina struthiopteridis Siemaszko, this becoming Taphrina siemaszkoi (Siemaszko) n. comb. IN compiling the following descriptive list of species of Taphrina, the object has been to render their identification as easy as possible. Descriptions have been reduced to essentials: a brief characterization of the effect on the host, a statement (if the point is on record) as to whether the mycelium grows intercellularly or merely beneath the cuticle, a description of the asci as to position, shape, size, presence or absence of a stalk cell, number, shape and size of spores. Dimensions of asci and spores are given in microns, but the symbol for microns is omitted from the text. In connection with each species references are given to the paper containing the original description and to other papers in which descriptions may be found. These references are cited by author's name and number, the number referring to the complete citation to be found in the first paper of this series (Mix, The Genus Taphrina I: An Annotated Bibliography. University of Kansas Science Bulle- tin £4:9: 113-149, 1936). References are also made to Sac.cardo's Sylloge Fiingorum by volume, page, and number. The original idea of Giesenhagen that Exoascus, Magnusiella, and Taphrina should be united to a single genus Taphrina has been accepted. This has necessitated the revision of a few scientific (151) 152 The University Science Bulletin names. Giesenhagen's objections to Sadebeck's division of the genus into Taphrina, Exoascus, and Magnusiella seem to be well taken, and although Giesenhagen later proposed a genus Magnusiella, this was made with special reference to species which were later found to belong to the genus Taphridium of the Protomycetaceae. Jaczew- ski places Taphrina entomospora Thaxt. in the genus Entomospora of Saccardo. Saccardo did not describe a genus Entomospora, but merely proposed it, and while this aberrant species might well be placed in a separate genus, it may be quite as well to await the dis- covery of other similar species before so doing. It has been necessary to assign a new specific name to the form described by Siemaszko on Matteucia struthiopteiis Tod. since the name Taphrina struthiopteridis had been previously used by Nishida. Although occurring on the same host, these two fungi are, from their published descriptions, quite different. For the present, at least, they must be considered as separate species. Only those species have been excluded or reduced to synonymy which have been so treated by other authors. Some species are probably synonymous. It seems highly probable, for example, that Taphrina rubrobrunnea (Pk.) Sacc. is actually T. caerulescens (Desm. and Mont.) Tul. Its reduction, however, must await the completion of studies of T. caerulescens now in progress in this lab- oratory. Perhaps certain other species should be regarded with sus- picion, but their true status can only be determined after careful morphological study combined with cross-inoculation experiments. Some studies of this nature are being undertaken, but their accom- plishment will take time. Meanwhile, it is hoped that the list which follows will eliminate some confusion in the deteraiination of species. 1. Taphrina cornu-cervi Giesenhag. Giesenhagen (77, 108),*Sadebeck: (80, 93), Jaczewski (170), Saccardo ll:i37. No. 2721. Host: Aspidium aristatum Sw. {Polystichum arislatum Presl.) Tumors on fronds. Asci: Clavate, rounded, 24 x 5-6, narrowing below to 2-4. Stalk cell 4-6 X 2-4. Spores: Lacking in type specimen. Distribution: Nepal and Ceylon. 2. Taphrina Jusca Giesenhag. Syn. : Exoascus fuscus (Giesenhag.) Sacc. and Syd. Giesenhagen (104, 108), .Jaczewski (170), Saccardo i6:803, No. 2921. Host: Aspidium pallidum Lk. Tumors on fronds. Mycelium : Subcuticular. Mix: The Genus Taphrina II 153 Asci: On upper surface, elongate-clavate, rounded, 20-24 x 5-7; stalk cell long, cylindric. Spores: Eight in ascus, ellipsoid, 3-4 x 2. Distribution: Sicily, Albania. 3. Taphrina moriforniis Bubak. Bubak (128), Jaczewski (170), Saccardo 16:766, No. 398. Host: Aspidmm rigiduni Sw. Outgrowths on fronds. Asci: Subcylindric, 30-45 x 4-6, narrowed at base. Spores: Not known. Distribution : Montenegro. 4. Taphrina filicina Rostr. Syn. : Ascomyces filicinus Rostr. ; Exoascus filicinus (Rostr.) Sacc. Johanson (56), Sadebeck (80, 93), Giesenhagen (94, 104, 108), Saccardo 8:819, No. 3354. Host: Aspidmm spinulosum (O. F. Mull) Sw. According to Jaczew-ski (170), also on Aihyrium filix-jemina Roth., and Nephrodium phegopteris {Phegoptcris phegopteris (L.) Underw.?). Spots on fronds. Mycelium : Subcuticular. Asci: Amphigenous, clavate, rounded, 29-38 x 5-9, attenuate at base to width 3.5-4.5; no stalk coll. Spores: Often eight in ascus, oblong or ovoid, 4-5 x 2. Distribution: Sweden, and (according to Jaczewski) Moscow district and Caucasus, Russia. 5. Taphrina lutescens Rostr. Syn. : Magnusiella lutescens (Rostr.) Sadeb. Rostrup (71), Sadebeck (80, 93), Giesenhagen (94, 108), Palm (152), Jaczewski (170), Saccardo 10:68, No. 4729. Host: Dryopteri^ thelypteris (L.) A. Gray. Yellow spots on fronds. Mycelium : Intercellular. Asci: Slender, 60-75 x 8-9; no stalk cell. Spores: Unknown. Conidia numerous, 4-5 x 0.5-1. Distribution : Denmark, Germany, Russia. 6. Taphrina athyrii Siemaszko Jaczewski (170). Host: Athyrium filix-femina Roth. Asci : Hypophyllous, clavate, tnmcate or rounded, 20-32 x 5-7. Spores: Usually 6 to 8 in ascus, ellipsoid, 4-5.5 x 3.3-4.5. Distribution : Caucasus. 7. Taphrina vestergrenii Giesenhag. Syn. : Exoascus vestergrenii (Giesenhag.) Sacc. and Syd. Giesenhagen (108), Jaczewski (170), Saccardo i6 :1152, No. 4789; J8 :196, No. 3849. Host: Dryoptens filix-mas Schott. Mycelium : Subcuticular. 154 The University Science Bulletin Asci: Amphigenous, cylindric, rounded or truncate, 25 x 6; stalk celt truncate. Spores: Eight in ascus, suboblong, 7 x 2.5-3. Distribution: Baltic Russia, Sweden, Switzerland, Alsace. 8. Taphrina struthiopteridis Nishida Nishida (140). Host : Matteucia struthioptcris Todaro. Yellow spots on fronds. Asci: Hypophyllous, clavate, rounded or truncate, 16-28 x 4-6; stalk cell 4-7 X 3-4. Spores: Usually eight in ascus, spherical, diameter 2. Distribution: Japan. 9. Taphrina siemaszkoi (Siemaszko) n. comb. Syn. : Taphrina struthiopteridis Siemaszko. Siemaszko (155), Jaczewski (170), Saccardo 2^:1304. Host: Matteucia struthioptens Tod. Yellow spots on fronds. Asci : Hypophyllous, clavate. rounded, 28-40 x 6-7 (170) . Spores: Six to eight in ascus, subglobose, 4-5.5 x 3.3-4.5. Distribution : Caucasus. 10. Taphrina fasciculata (Lagerh. and Sadeb.) Giesenhag. Syn. : Magnusiella fasciculata Lagerh. and Sadeb. Sadebeck (93), Giesenhagen (94, 108). Host: Nephrodiuni sp. Spots on fi'onds. Asci: Resembling those of T. potentillae, 50-70 x 9-12; stalk cell present, narrower than ascus, dimensions not given. Spores: Ovate, 5-8 x 4. Distribution: Quito, Ecuador, South America. 11. Taphrina hiratsukae Nishida Nishida (140), Saccardo 22:766, No. 5878. Host: Onoclea sensibilis L., Nephrodium thelypteris Desv. (Dryoptcris thelypteris (L.) A. Gray?). Asci: Clavate or fusiform, rounded or truncate, 18-30 x 4-6; stalk cell 7-8 x 3-4. Spores: Usually 8 in ascus, elliptic, 4-5 x 2-3. Distribution : Japan. 12. Taphrina osmundae Nishida Nishida (140). Host: Osmunda regalis L. var. japonica Willd. Spots on fronds. Asci : Hypophyllous, rarely epiphyllous, oblong or oblong clavate, 32-44 x 1 7- 25 ; no stalk cell. Spores: Numerous, minute. Distribution : Japan. Mix: The Genus Taphrina II 155 13. Taphrina wettsteiniana Herzf. Herzfeld (136), Palm 152), Jaczewski (170), Saccardo 2-}:1303. Host: Polystichum lonchitis (L.) Roth. Mycelium : Intercellular. Asci: Slender-cylindric, rounded or blunt, 50-70 x 5-7; stalk cell present or absent, half length of ascus. Spores: Eight in ascus, spindle shaped, sometimes slightly constricted in middle, 5 x 12. Distribution : North Tyrol. 14. Taphrina tonduziana P. Henn. Hennings (113), Jaczewski (170), Saccardo JS:197, No. 3855. Host: Pteris aculeata Sw. Spots on fronds. Asci: Hypophyllous, clavate, rounded or truncate, 16-24 x G-S. Spores: Eight in ascus, fusiform or clavate, 6-7 x 1.5-2.5. Distribution: Central America. 15. Taphrina uleana (P. Henn.) Giesenhag. Syn. : Exoascus vleantis P. Henn. Hennings (119), Jaczewski (170), Saccardo iS:196, No. 3850. Host: Pteris decurrens Presl. Spots on fronds. Asci : Hypophyllous, cylindric-clavate, truncate, 30-35 x 12-15. Spores: Eight in ascus, subglobose or ovoid, 6-10x6-8. Distribution: Brazil. 16. Taphrina laurencia Giesenhag. Ciesenhagen (77, 94, 108), Sadebeck (80, 93), Jaczewski (170), Saccardo i/:437, No. 2722. Host: Pteris qiiodriaurita Retz. Bushy outgrowths on fronds. Mycelium : Intracellular. Asci: Formed vvithm epidermal cells, clavate, 24 x 7; stalk cell cylindric 19-6-7. Spores: Not present in type-specimen. Later reported by Giesenhagen (108) as right in a.scus and in size near those of T. vcstcrgrcnii (7x2.5-3). Distribution : Ceylon. 17. Taphrina rhomboidalis Syd. and Butl. Sydow and Butler (141), Jaczewski (170), Saccardo 2.} :1303, No. 7559. Host: Pteris quadriaurita Retz. Spots on leaves. Asci: Amphigenous, clavate, broader above, rounded, 30-32 x 6.5-8. Spores: Ellipsoid. 5-7.5 x 2-3. Distribution: India. 18. Taphrina acericola Massal. Syn.: Exoascus acericola (Massal.) Sacc. Massalongo (84), Giesenhagen (94, 108), Jaczewski (170), Saccardo J/:436, No. 2710. Host: Acer campestre L. Spots on leaves. 156 The University Science Bulletin Asci: Hypophyllous, rarely amphigenous, cylindric-clavate 16-20 x 6-8; stalk cell variable, 10-12 x 4-6. Spores: Usually eight in ascus, subglobose, diameter 2.5-3.5. Distribution : Italy. 19. Taphrina jaczewski Palm. Syn. : Excascus covfvsus Jacz. Jaczewski (111), Palm (152), Jaczewski (170), Saccardo 18:196, No. 3848, 2^:1301, No. 7551. Host: Acer campesire L. Witches' brooms. Mycelium: Intercellular. Asci: Hypophyllous, oblong-cylindric, 16-20 x 8-10; stalk cell roundish or irregular in form. Spores : Usually 6 in ascus, ovoid, 5-6 x 3-4. Distribution : Caucasus. 20. Taphrina aceris (Dearn. and Barth.) n. comb. Syn. : Exoascus aceris Dearn. and Barth. Dearness (150), Saccardo 24:1300, No. 7547. Host: Acer grandidentatum Nutt. Spots on leaves. Asci: Hypophyllous, short-clavate or cylindric, 30 x 9. Spores: Subglobose or irregular, 4.5-6 x 3-4.5. Distribution: Utah, U. S. A. 21. Taphrina acerina Eliass. Syn.: Exoascus acerinus (Eliass.) Sacc. Eliasson (91), Palm (152), Jaczewski (170), Saccardo 1^:823, No. 3075. Host: Acer platanoidcs L. Deforming whole leaves. Mycelium : Subcuticular. Asci: Amphigenous, more often hypophyllous, broad cylindric or clavate, round or truncate, often attenuate at base, 15-23 x 9-12 ; stalk cell rounded at base, 7-9 x 12-15. Spores: Eight in ascus, globose, diameter 4-5. Distribution: Sweden, Russia. 22. Taphrina pseudoplatani (Massal.) Jaap. Syn.: Taphrina polyspora (Sor.) Joh. var. pseudoplatani Massal.; Taphrina acericola Massal. (var.) pseudoplatani Massal. Massalongo (76, 84), Jaap (151), Palm (152), Jaczewski (170), Saccardo JJ :436, No. 2710. Host: Acer pseudoplatanus L. Spots on leaves. Asci : Hypophyllous, subcylindric or clavate, rounded, 16-24 x 10-12 (76) or 18-26 X 7-10 (84) ; no stalk cell (76), stalk cell 7 x 15 (84). Spores: Numerous, globose or ellipsoid, diameter 2-2.5. Distribution : Italy. Mix: The Genus Taphrina II 157 23. Taphrina nikkoensis Kusano Syn. : Exoascus nikkoensis (Kus.) Sacc. and Trott. Kusano (130), Jaczewski (170), Saccaido 23:763, No. 5867. Host: Acer purpurascens Fr. and Sav. Spots on leaves. Mycelium : Subcuticular. Asci: Mostly hypophyllous, cylindric or somewhat clavate, rounded or subtruncate, 40-50 x 10-13; stalk cell 10-15 x 10-13. Spores : Uusally eight in ascus, globose, diameter 5, ellipsoid, 4-5 x 7. Distribution : Japan. 24. Taphrina lethifera (Peck) Sacc. Syn. : Ascomyces letifer Peck. Peck (58), Jaczewski (170), Saccardo 10:67, No. 4723. Host: Acer spicatum Lam. Deforming leaves. Asci: Hypophyllous, cylindric, obtuse or subtruncate, 0.0016 to 0.0020 in. X 0.0006 to 0.0008 in. (Peck), 40-50 x 15-20 (Saccardo). Spores: Subglobose or narrow elliptic, 0.00016-0.00020 in. x 0.00008 x 0.00012 in. (Peck), 4-5 x 2-3 (Saccardo). Distribution: Elizabethtown, New York, U. S. A. 25. Taphrina polyspora (Sorok.) Johans. Syn. : Ascomyces polysporus Sorok. ; Exoascus aceris Linh. Sorokine (24), Fisch (49), Johanson (52), Sadebeck (80, 93), Giesenhagen (94, 108), Palm (152), Jaczewski (170), Saccardo 8:813, No. 3327. Host: Acer totaricum L. Spots on leaves. Asci: Usually epiphyllous, cylindric, rounded, 33-47 x 12-17; no stalk cell. Spores: Eight in ascus, globose, diameter 4-5, rarely seen. Asci filled with conidia, diameter 3-4. Distribution: Western Europe. 26. Taphrina aesculi (Patterson) Giesenhag. Syn.: Ascomyces deformans var. aesculi Ell. and Ev. Exoascus aesculi (Ell. and Ev.) Patterson. Harkness (68), Patterson (92), Giesenhagen (94, 108), Jaczewski (170), Saccardo 22:763, No. 5867. Ellis and Everhart, North American Fungi, No. 1887. Host: Aesculus calif omica Nutt. Young shoots and leaves. Asci: Amphigenous, cylindric, rounded, 16-18 x 7-9; no stalk cell. Giesen- hagen (94) gives dimensions of ascus as 25-30 x 9, and of stalk cell 5-8 x 15. Spores : Elliptic, 6x3. Distribution: California, U. S. A. 27. Taphrina media Palm. Syn. : Exoascus medius (Palm) Jacz. Palm (152), Jaczewski (170), Saccardo 2i :1301, No. 7552. Host: Alnus glutinosa Medic. Witches' brooms. Mycelium : Subcuticular. 158 The University Science Bulletin Asci: Broad cylindric, truncate at apex, 25-90 x 10-12. Spores: Eight in ascus, globose, diameter 3-5. Distribution: Sweden. 28. Taphrina sadebeckii Johans. Syn. : Exoascus alni de By. in part; Exoascus flavus Sadeb. ; Exoascus epiphyllus Sadeb. var. maculans; Ascomyces tosquinetii Westend. in part. Westendorp (12), Tulasne (14), Sadebeck (47, 61, 73, 80, 93), Johanson (52), Giesenhagen (108), Palm (152), Jaczewski (170), Saccardo 8:816, No. 3338. Host: Alnus glutinosa Medic; A. tinctoria Sarg. {A. hirsuta Turez) ; A. hybrida A. Br.; A. incana Willd. var. sibirica Spach. (A. hirsuta Turcz. var. sibirica Schneid.). Yellow spots on leaves. Asci: Hypophj'llous, cylindric, truncate, 41-55 x 14-18, with yellow con- tents; stalk cell 18-22 x 18-22. Spores: Globose, diameter 6.5. Distribution: Europe. 29. Taphrina tosquinetii (Westend.) Magnus Syn. : Ascomyces tosquinetii Westend. ; Taphrina alnitorqua Tul. ; Exoascus alnitorquus Sadeb.; Exoascus tosquinetii Magnus; Exoascus alni de By. in part. Westendorp (12), Tulasne (14), Fuekel (15), Sadebeck (47, 73, 80, 93), Magnus (70), Giesenhagen (94, 108), Palm (152), Jaczewski (170), Saccardo S:817. Host: Alnus glutinosa, A. hybrida A. Br. Deforming leaves and twigs. Asci: Cylindric, truncate, 31-37 x 6-7; stalk cell 11-20 x 6-7, inserted be- tween epidermal cells. Spores: Usually 8, globose, diameter 3-5. Distribution : Europe. 30. Taphrina alni-incanae (Kiihn) Magn. Syn.: Taphrina alni-incanae (Kiihn) Sadeb.; Ascomyces alni Berk, and Br.; Exoascus alni- torquus (Tul.) forma alni-incanae Kiihn. in Litt. ; Exoascus alnitorquus (Tul.) Sadeb. in part: Exoascus alni de By. in part; Exoascus alni de By. var. strobilinus Thm. ; Ascomyces tosquinetii strobilina Rostr. ; Exoascus amentorum Sadeb. Berkeley and Broome (25), Rostrup (71), Sadebeck (47, 73, 80, 93), Magnus (70), Patter- son (92), Giesenhagen (94, 108), Palm (152), Jaczewski (170), Saccardo iO :69, No. 4733. Host: Abius incana Willd., A. glutinosa Medic, A. hybrida A. Br., A. rubra Bong., A. rugosa (Du Roi) Spreng. Deforming female catkins. Asci: Cylindric, truncate, 40-45 x 10 (Sadebeck), 40-50 x 10 (Giesenhagen), 35-41 x 9-10 (Patterson), inserted for about one-third of length between epi- dermal cells; no stalk cell. Spores: Usually 8 in ascus, globose, diameter 5. Distribution: Europe, North America. 31. Taphrina epiphylla Sadeb. Syn. : Exoascus epiphyllus Sadeb. ; Taphrina sadebeckii Johans. var. borealis Johans. ; Taphrini borealis Jolians. Sadebeck (47, 73, 80, 93), Johanson (52), Rostrup (71), Giesenhagen (94, 108), Palm (152), Jaczewski (170), Saccardo 8:816; No. 3339, 3340. Host: Almis incana Willd. Witches' brooms. Mycelium : Intercellular. Mix: The Genus Taphrina II 159 Asci: On both surfaces of leaf, broad cj-lindric, truncate, 33-40 x 15-20, in- serted in stalk cell. Latter truncate below, 8-9 x 20-33, or pointed, 15-20 x 20-30. (Johanson's variety borealis had asci 37-54x15-17, stalk cells 12-25 x 17-24.) Spores: Eight in ascus, globose, diameter 5-6. Distribution : Europe. 32. Taphrina klebahni Wieben. Wieben, (176). Host: Alnus incana Willd. Yellowish spots on leaves. Mycelium : Subcuticular. Asci : Cylindric, broader and truncate at apex, acute at base and somewhat inserted, 35-45 x 15-20. Spores : Eight in ascus, globose, diameter 5. Distribution: Hamburg, Germany. 33. Taphrina robinsoniana Giesenhag. Syn. : Exoascus rohinsonianus (Giesenhag.) Sacc. and Trott.; Taphnna alnitorqua Robins. not Tul. Giesenhagen (94, 108), Saccardo 22:765, No. 5873. Host: Alnus incana Willd. Deforming carpels. Mycelium: Subcuticular. Asci: Cylindric, subround at apex, truncate at base; 29-37 x 6-10. Stalk cell 15-17 X 6-10. Spores: Eight in ascus, globose, diameter 3.5-6. Distribution: North America. 34. Taphrina japonica Kusano. Syn. : T. alni-japonicae Nishida. Kusano (125), Nishida (140), .laczewski (1"0), Saccardo i8 :197, No. 3854. Host: Alnus japonica Sieb. and Tsug. Witches' brooms. Asci: Hypophyllous, cylindric, round or obtuse, sometimes slightly broader at base. 60-90 x 16-25. No stalk cell. Spores: Numerous, diameter 2-3. Distribution: Japan. 35. Taphrina viridis (Sadeb.) Maire. Syn.: Exoascus viridis Sadeb.: Taphrina altiastri Lagerh. Jaap (110), Maire (137), Saccardo 24:1300, No. 7548. Host: Alnits viridis DC. Spots on leaves. Asci: Ellipsoid oblong, rounded, 16-22 x 10; stalk cell inserted between epidermal cells. Spores : Subglobose or ellipsoid. 5-6 x 3-4. Distribution: Germany, Italy. 36. Taphrina willeana Svends. Svendsen (115), Juel (133), Palm (152), Jaczewski (170). Host: Betula alpestris Ft. Spots on leaves. Mycelium: Subcuticular. 160 The University Science Bulletin Asci : Amphigenous, broad cylindric, truncate or rarely emarginate, 40-50 x 15-20; stalk cell broad, 20-24 x 24-30. Spores: Numerous, 6-8 x 4-5. Distribution: Norway, Sweden. 37. Taphrina betulicola Nishida Nishida (140). Host: Betula ermanni Cham. var. nipponica Maxim. Witches' brooms. Asci: Cylindrical, rounded or truncate, 36-42 x 14-22; stalk cell broad, 8-20 x 20-26. Spores: Numerous, minute. Distribution : Japan. 38. Taphrina alpina Johans. Syn. : Exoascus alpinus (Johans.) Sadeb. Johanson (56), Sadebeck (80, 93), Giesenhagen (94, 108), Juel (133), Palm (152), Jac- zewski (170), Saccardo 8:818:3349. Host: Betula nana L., B. verrucosa Ehrh., B. ermanni Cham. Witches' brooms. Mycelium : Subcuticular. Asci: Hypophyllous (more often epiphyllous according to Jaczewski), broad cylindric, rounded or truncate, often constricted at base, 20-27 x 9-14; stalk cell often somewhat inserted between epidermal cells, 8-14 (rarely 17- 18) x 12-20 (mostly 15-17). Spores: Usually eight in ascus, globose, diameter 3-5. Distribution: Sweden, Russia, Kamchatka. 39. Taphrina bacteriosperma Johans. Syn. : Exoascus bacteriospermus (Johans.) Sadeb. Johanson (56), Sadebeck (80, 93), Patterson (92), Giesenhagen (94, 108), Juel (133). Palm (152), Jaczewski (170), Saccardo 8:814, No. ?330. Host: Betula nana L., B. odorata Bechst.. B. alpcstris Fr., B. glandulosa Michx. Deformation of leaves and twigs. Mycelium : Subcuticular. Asci : More often epiphyllous, broad cylindric, rounded or rounded-truncate, often dilated, 47-80 x 14-20, at base often 28-30 broad; no stalk cell. Spores: Globose, diameter 3.6-4.5 (Jaczewski). Conidia numerous, rod- shaped, 6.8-7 X 1. Distribution: Sweden, Greenland, Russia, North America. 40. Taphrina nana Johans. Syn.: Exoascus nanus (Johans.) Sacc. ; Exoascus nanus (Johans). Sadeb. Johanson (52), Sadebeck (80, 93), Giesenhagen (94, 108), Juel (133), Palm (152), Jac- zewski (170), Saccardo 8:818, No. 3348. Host: Betula nana L., B. udorata Bechst., B. alpestris Fr. Witches' brooms. Mycelium : Intercellular. Mix: The Genus Taphrina II 161 Asci: Mostly epiphyllous, cylindric, truncate or rounded, 18-24x7-9, rarely 24-30 X 7-9; stalk cell 7-10, rarely 12-15 x 8-17, truncate at base. Spores: Often eight in ascus, globose, diameter 3-5. Distribution: Swedish Lapland, Kamchatka. 41. Taphrina carnea Johans. Johanson (52), Sadebeck (80, 93), Giesenhagen (94, 108), Juel (133), Patterson (92), Palm (152), Jaczewski (170), Saccardo 8:813, No. 3329. Host: Betula gmelini Bge. {B. fruticosa Pall) B. humilis Schr., B. nana L., B. intermedia Thomas, B. ordorata Bechst., B. pubescens Ehrh., B. tortuosa Led. Spots on leaves. Asci: Mostly epiphyllous, broad cylindric, rounded or truncate, 44-80 x 14-30; no stalk cell. Spores: Numerous (conidia). Distribution: Sweden, Russia. 42. Taphrina lagerheimi Palm. Palm (152), Jaczewski (170), Saccardo 24:1302, No. 7553. Host: Betula odorata Bechst. Deformation of leaves and twigs. Mycelium: Subcuticular. Asci: Hypophyllous, cylindric, rounded truncate, 53-60 x 13.5-17; stalk ceil 19.5-23.5 x 19.5-26. Spores: Globose, diameter 4.5-6, condidia globose, minute. Distribution: Sweden. 43. Taphrina lata Palm. Palm (152), Jaczewski (170), Saccardo 24:1302, No. 7555. Host: Betula odorata Bechst. Deforming leaves and twigs. Mycelium : Subcuticular. Asci: Hypophyllous, cylindric, rounded, 40-45 x 18-22; stalk cell 16-20 x 25-33. Spores: Ascospores not seen, conidia numerous, 3-5 long. Distribution: Sweden. 44. Taphrina splendens Palm. Palm (152), Jaczewski (170), Saccardo 24:1302, No. 7556. Host: Betula odorata Bechst. Witches' brooms. Mycelium : Subcuticular. Asci : H3rpophyllous, subcylindric, truncate-rounded, slightly constricted to- ward base, 59-71 x 20-23; stalk cell truncate, 20-25 x 20-24. Spores: Ascospores not seen, conidia numerous, minute, globose. Distribution: Lapland, Sweden. 11— €037 162 The University Science Bulletin 45. Taphrina lapponica Juel. Syn. : Exoascus lappcmicus (Juel) Jacz. Juel (142), Palm (152), Jaczewski (170), Saccardo 24:1302, No. "554. Host: Betula odorata Bechst., B. alpestris Fr. Deforming leaves and twigs. Mycelium: Intercellular. Asci: Hypophyllous, 40x16; stalk cell present (as in T. alpina according to Jaczewski). Spores: Numerous, globose, diameter 3.5, or ellipsoid, 4 x 2.5 (Jaczewski). Distribution: Sweden. 46. Taphrina betulina Rostr. Syn.: Exoascus betuUnus (Rostr.) Sadeb. ; Exoascus turgidus Sadeb. in part. Rostrup (43, 71), Johanson (52), Sadebeck (80, 93), Giesenhagen (94, 108), Juel (133), Palm (152), Jaczewski (170), Saccardo 8:818. Host: Betula odorata Bechst., B. carpatica Waldst., B. pubescens Ehrh., B. tortuosa Led. Witches' brooms. Mycelium : Subcuticular. Asci: Hypophyllous, cylindric, rounded, 45-55x15-20 (sometimes only 30-40 long) ; stalk cell variable, 20-25 x 15-20, or broadened below, 10 x 20-25, not inserted. Spores: Globose, diameter 5. Distribution: Europe. 47. Taphrina betulae (Fkl.) Johans. Syn. : Exoascus betulae Fk. ; Ascomyces betulae Magn. Fuckel (17), Sadebeck (47, 80, 93), Johanson (52), Rostrup (71), Giesenhagen (94, 108), Juel (133), Palm (152), Jaczewski (170), Rabenhorst, Fungi Europaei 2734, Saccardo 8:818, No. 3346. Host: Betula odorata Bechst., B. pubescens Ehrh., B. verrucosa Ehrh. Spots on leaves. Asci: On both surfaces, broad cylindric, truncate and wider at base. 25-40 X 8-12. Basal cell broad, 8-12 x 12-25. Spores: Eight in ascus, subglobose, 3-5. Distribution: Europe. 48. Taphrina auctumnalis Palm. Sj'n. : Taphrina betulae (Fkl.) Johans. var. auctumnalis Sadeb. Sadebeck (47), Juel (133), Palm (152), Jaczewski (170). Host: Betula odorata Bechst., B. pubescens Ehrh., B. verrucosa Ehrh. Reddish spots on leaves. Asci: Amphigenous, 15-27x6-9; stalk cell often broad, but not beyond 2-5 (Jacz.). Spore.s: Globose, diameter 3-5. Distribution : Europe. Mix: The Genus Taphrina II 163 49. Taphrina janus (Thomas) Giesenhag. Syn. : Exoascus janus Thomas. Thomas (99), Giesenhagen (108), Juel (133), Palm (152), Jaczewski (170), Saccardo J4:824, No. 3076. Host : Betula odorata Bechst., B. verrucosa Ehrh. Spots on leaves. Asci : Amphigenous, 52-63 x 9-15. Spores: Globose, diameter 3-5. Distribution: Switzerland, Sweden. 50. Taphrina fiava Farl. Syn.: Magnusiella fiava (Farl.) Sadeb. Farlow (44), Sadebeck (93), Giesenhagen (94, 108), Jaczewski (170), Saccardo 8:813, No. 3328. Host: Betula populifolia Marsh., B. alba L. var. papyrijcra (Marsh.) Spach. Yellow spots on leaves. Mycelium : Intercellular. Asci: Cylindric, obtuse or truncate at both extremities, 38-57x20-23, no stalk cell. Spores: Oblong, tenuous, 3-7 x 1.5-2. Distribution: North America. 51. Taphrina turgida (Sadeb.) Giesenhag. Syn. : Exoascus turgidus Sadeb. Sadebeck (47, 80, 93), Giesenhagen (94, 108), Juel (133), Palm (152), Jaczewski (170), Saccardo 8:818, No. 3347. Host: Betula verrucosa Ehrh. Witches' brooms. Mycelium : Subcuticular. Asci: Hypophyllous, cylindric, tnmcate, 46-50x15; stalk cell tapering, in- serted between epidermal cells, 17-30 x 15. Spores: Globose, diameter 3-4. Distribution : Europe. 52. Taphrina carpini (Rostr.) Johans. Syn. : Exoascus carpini Rostr. Rostrup (35, 71), Sadebeck (47, 73, 80, 93), Johanson (52, 56), Giesenhagen (94, 108), Palm (152), Jaczewski (170), Saccardo 8:814, No. 3331. Host: Carpimis betvlus L., C. orientalis Mill. Witches' brooms. Mj'celium : Subcuticular. Asci : Hypophyllous, cylindric, rounded, broad and truncate at base, 25 x 8-12. 20-24 broad at base; no stalk cell. Spores : Globose, diameter 4. Distribution: Northern and central Europe. ]64 The University Science Bulletin 53. Taphrina australis (Atk.) Giesenhag. Syn. : Ezoascus australis Atk. Atkinson (86), Giesenhagen (94, 108), Jaczewski (170), Saccardo llAZl, No. 2719. Host: Carpinus caroliniana Walt. Spots on leaves. Mycelium : Subcuticular. Asci: On upper surface, cj'lindric truncate, base narrowed to short foot, 30-60x7-10; no stalk cell. Spores: Globose, diameter 4-6. Distribution: Alabama, U. S. A. 54. Taphrina castanopsidis (Ell. anci Ev.) Jenkins. Jenkins (188). Host: Castanopsis chrysophylla A. DC. Spots on leaves. Asci: Hj'pophyllous, cylindric, rounded, 80-165x13-17, inserted basal por- tion variable in form and size. Spores: Eight in ascus, diameter up to 10, conidia 3-5x1.5-2.5. Distribution: California, U. S. A. 55. Taphrina celtis Sadeb. Syn.: Exoascus celtidis (Sadeb.) Sacc. ; Exoascus aemiliae Passer. Sadebeck (73, 80, 93), Passerini (67), Giesenhagen (94, 108), Jaczewski (170), Saccardo 10:69, No. 4731, 4732. Host: Celtis australis L. Spots on leaves. Asci: Hypophylloiis, cylindric, blunt-rounded, 25-28x8-10; stalk cell flat- tened, 8-10 X 25-30. Spores: Globose, diameter 3-5.5. (Passerini's Exoascus aemiliae, declared by Sadebeck to be identical with T. celtis, had asci 12-15x6-7; stalk cell not found, spores globose, diameter 2.5-3.) Distribution: Central Europe. 56. Taphrina coryli Nishida Nishida (140), Martin (167), Jaczewski (170). Host: Coryliis americana Walt., C heterophylla Fisch., C. rostrata Ait. var. sieboldiana Maxim. Yellow spots on leaves, and leaf-curl. Asci : Usually hypophyllous, cylindric or cylindric clavate, rounded, 20-24 x 8-12; stalk cell roundish flattened, 8-12x8-16. Spores: Usually eight in ascu.s, globose, diameter 4-6. Distribution: Japan, North America. 57. Taphrina crataegi Sadeb. Syn.: Exoascus crataegi (Sadeb.) Sacc; Exoascus bullatus Sadeb. in part. Sadebeck (47, 73, 80, 93), Giesenhagen (94, 108), Palm (152), Jaczewski (170), Saccardo i0:70. No. 4735. Host: Crataegus oxyacantha L., C. m,onogyna Jacq., C. sanguinea Pall. Deforming leaves and rarely flowers. Witches' brooms recorded by Sadebeck and Palm. Mycelium : Subcuticular. Mix: The Genus Taphrina II 165 Asci: Hypophylloiis, cylindric, truncate, 25-35x8; stalk cell 6-8x6-8. Spores: Eight in ascus, globose, diameter 4.5. Distribution: Europe, comparatively rare. 58. Taphrina maculans Butl. Butler (139), Sydow and Butler (141), Jaczewski (170), Saccardo 24:1303, No. 7560. Host: Curcuma longa L., C. angustifolia Roxb., C. amadae, Zingiber cosunmnar Roxb., Z. zerumbet Rose. Yellow-brown spots on leaves. Mycelium: Growing in walls of cells, possessing haustoria. Asci : Clavate, rounded or truncate, often attenuate at base, 20-30 x 6 ; pro- vided with a stalk cell or even a row of two or three. Spores: Five to ten, mostly eight, in ascus, ovoid or sub-elongate, 4-6.5 X 2-2.5. Distribution: India. 59. Taphrina linearis Sydow. Sydow and Sydow (146), Saccardo 24:1303, No. 75G1. Host: Globba marantina L. Spots on leaves. Asci: Clavate, rounded or truncate, 25x7-9 (immature); with one or a few stalk cells. Spores: Not known. Distribution : Philippine Islands. 60. Taphrina entomospora Thaxt. Syn. : Exoascus entomosporus (Thaxt.) Sacc. and Trott. ; Entomospora antarctica (Sacc.) Jacz. Thaxter (138), Jaczewski (170), Saccardo 22:765, No. 5874. Host : Nothojagus antarctica Oerst. Spots on leaves. ■ Asci: Subcylindric, rounded or subtruncate, 55-60 x 13-15; stalk cell broader than ascus. Spores: Eight in ascus, appendiculate, 9-10x3-4. Appendages of two ordera, terminal 8-12 x 3-5, subterminal 15-25 x 0.8. Distribution: Punta Arenas, Patagonia, South America. 61. Taphrina ostryae Massal. Syn. : Exoascus ostryae Massal. Massalongo (60), Sadebeck (73, 80, 93), Giesenhagen (94, lOS), Jaczewski (170), Saccardo 8:818, No. 3350. Host: Ostrya carpinifolia Scop. Spots on leaves. Asci: Hypophyllous, oblong, obtuse, 20-24x12-14; stalk cell variable in form and size, inserted. Spores: Eight in ascus, globose, diameter 5-7. Distribution : Italy, Caucasus. 62. Taphrina virginica Seym, and Sadeb. ), Sadebeck (93), Giesenhagen (94, 108), Ji Host: Ostrya virginica Willd. Spots on leaves. Patterson (92), Sadebeck (93), Giesenhagen (94, 108), Jaczewski (170), Saccardo 18:824, No. 3078. 166 The University Science Bulletin Asci: Hypophyllous, cylindric, rounded or truncate with truncate base, 25 X 10; no stalk cell. Spores: Numerous, minute. Distribution : North America. 63. Taphrina kusanoi Ikeno Ikeno (118), Jaczewski (170), Saccardo 22:765, No. 5875. Host: Pasania cuspidata Oerst. Swellings on leaves. Asci: Hypophyllous, cylindric, rounded, 102-117x13-19; no stalk cell. Spores: Numerous (conidia), small, ellipsoid. Distribution : Japan. 64. Taphrina rhizophora Johans. Syn. : Taphrina aurea auct. in part. ; Exoascus aureus auct. in part ; Exoascus rhizophorus (Johans.) Sadeb. Johanson (56), Sadebeck (73, 80, 93), Giesenhagen (94, 108), Palm (152), Jaczewski (170), Saccardo 8:812, No. 3326. Host: Populus alba L., and P. tremula L., P. bachofeni (P. tremula x alba). Mycelium : Subcuticular. Asci: With yellow contents, elongate, clavate, rounded, attenuate at base to root-like appendage, often forked, inserted between epidermal cells, 120- 160x20-22, inserted part narrowing to diameter 6; no stalk cell. Spores: Globose, diameter 4. Distribution: Europe, North America. 65. Taphrina aurea Fr. Byn. : Taphrina popuUna Fr. ; Ascomycts aureus (Pers.) Magn. ; Exoacus aureus (Fr.) Sadeb. ; Exoascus populi Thm. ; Exoascus flavo-aureus Cocc. Fries (1, 2, 3, 4, 7), Sadebeck (47, 73, 80, 93), Johanson (52, 56), Cocconi (88), Patterson (92), Giesenhagen (94, 108), Palm (152), Jaczewski (170), Saccardo 8:812, No. 3325. Host: Populus nigra L. and Populus spp. Yellow spots on leaves. Asci: With yellow contents. Form variable, cylindric or clavate, rounded or truncate, narrowed toward base, size as reported by various authors 47-112 X 15-27; stalk cell present or absent, variable in size and form, 4-27x8-17. Spores: Globose, diameter 4, forming numerous conidia. Distribution: Europe, North America. 66. Taphrina johansonii Sadeb. Syn. : Taphrina rhizophora Johans. in part. ; Exoascus johansoni Sadeb. ; Exoascus aurew Sadeb. in part. Sadebeck (73, 80, 93), Rostrup (71), Patterson (92), Giesenhagen (94, 108), Jaczewski (170), Saccardo 10:68, No. 4725. Host: Populus tremula L., P. tremuloides Michx., P. grandidenlata Michx., P. jremontii Wats. Deforming carpels. Mycelium : Subcuticular. Mix: The Genus Taphrina II 167 Asci: With yellow contents, clavate, rounded, deeply inserted, 46-105 x 14-20 (exserted part) or 8 (inserted part) according to Patterson; 92-105x16-25 according to Sadebeck; no stalk cell. Spores: Globose, diameter 4. Distribution: Europe, North America. 67. Taphrina potentillae (Farl.) Johans. Syn. : Exoasai-^ potentillae (Farl.) Sacc. ; Ascomyces potentillae Farl.; Exoascus deformans (Berk.) Fkl. var. potentillae Farl.; Taphrina tormentillae Rostr. ; Magnusiella potentillae (Farl.) Sadeb. Farlow (44), Rostrup (50), Johanson (52), Sadebeck (73, 80, 93), Patterson (92), Giesen- hagen (94, 108), Palm (152), Jaczewski (170), Saccardo 8:819, No. 3352. Host: Polcntilla silvestris Neck., P. geoides L., P. canadensis L., P. silves- tris X procumbens. Deforming leaves and stems. Mycelium: Intercellular, hymenium subepidermal. Asci: Clavate, rounded or truncate, prolonged to pedicel below, 40-55x8-10. Spores: Ovate-oblong, 4 x 5.8. Distribution: P]urope, North America. 68. Taphrina pruni-acidae (Jacz.) Syn. : Exoasciis pruni-acidae Jacz. ; Taphrina sp. Laubert. Laubert (143), Jaczewski (170). Host: Prunus acida (Dum.) Koch. Deforming fruits, flowers, and flower buds. • Asci: 30-36x9-10; stalk cell pedicellate, sometimes forked, 6-16x3-10. Spores: 6-9x5. Distribution : Germany. 69. Taphrina communis (Sadeb.) Giesenhag. Syn. : Exoascus communis Sadeb. Sadebeck (80, 93), Patterson (92), Giesenhagen (94, 108), Jaczewski (170), Saccardo iJ:43C, No. 2709. Host: Prunus amencana Marsh., P. pumila Wang., P. maritinw Wang., and P. subcordnta Benth. Deforming fruits. .\sci: Clavate, rounded or blunt, 30-40x8; stalk cell 15-20x3-5. Dimen- sions of asci given by Giesenhagen as 25-45x6-10; stalk cell 15-25x3-6. Spores : Eight in ascus, globose or broad ellipsoid, 5 x 3-4. Distribution: North America. 70. Taphrina decipiens (Atk.) Giesenhag. Syn. : Exoascus decipiens Atk. Atkinson (86), Giesenhagen (94, 108), Sadebeck (93), Jaczewski (170), Saccardo ii :436, No. 2714. Host: Pninus americana Marsh. Leaf-curl. Asci: Hypophyllous, 20-40x7-10; stalk cell 6-13x7-12. Spores : Oval or broad-elliptic, diameter 3-4. Di-stribution : New York, U. S. A. 168 The University Science Bulletin 70a. Var. superficialis Atk. Deforming your fruits of P. americana. Asci 25-30x8-11; stalk cell often short. 71. Taphrina longipes (Atk.) Giesenhag. Syn. : Exoascus longipes Atk. Atkinson (86), Giesenhagen (94, 108), Sadebeck (93), Jaczewski (170), Saccardo ii :436, No. 2713. Host: Prunus americana Marsh. Deforming fruits. Mycelium : Intercellular. Asci: 30-40x7-10; stalk cell 25-35x3-5, inserted between epidermal cells. Spores: Not described by Atkinson. Given by Jaczewski as globose or ellipsoid, diameter 3-4. Distribution: New York, U. S. A. 72. Taphrina mirabilis (Atk.) Giesenhag. Syn. : Exoascus mirabilis Atk. Atkinson (86), Giesenhagen (94, 108), Sadebeck (93), Jaczewski (170), Saccardo ii:436: 2715. Host: Prunus angustijolia Marsh., P. hortulana Bailey, P. americana Marsh. Deforming shoots. Asci: Subclavate, rounded or truncate, 25-45x8-10; stalk cells rounded below, not inserted, 10-18 x 5-8. Spores; Elliptic. Distribution: North America. 72a. Var. tortilis Atk. Deforming fruits of P. angustijolia, affecting only one side of fruit. Asci and stalk cells usually a little longer than in species. 73. Taphrina cerasi (Fkl.) Sadeb. Syn.: Exoascus deformans f. cerasi Fkl.; Exoascus wiesneri Rathay. ; Exoascus cerasi (Fkl.) Sadeb. ; Taphrina gilgii Henn. and Lindau. ; Exoascus and Taphrina deformans Auct. in part. Fuckel (15), Rathay (31), Rostrup (71), Sadebeck (73, 80, 93), Hennings (82), Patterson (92), Giesenhagen (94, lOS), Palm (152), Jaczewski (170), Saccardo iO :69, No. 4734: ii:436. No. 2711. Host: Prunus avium L., F. ceras^is L., P. fruticosa Pall. Witches' brooms. Also, according to Nishida (140) on P. pseudoccrasus Lindl. var. sieboldi Maxim., and var. spontanea Maxim., and on P. miqueliana Maxim. Asci: Slender, clavate, rounded, 35-50x5-12; stalk cell 6-16x5-9. Spores: Ellipsoid 6-9 x 5-7. Distribution : Europe, North America, Japan. Mix: The Genus Taphrina II 169 74. Taphrina reichei Werd. Werdermann (158). Host: Prunus capollin Koehne. Leaf-curl. Asci: Cylindnc, obtuse, 26-34 xS-i2; stalk cell 15-18x8-10. Spores: Eight in ascus, subellipsoid, diameter 4-5.5. Distribution : Mexico. 75. Taphrina amygdali (Jacz.) n. comb. Syn. : Exoascus amygdali Jacz. ; Exoascus deformans (Berk.) Fkl. in part. Jaczewski (170). Host: Prunus communis (L.) Fritsch var. dulcis, var. amara. Mycelium : Intercellular. Asci: Hypophyllous, clavate, rounded, 24x9-11; stalk cell 11x8.3. Spores: Usually four to eight in ascus, globose, diameter 3-4. Distribution: Southern Europe. According to Jaczewski this fungus is distinct from the variety of T. de- jormans described by Campbell (165) from the same hosts. 76. Taphrina pruni Tul. Syn. : Exoascus pruni Fkl. Tulasne (14), Fuckel (15), Sadebeck (47, 73, 80, 93), Giesenhagen (94, 108), Patterson (92), Palm (152), Jaczewski (170), Saccardo 8:817, No. 3342. Host: Prunus domestica L., P. padus L., P. spinosa L. Deforming fruits. Also, according to Nishida (140) on P. tom,entosa Thunb., and P. incisa Thunb. Mycelium : Intercellular. Asci: Cylindric-clavate, rounded, 30-60x8-15; stalk cell 10-20x8, not in- serted. Spores: Eight in ascus, rarely more. Subglobose, diameter 4-5. Distribution: Europe, North America, Japan. Jaczewski would confine the species to the form on Prunus domestica and makes the following varieties: 76a. Var. divaricata Jacz. On Prunus divaricata Led. Russia. 76b. Var. padi Jacz. On fruit and twigs of Prunus padus L. Asci elongate- cylindric, rounded, 30-35x11; stalk cell 19x8. Spores globose, diameter 4. Everywhere that host grows. 76c. Var. ussuriensis Jacz. On Prunus u^suriensis. Asci cylindric, 41.5 x 11; stalk cell 19.4-24.9x5.5-8.3. Spores globose, diameter 5-6. Russia. 77. Taphrina minor Sadeb. Syn.: Exoascus minor (Sadeb.) Sacc. Sadebeck (73, 80, 93), Giesenhagen (94, 108), Palm (152), Jaczewski (170), Saccardo iO:70, No. 4736. Host: Prunus jruLicosa Pall. {P. chajyiaeceras us J acq.). Deforming leaves, buds, and branches. Also, according to Cunningham (161), and Palm (152), on P. cerasus L. and, according to Jaczewski (170), on P. avium L., P. cerasus L., and P. microcarpa C. A. Mey. Mycelium : Subcuticular. 170 The University Science Bulletin Asci : Terete-clavate, 30-35 x 6-8. Basal cell broader toward base, 8-10 diam. Giesenhagen gives dimensions of asci as 18-35x6-8, of stalk cells as 8-10x6-10. Spores: Eight in ascus, 6-7x5. Distribution: Germany, Russia, New Zealand (Siberia). 78. Taphrina truncicola Kusano. Kusano (124), Jaczewski (170), Saccardo i8:197, No. 3851. Host: Prunus incisa Thunb. Deforming shoots and inflorescences. Also, according to Nishida (140) on P. maximouiczi Rupr. Asci: Cylindric, rounded, 37-45x8.7-10; stalk cell cylindric, 13-20x10-13. Spores: Globose, diameter 4-5.8. Distribution : Japan. 79. Taphrina instititiae (Sadeb.) Johans. Syn. : Exoasrus insititiae Sacicl). Sadebeck (47, 73, 80, 93), .Tohar.son (52), Rostrup (71), Giesenhagen (94, 180), Palm (152), Jaczewski (170), Saccardo 8:817, No. 3344. Host: Prunus insititia L., and P. domestica L. Witches' brooms. In North America on P. pennsylvanica L. (Giesenhagen, Jaczewski). Asci: Cylindric-clavate, rounded, 25-30x8-10; stalk cell subcubical, sub- acute at base, inserted, 6-8x7-10. Spores: Globose, diameter 3.5. Distribution: Northern and Central Europe, North America. 80. Taphrina cerasi-microcarpae (Kiischke) Laubert. Syn. : Exoascus cerasi-microcarpae Kuschke. Kuschke (144), Jaczewski (170), Saccardo 2^:1301, No. 7550. Host: Prunus ynicrocarpa C. A. Mey. {Cerasus microcarpa Boiss.). De- forming fruits. Mycelium : Intercellular. Asci: Clavate, short stalked, 35-50x10-13. Spores: Globose, oval, or ovate, 5-7.5x5-6.25. Distribution : Transcaucasus. 81. Taphrina mexicana Sy(i. Sydcw and Sydow (150), Jaczewski (170), Saccardo 24:1302, No. 1557. Host: Prunus microphyUa Hemsl. Small witches' brooms, about 5cm. m diameter. Asci: Cylindric, round or more often truncate, 28-35x6-9; stalk cell 9-13 x 7-10. Spores: Eight in ascus, ellipsoidal. 3.5-4x2.5-3.5. Distribution: Mexico. 82. Taphrina mume Nishicia Yoshino (127), Nishida (140), Saccardo 22:705, No. 5877. Host: Prunus murnc S. and Z., P. nrmcninca L. var. ansu Maxim. Deform- ing shoots. Mix: The Genus Taphrina II 171 Asci: Cylindric, rounded, 25-52x8-15; stalk cell 8-12x5-15. Spores: Usually eight in ascus, globose, diameter 4-6. Distribution : Japan. 83. Taphrina dejormans (Berk.) Tul. Syn. : Exodscus deformans (Berk.) Fkl. ; Ascomyces deformans Berk.; Ascosporium deformans Berk. Berkeley (10), Tulasne (14), Fuckel (15), Rathay (28), Sadebeck (47, 73, 80, 93), Rostrup (71), Giesenhagen (94, 108), Patterson (92), Campbell (165), Jaczewski (170), Saccardo 8:816, No. 3341. Host: Prunius persica L., P. communis (L.) Fritsch, deforming leaves, twigs, and rarely flowers. Mycelium : Intercellular. Asci: Described by most authors as epiphyllous, but occasionally also hypophyllous (Mix 186), according to Jaczewski exclusively hypophyllous (!). Cylindric, rounded, 25-50 x 8-12 ; stalk cell 6-8 x 6-10. Spores: Usually 8, .sometimes 4 in ascus, globose, diameter 3-5. Distribution: World wide. The form on almond, noted by Rathay and by subsequent authors is described by Campbell (165) as a variety, without naming it as such. Asci are 34x11-12, without stalk cells; spores 7x5. 84. Taphrina pseudocerasi (Shirai) Sacc. Syn. : Taphria pseudocerasi Shirai Shirai (96), Jaczewski (170), Saccardo i^ :824, No. 3077. Host: Prunxis pseudocerasus Lindl., and P. miquelliana Maxim. (P. subhirtella Miq.). Witches' brooms. Asci : Terete-clavate, rounded, 40-44 x 6-9 ; stalk cell 6-9 long. Spores: Many in ascus, subglobose, 4.5-5x3.5-4.5. Distribution: Japan. 85. Taphrina andina Palm. Syn.: Exoascus andinus (Palm.) Sacc. and Trott. Palm (134), Jaczewski (170), Saccardo 22:764, No. 5871. Host: Pninus salicifolia B.BK. Spots on leaves. Asci: Epiphyllousor hypophyllous, cylindric, rounded or truncal o, 27-3 1 X 9.5-13.6; stalk cell cuboidal, 9.5-13.6x9.5-10.2. Spores: Eight in ascus, subglobose, 4.5x3.5. Distribution: Quito and Ambato, Ecuador. 86. Taphrina jarlowi Sadeb. Syn. : Exoascus farlowii (Sadeb.) Sacc. Sadebeck (73, 80, 93), Giesenhagen (94, lOS), Patterson (92), Jaczewski (170), Saccardo i0:70, No. 4737. Host: Primus serotina Ehrh., deforming fruits. Mycelium : Intercellular. 172 The University Science Bulletin Asci : Scattered, terete-clavate, rounded 20-30 x 8-9 ; stalk cell elongate, 15- 25 X 8-9, not inserted. Spores: Globose, diameter 4. Distribution: North America. 87. Taphrina varia (Atk.) n. comb. Syn. : Exoascus varius Atk. Atkinson (8C), Giesenljagen (94), Saclebeck (93), Jaczewski (170), Saccardo llAZQ, No. 2717. Host: Prunus serotina YAwh. Deforming shoots. Asci : 20-27 x 8-10, stalk cell 12-17 x 8-10. Spores: Not described. Distribution: Alabama, U. S. A. Perhaps also on P. demissa Walp. in Colorado (Atkinson). Giesenhagen makes this species synonymous with T. jarlowi, which, how- ever, occurs on fraits. They should be considered distinct species until further proof of their identity is forthcoming. 88. Taphrina rostrupiana (Sadeb.) Giesenhag. Syn. : Exoascus rostrupianus Sadeb. Sadebeck (80, 93), Giesenhagen (94, 108), Jaczewski (170), Saccardo iJ :435, No. 2708. Host: Prumis spinosa L., deforming finiits. Asci: Cylindric-clavate, rounded, 35-50x7-8; stalk cell variable, 10-16x2-6 or 1.5, often pointed below but not inserted. Spores: Eight in ascus, ellipsoid, 6-7x3-4. Distribution : Europe. 89. Taphrina pruni-subcordatae (Zeller) n. comb. Zeller (177). Host : Prunus subcordata Benth. Deforming fruits. Asci: Clavate, often truncate, sometimes narrowed above, 54-61x10-13; stalk cells not inserted, 12-16x5-11. Spores: Ellipsoid to ovate, 6-10x3.5-5. Distribution: Oregon, California, U. S. A. 90. Taphrina rhizipes (Atk.) Giesenhag. Syn. : Exoascus rhizipes Atk. Atkinson (86), Sadebeck (93), Giesenhagen (94, 108), Jaczewski (170), Saccardo ii:436, No. 2716. Host : Prunus triflora Roxb., deforming shoots and fruits. Asci: Clavate, 30-40x8-10; stalk cell 25-40x3-5. Rhizoidal outgrowths from asci and basal cells penetrate between epidermal cells. Spores: Not described. Distribution : Alabama, U. S. A. Mix: The Genus Taphrina II 173 91. Taphrina ceddomophila (Atk.) Giesenhag. Syn. : Exoascus cecidomophilus Atk. Atkinson (86), Sadebeck (93), Jaczewski (170), Saccardo Ji :436, No. 2718. Host: Prunus virginiana L., occurring on insect galls on fruits. Mycelium : Intercellular, but superficial. Asci: Cylindric or rarely clavate. 30-40 x 6-10; stalk cell broad, rountled below, 6-10 X 10-15. Spores: Not described. Distribution: New York, U. S. A. 92. Taphrina confusa (Atk.) Giesenhag. Syn. : Exoascus confusus Atk. Atkinson (86), Giesenhagen (94, 108), Sadebeck (93), Jaczewski (170), Saccardo 11AS6, No. 2712. Host: Prunus virginiana L. Deforming fruits and floral envelopes. Asci: 30-45 x 8-12; stalk cell 15-30 x 6-10. Spores: Not described. Distribution: North America. 93. Taphrina unilateralis (Pk.) n. comb. Syn. : Exoascus unilateralis Pk. Peck (102), Jaczewski (170), Saccardo i6 :803, No. 2920. Host: Prunus virginiana L. Leaf curl. Asci: Subcylindric, slightly contracted above basal cell, 40-52 x 13-16; stalk cell 13-16 broad and about as long. Spores: Eight in ascus, globose or broad elliptic, 6-6.5 x 5-8. Distribution: New York, U. S. A. 94. Taphrina bullata (Berk, and Br.) Tul. Syn.: Taphrina bullata (Berk, and Br.) Sadeb. ; Exoascus bullatus (Berk, and Br.) Fkl. ; Oidium bullatum Berk, and Br.; Ascomyces bullatus Berk.; Ascosporium bullatum Berk. Berkley (8), Tulasne (14), Sadebeck (73, 80, 93), Johanson (52), Rostrup (71), Giesen- hagen (94, 108), Palm (152), Jaczewski (170), Saccardo 8:817, No. 3343. Host: Pyrus communis L., leaf curl. Also on Chaenomeles lagenaria Koidz. (Rostrup). Mycelium : Subcuticular. Asci: Cylindric, truncate, 36-40 x 8-9; stalk cell 10-15 x 8-9. Spores: Globose, diameter 5. Distribution: Europe. 95. Taphrina piri Kusano. Syn. : Exoascus piri (Kusano) Sacc. and Trott. Kusano (124), Jaczewski (170), Saccardo iS :197, No. 3852 and 22:764, No. 5870. Host: Pynis miyabe Sarg. Spots on leaves. 17-1 The University Science Bulletin Asci: Plurap-cylindric, rounded-truncate, 29^42 x 9-13; no stalk cell. Spores: Globose, diameter 3-5. Distribution : Japan. 96. Taphrina caerulescens (Desm. and Mont.) Tul. Syn. : Exoascus coerulescejis Sadeb. ; Ascoviyces caerulescens Desm. and Mont. ; Ascomyces quercus Cke. ; Ascomyces alutaceus Thm. ; Ascomyces extensus Pk. ; Taphrina extensa (Pk.) Sacc. ; Taphrina alutacea (Thum) Sacc. ; Taphrina quercus (Cooke) Sacc. Desmazieres (6), Tulasne (14), Cooke (26), Thunien (33), Saccardo (42), Sadebeck (47, 73, 80, 93), Peck (51), Johanson (56), Patterson (92), Giesenhagen (94, 108), Palm (152), Jaczewski (170), Saccardo 8:814, No. 3332, 3333, 815, No. 3334. Host: Quercus spp. Spots on leaves. Asci: Often amphigenous, broad-cylindric, rounded or truncate, abruptly attenuate at base, with rhizoidal appendage, 55-70 x 15-20; no stalk cell. Spores: Numerous (conidia) 2.5-3 x 1.5-2. Distribution: Europe, N. America. 97. Taphrina rubrobrunnea (Pk.) Sacc. 8yn. : Ascomyces rubrobrunneus Pk. Peck (58), Jaczewski (170), Saccardo iO :67, No. 4724. Host: Quercus rubra L. Spots on leaves. Asci : Oblong, truncate, 50-75 x 15-22. Spores: Numerous (conidia) subelliptic, 3-4x1.5-2. Distribution: Sandlake, New York, U. S. A. ' 98. Taphrina kruchii (Vuill.) Sacc. Syn. : Exoascus kruchii Vuill. Kruch (69), Vuillemin (75), Sadebeck (80, 93), Giesenhagen (94, 108), Jaczewski (170), Saccardo 10 :68, No. 4727. Host: Quercus ilex L. Witches' brooms. Mycelium : Subcuticular. Asci: Subclavate, 72-80x21-23; no stalk cell. Sadebeck and Giesenhagen give dimensions of asci as 65-75 x 15-20, broadening below to a width of 30-40. Spores: Eight in ascus, globose, diameter 4, conidia 2.5x2. Distribution : France and Italy. 99. Taphrina randiae Rehm. Rehm (112), Jaczewski (170), Saccardo i6:1152. No. 4790. Host: Randia sp. Spots on leaves. Asci: Sessile, between epidermal cells, oblong, truncate, narrowed at base, 50x25; no stalk cell. Spores: Globose, diameter 3-4. Distribution: Serra Orgaos, Brazil. Mix: The Genus Taphrina II 175 100. Taphrina purpurascens Robins. Syn. : Exoascus purpurascens (Robins.) Sacc. ; Ascomyces deformans (Berk.) var. purpurascens Ell. and Ev. Robinson (55), Sadebeck (80, 93), Giesenhagen (94. 108), Patterson (92), Jaczewski (170), Ellis and Everhart, North American Fungi No. 1886, Saccardo 8:819, No. 3353. Host: Rhiis copallina L. leaves. Asci : Dumbbell-shaped, constricted in the middle. 24-32 x 9-14, 6-11 broad in constricted part, 9-21 at base; no stalk cell. Spores : Ellip.soidal, 3.5-5 x 2.5-4. Distribution: North America. 101. Taphrina sebastianae (Sadeb.) Jacz. Exoascus sebastinae Sadeb. Sadebeck (120), Jaczewski (170), Saccardo 22:764, No. 5872. Host: Sebastiana brasiliensis (Mull.) Arg. Leaf curl. Asci: Polymorphic, narrow-cylindric, 90-130x8, or 70-80x6-9 and 2-3 broad at base; or clavate to oblong-ellipsoid, rounded or subtenuate, 50-90x9-20 (in- cluding stalk cell) ; stalk cells one or two. Spores: Eight in ascus, ellipsoid, 5.5-6.5x4.5-5. Distribution: Tubarao, Brazil. 102. Taphrina sorbi (Jacz.) n. comb. Syn. : Exoascus sorbi Jacz. Jaczewski (170). Host: Sorbus terminalis Crantz. Spots on leaves. Mycelium: Subcuticular. Asci: Hypophyllous, clavate, truncate, 16-20x9-11; stalk cell flat, not in- serted, diameter 6-8. Spores: Usually eight in ascus, diameter 3.7-4; conidia 3.7x1.8. Distribution : Caucasus. 103. Taphrina bussei v. Fab. Syn. : Exoascus bussei v. Fab. ; ? Exoascus theobromae Ritz. Bos. Von Faber (129, 131), Ritzema Bos. (106), Jaczewski (170), Saccardo 22:764, No. 5869. Host: Theobroma cacao L. Witches' brooms. Mycelium : Intracellular. Asci : On leaves only, amphigenous, 15-17 x 5. Spores: Eight in ascus, ellipsoid, 2.5x1.7. Distribution: Kamerun, South Africa. (Surinam?). 104. Taphrina ulmi (Fkl.) Johans. Syn. : Exoascus ulmi Fkl. ; Exoascus campestris Sacc. Fuckel (17), Saccardo (41), Sadebeck (47, 73, 80, 93), Johanson (52), Rostrup (71), Gie.senhagen (94, 108), Patterson (92), Palm (152), Jaczewski (170), Saccardo 8:819, No. 3351 ; 820, No. 3356. 176 The University Science Bulletin Host: Ulmus campestris L., U. glabra Mill., U. montana With., U. suberosa Monch., U. americana L. Spots on leaves. Asci: Cylindric, or oval, rounded, 12-20x8-10; stalk cell very broad, 3-8 X 15-20. Spores: Eight in ascus, globose, diameter 3.5. Distribution : Europe, North America. DOUBTFin. SPECIES 1. Taphrina githaginis Rostr. Rostrup (71), Sadebeck (73), Saccardo 10:68, No. 4728. Host: Agrostemma githago L. Swellings of stems and leaves. Mycelium : Intercellular. Asci: Ellipsoid, 48-58x30-45; no stalk cell. Spores : Numerous, 4-6 x 2-3. Distribution : Europe. Apparently belongs in the Protomycetaceae. EXCLUDED SPECIES 1. Exoascus anomalus Sacc. Is Ascocorticium albidum Bref. 2. Taphrina candicans Sacc. on Teucrium chamaedrys L. Not a fungus, but the effect of mite injury (Jaczewski, 170). Ex- cluded by Giesenhagen (94). 3. Taphrina cissi Zoll. On Cissus sp. Never recognized by any student of the genus. 4. Exoascus fulgens (Cke.) Sacc. {Ascomyces fulgens Cke). on Arctostaphylus sp. An insect injury. Excluded by Giesen- hagen (94). 5. Taphrina marginata Lamb et Fautr. On Crataegus oxyacan- tha L. Not a fungus, but injury due to Erineum mites. (Palm 152, Jaczewski 170.) 6. Taphrina rhaetica Volk. on Crepis blattariodies Vill. Belongs to Taphridium (Volkartia) of the Protomycetaceae. (Juel 157.) 7. Taphrina umhellijerarum Rostr. {T. oreoselini Massal.) Is Taphridium umhellijeraram (Rostr.) Lagerh. and Juel. (Juel 157.) 8. Exoascus Quercus lobatae Mayr. No fungus was ever seen. THE UNIVERSITY OP KANSAS SCIENCE BULLETIN YoL. XXIV.] July 15, 1936 [No. 11. Two New Sunfish of the Family Centrarchidae from the Middle Pliocene of Kansas CLAUDE W. HIBBARD, The Kansas University Museum of Vertebrate Paleontology' Abstract: Two new species of Centrarchid fishes. Chaenobryttus kansasensis sp. nov., and Pomoxis land sp. nov., are described from specimens in the Kan- sas University Museum of Vertebrate Paleontology. Both species are from the diatomaceous marl bed, Middle Pliocene, Logan county, Kansas. DURING the summer of 1924, Mr. H. T. Martin made his first visit to what is known as "Rhino Hill Quarry" in Wallace county, Kansas. While visiting this locality he discovered the white diatomaceous marl bed near the Marshall ranch. In this bed he found a few scattered fish scales and a few dicotyledonous leaves. This information was given to Dr. M. K. Elias of the Kansas Geo- logical Survey. Doctor Elias in his intensive study of the geology of Wallace county, located a rich deposit of fossil fish in the diatoma- ceous marl, which is associated with a deposit of dicotyledonous leaves. The latter deposit was first opened in the summer of 1931 iind only one day was spent collecting in this locality. The quarry is located at sec. 7, T. 11 S., R. 37 W. in Logan countj^, about a mile east of the "Rhino Hill Quarry," which lies near the cast line of Wallace county and is at the base of the diatomaceous marl. The marl was laid down in a large, deep, clear lake that ex- isted during the middle Pliocene. It is interesting that under the lake bed is the old river channel filled with sand and assorted gravel, iu which the "Rhino Hill" fauna is found in Wallace county. The geology of this area has been fully discussed by Doctor Elias in "The Geology of Wallace County, Kansas," 1931, Kansas Geol. (177) 12—6037 178 The University Science Bulletin Surv. Bull., No. 18. An excellent detailed description of this dia- tomaceous marl is also given by Doctor Elias in Circular No. 3, Kansas Geol. Surv., 1931, "Diatomaceous Marl from Western Kan- sas, a Possible Source of Hydraulic Lime." In collecting the specimens it was found that the procedure used in preserving many other types of fossils could not be used with those from the diatomaceous marl. The marl has been exposed to weathering, and has been attacked by the roots of plants from above which have caused it to crack in many pieces. In many places sur- face water has worked down through the joint planes, carrying sedi- ment, and making it almost impossible to secure unweathered or complete specimens. Tliere are also fine layers of clay intercalated in the marl, which hold moisture for a considerable time, so that upon exposure to the dry, hot climate of that area tlic clay dries rapidly, warps and checks, causing the thin layer of marl above to slough off, oftentimes destroying good specimens. The marl itself, if damp when first exposed to the atmosphere, likewise loses its moisture rapidly, causing it to crack and form fine laminae, destroy- ing fossils in it. Great caution must be used in removing the fossils and special care must be taken to place them in a shaded area im- mediately upon removal. The fossils cannot be treated with gum- arabic or shellac solution with any success, since this treatment causes them to break into many pieces. Due to apparently close relationship of these forms to our modern fishes, it seems advisable at present to refer the following species to living genera, since many of the characters by which our modern genera are distinguished are those not found preserved in fossil forms. These species have been placed with living genera because they have certain skeletal characters in common. Future collecting may provide better preserved material which will further substan- tiate this association or give sufficient grounds for separation. Other species from this fauna will be described when sufficient modern material is available for comparative study. Chaenobryttiis k(tn!^asensis, sp. nov. (Platf VI) Holotype. No. 792F, University of Kansas Museum of A^-rtcbrate Paleontology. Complete skeleton, lacking part of premaxillae. Horizon (Old type locality. From the Diatomaceous Marl Bed, Middle Pliocene. Collected on sec. 7, T. US., R. 37 W., Logan county, Kansas, by the University of Kansas Museum Expedition, summer of 1931. Hibbard: Two New Sunfish 179 Diagjiosis. Body sunfish-shaped, oblong and slightly arched in front of dorsal fin. Vertebrae 13 + 16. D. X, 12; A. Ill, 8; V. I, 5. Length of attachment of anal fin less than one half of dorsal fin at- tachment. Longest anal and dorsal spines over half as long as soft rays. Longest dorsal spine equal to distance from snout to about the center of the eye. Neither pectoral nor ventral fins reaching anal fin. Maxillary, premaxillary, vomer, and pterygoid bearing teeth. Scales ctenoid. Caudal fin emarginate. Description of type. (See plate VL) The specimen is that ap- parently of a young adult fish. Length of fish without tail, 94 mm. Length of head from end of snout to atlas, 31 mm. Head 3 in length; depth, 2%. Depth at caudal peduncle, 13mm. The dorsal fin attachment is 35 mm. long. The lengths of dorsal spines are as follows : I, 8 mm. ; II, 10 mm. ; III, 12.5 mm. ; IV, 13.5 mm. ; V, VI, VII, VIII, IX, X, respectively, 14 mm. A distance of 18 mm. sepa- rates the posterior attachment of the dorsal fin from the caudal fin. The head is badly crushed, but the dentition of the maxillaries and premaxillaries is very distinct. They are set with rows of small sharp conical teeth. On the anterior part of the maxillaries and pre- maxillaries in the outer row the canines arc well developed. These are about three times the size of the small conical teeth. The vom- erine and pterygoid teeth are small and conical. Only four branch- iostegals are visible. The pectoral fin shows only ten rays, which are nearly hidden by the ribs. The ventral fin (I, 5) is so broken that only about half of the well-developed spine is present. The soft rays reach nearly to the first anal spine. The anal fin is well developed and is attachetl almost directly below the IX dorsal spine. The length of the attachment of the anal fin is 16 mm. and the posterior point of attachment is 17 mm. from the caudal fin. The anal sinnes are well developed and heavier than the dorsal spines. Their lengths are as follows: I, 11 mm.; II, 13 mm.; Ill, 15.75 mm. Three interneurals are present anterior to the dorsal spines. Pom oris lanei, sp. nov. (Plate vin Holotijpe. No. 789F, K. U. IM. V. P. Complete crushed skeleton lacking most of caudal fin. Horizon and type locality. From the diatomaceous marl bed, Mid- dle Pliocene. Collected on sec. 7, 11 S., R. 37 W., Logan county, Kansas, by the LTniversity of Kansas Museum Expedition, summer of 1931. 180 The University Science Bulletin Diagnosis. Body sunfish-shaped, obiong, and symmetrical. Ver- tebrae 14 + 17. D. VII, 15; A. VI, 12; V. I, 5. Anal fin nearly as long as dorsal. Longest anal and dorsal spines over half as long as soft rays. Scales ctenoid. Description of type. Complete impression of a small sunfish, with the exception of the tip of the caudal fin ; many of the bones and fin rays present. The skull is badly crushed. The dentary and pre- maxilla are covered by rows of sharp conical teeth. The vomer, palatines, pterygoids and pharyngeals are missing or so crushed that they cannot be studied. Length of the head from the tip of the pre- maxilla to the posterior border of the opercular, in line with the vertebrae, 25 mm. Only three branchiostegals are visible. The pectoral fin is so folded under the body that it is impossible to study it in detail. The ventral fin (I, 5) is attached 11.5 mm. anterior to the anal fin. The spine is well developed, being 9.5 mm. long. The soft rays just reach the first spine of the anal fin. The length of basal attachment of the anal fin is 20.5 mm. The first anal spine is attached directly under the attachment of the fourth dorsal spine. Anal spines I and II are rather slender. The lengths of the anal spines are as follows: I, 3.5 mm.; II, 5 mm.; Ill, 6 mm.; IV, 9 mm.; V. 10.5 mm. ; VI, 12 mm. The first soft rays of the anal fin are 15 mm. long. The soft anal rays reach only to the center of the caudal peduncle. The dorsal fin has a basal attachment of 22 mm. The soft rays are longer than the longest dorsal spines. The lengths of the dorsal spines are as follows: I, 4 mm.; II, 6 mm.; Ill, 10 mm.; IV, 11 mm.; V, VI and VII, respectively, 13 mm. Six interneurals are present anterior to the first dorsal spine. Vertebrae (impres- sions) 14 -j- 17. Only four vertebrae are actually present. The greatest depth of 26 mm. is just anterior to the dorsal fin. Length of fish to last caudal vertebrae is 70 mm. The skeleton is that of a young fish. The paratype is the skeleton of a still younger fish 45 mm. in length to the last caudal vertebrae. The caudal fin is entirely lack- ing. The skeleton is nearly complete except for the badly crushed condition of the head. Small conical vomerine teeth are present. The ventral, anal and dorsal fins are the same as in the holotype. The spinous rays have the same proportionate length. Pectoral fin shows at least ten rays. Vertebrae are complete, being 14 + 17. The eye is contained four times in the length of the head. Remark. This species is named for Dr. H. H. Lane, under whose guidance the expedition worked the summer of 1931, and who has offered many helpful criticisms and suggestions. 182 The University Science Bulletin PLATE VI Chaenobryttus kfui.soscnsis, sp. nov., nearly complete skeleton, K. U. Mus, Vert. Paleo. No. 792F. Approximately Vs natural size. See text for measure- ments and description. Hibbard: Two New Sunfish 188 PLATE VI •^ f' ■^ '>,«^Vjf^ X V. "i(.'-".-« \ J .4 184 The University Sc:ence Bulletin PLATE VII Ponioxis land, sp. nov., nearly complete skeleton, K. U. Mus. Vert. Paleo. No. 789F. Approximately natural size. See text for measurements and descrip- tion. Hibbard: Two Xew Sunfish 185 PLATE VTI ^^^V. THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 12. The Family Apioceratidae (Diptera) in North America* REGINALD H. PAINTER Maiiluittim, Kansas Abstr.\ct: Notes on most of the type series, keys to all species, and ad- ditional distribution records are given for the known North American species. The following new species of Apiocera are described: trimaculato, calons, aldrichi, bcamert, clnvator, iidvrrnpla, notnla, coiivcrgeiu, martinorum and figures are given of genitalia and other characters. There are seventeen de- scribed species in this family in North America. IN a recent review of the genus Apiocera (4) the writer redescribed the two previously known North American spec'cs <^f t' - • '-pniis and added one new species. Through the kindness of the late Dr. J. M. Aldrich, United States National Museum, it has been possible to study a number of additional specimens representing six new species of this genus. During a recent ir'ip for the study of the types of Bombyliidaef it was jiossible to examine also all of the type specimens of the family Apioceratidae which still exist in North American museums. A few notes on these have also been included. These records have extended the range of several species and our knowledge of this unusual family. The species described here bring the total munber of North American species to seventeen and the total known for the world to twenty-five. * A study of the material in the Francis Huntington Snow Entomological Museum, Uni- versity of Kansas, the United States National Museum and several others. This is Con- tnl)Ution No. 41.5, from tlie Department of Entomology, Kansas State College. t The writer is indebted to the directors of the Bach fund for funds to permit the ex- amination of these types. The author is also indebted to the following persons for the loan of personal collections or specimens in their care: Dr. J. M. Aldrich, Dr. R. H. Beamer, Messrs. E. P. VanDuzee, C. H. Martin. J. Wilcox, E. R. Tinkham, F. M. Hull and E. T. Cresson, and to his wife fur making the illustrations which are given here. (187) 188 The University Science Bulletin RHAPHIOMIDAS A study has been made of a few specimens of each of the de- scribed species of this genus and of the types which still exist. The key given below is adapted from Townsend (5) : 1. First anal cell open widely Apomidas troch'dus Coq., 188 First anal cell closed (Rhaphionuda^) 2 2. Abdomen wholly brown, wings with a smoky tinge, antennae blackish . .episcopusO. S., 188 Abdomen not wholly black; wings hyaline, antennae yellowish or reddish, at least on the third joint 3 3. Abdomen almost wholly yellow santhos Townsend, 189 None of the abdominal segments entirely yellow, the second and fourth each partly yellow, partly blackish 4 4. Ground color of abdomen yellowish, only the front border and a median spot on sec- ond to fourth segments blackish; bristles of legs and scutellum yellowish, .acton Coci., 188 Ground color blackish, only the hind border of second to fourth segments yellowish ; bristles of hind border of scutellum and middle and hind legs blackish, mellijex Townsend, 189 1. Apomiflas trochilus Coc}. (2)* This species is included in the key on account of its great simi- larity to Rhaphiomidas. The type female in the United States National Museum lacks the antennae and is greasy. It is in all respects like R. acton or xanthos except in the open first and anal cell. A male intermediate in this character is mentioned under R. acton. It appears that this single character is hardly sufficient grounds for generic segregation. No other specimens appear to have been collected or recorded and additional material will probably be necessary before the exact generic status can be determined. 1. Rhaphiomidas episcopus 0. S. (3) (5) The type female in the Museum of Comparative Zoology is very greasy, but so far as is visible, the identification and description given by Townsend from fresh specimens is correct. The black pile, mentioned by Osten Sacken, on the last three segments consists of restrose hairs similar to those found on female Apiocera. The antennae are like those of acton in size and shape. There is a female irom the Townsend collection in the University of Kansas collection taken at El Taste, Baja Cal., Sept. '95 (Eisen). 2. Rhaphiomidas acton Coq. (1) The type male is in the United States National Museum. The several specimens which have been seen vary considerably in re- spect to the amount of black on the abdomen. In a typical male * Numbers refer to literature cited. Painter: The Family Apioceratidae 189 specimen the antenna measured first joint, 0.65 mm.; second joint, 0.25 mm.; third joint. 1.85 mm. The third segment is elongate oval in shape. The following specimens have been seen: Three males, two females, Cajon, Cal., July 24, 1930 (T. F. Winburn and R. H. Painter, coll. I ; several specimens "California." A male, Mohave Desert, Cal., July 20, 1930 (C. H. Martin, Coll.), is apparently this species. The first anal cell is narrowly open and in this respect is about half way between the typical acton and that in the type of .4. trochilus Coq. The genitalia and other structures are identical with other males of acton; the amount of black color- ation on the abdomen is a little less than usual. 3. Rhaphiomidas xanthos Townsend (5, 6) The type of this species was destroyed in the San Francisco fire in April, 1906. There are, however, three males in the Snow collec- tion of the University of Kansas which were received from Town- send and appear to belong to the type series. They are labeled "type" in the same handwriting as are the specimens of mellifex. The locality is given as "El Taste, B. C. Sept. (Eisen)." A single teneral male from the type locality is in the National INIuseum. The type series consisted of one male mentioned at the first of the de- scription and ten others mentioned later. These were said to be from San Jose del Cabo. The two localities are very close together and it appears that the author designated the town rather than the moun- tain as the type locality. The only specimens seen in addition to those reported by Town- send and Colquillett were a male and female. State College, New Mexico, Sept. 23 (W. J. Beargl. The genitalia are different from mellifez and acton, but the antennae are like the former species in size and shape. In the New Mexico specimen the first joint of the antenna measured 0.5 mm.; second, 0.35 mm.; third, 1.2 mm. The third joint is "flask-shaped" with the neck near the second joint. In one of the Townsend specimens the anal cell is narrowly open in one wing and closed in the other. In both the other specimens it is closed and more or less petiolate. 4. Rhaphiomidas mellifex Townsend (5, 6) The type of this species was also destroyed in the San Francisco fire. In the Snow collection at Kansas University there are two fe- males which apparently belong to the type series. Both are marked 190 The University Science Bulletin "type" in the same handwriting as are other types in the Townsend collection there. These two specimens are labeled "El Taste, L. Cal., Sept." This locality reference is evidently to a mountain very near San Jose del Cabo, the type locality mentioned in the original de- scription. In the Snow collection there is also a female, San Jacinto Mts., Cal., 7-21-29 (L. D. Anderson), which is identical with these others except that the third joint of the antennae is a little longer. In this species this joint is like xanthos, thus shorter and broader than in episcopiis or acton (text, figure 1). Genus Apiocera In this genus the males and females sometimes differ appreciably in coloration. The males are easily distinguished either on the basis of abdominal coloration or by the structure of the genitalia. The females of several species, however, look much alike and it is only by comparative study that it has been possible to identify these with certainty. The shape and vesture of the ninth sternite and tergite and of the eighth somite provide excellent characters for distinguishing the males of the several species. These structures are figured for all the North American species. The details of head, thorax and wings appear to present little of use in distinguishing the species. The structure of the vesture was discussed in a previous paper (4). The key given below is an attempt to provide for identi- fication of both males and females where possible. J\Iain de- pendence must be ])laced on the structure and coloration of the males for correct determinations (Plate Villi. Key to the Species of Apiocera in North America 1. Metupleura with a more or less conspicuous tuft of hair in front of the spiracle. . 2 Metapleura bare or uniformly short pilose 5 2. Dorso-caufla! angle of mesopleura with a tuft of white hair (see also martiiioruin) triinaiiilata. u. sp., 195 Dorso-caudal angle of mesopleura with three or four strong setae 3 3. Setae of body and legs largely white, hairs of thoracic dorsum white, raloria, n. sp., 194 Setae of body and legs largely black, hair? of thoracic dursvuu white 4 4. Abdominal segments 2, 3, and 4 of male banded black and white, .aldricin, n. sp., 19.3 Abdominal segments 2 and 3 of male spotted lilack and white or forming irregular bands as in haruspei beaineri, n. sp., 198 .'). Dorso-caudal angle of mesopleiua uitli tluce or fdur .-^tmiii; black setae (see, also, augur) cinvator, n. sp., 196 Dorso-caudal angle of mesopleura bare or slightly hairy 6 (>. Mo.st of short hairs on mesonotum and male genitalia lilack in color; setae mostl\- black 8 Most of short hairs white on mesonotum, and male genitalia; setae white 7 7. Abdominal spots convex above, straight below; pulvilli usually more than half as long as claws; two black spots on each side in females hitn-riiptii. n. sp., 192 -■Wxlominal spots quadrate; pulvilli about half as long as claws; three Ijlack spots on each side in female bilineata Painter, 191 Painter: The Family Apioceratidae 191 8. Dorsum of abdominal segments 3 and 4 in male broadly black 9 Black on segments 3 and 4 of male forming spots 10 9. Male genitalia brown, ninth tergite notched at apex hanispex O. 8., 191 Male genitalia black, ninth tergite not notched at apex notata, n. sp., 199 10. Thorax mostly gray pollenose; dark brown of abdominal segments not extending across dorsum in male; sometimes almost lacking or confined to two small trian- gular spots on segments two and three augur O. S., 192 Thorax brownish pollenose; in male two white stripes the length of the abdomen show conspicuously against the dark brown, three black, subquadrate spots on each side in female convergeiis, n. sp., 19(i Thorax gray or brownish jiollenose ; in the male a central lilack stripe and lateral black stripes constricted or interrupted at the base or apex of segments three and four; usualK- four triangular spots on abdomen of female martiiioruin, n. sp., 197 1. Apiocera bilineata Painter (4) Drawings of the male genitalia of this species are shown on Plate VIII. In studying over the specimens previously reported the long hairs on the ventral lobe of the genitalia have been found to be black in cigiit specimens, orange-yellow in four and with some of each color in seven specimens. Variation in this character has been found in other species, notably haruspex 0. S. A male and a female in the Jas. S. Hine collection from Bill Williams Fork, Ariz. (F. H. Snow), appear to be the same species, but both specimens lack the genitalia. In the male the pulvilli are longer than usual for this species and the abdominal stripes are hardly interrupted at the incisures. It is possible that these ma}^ represent another species, possibly an undescribed one. One male. White Sands, N. Mex., June 30, 1932, R. H. Beamer collector; two females. White Sands, N. Mex., July 23, 1933 (W. Benedict), in the Kansas University collection, belong to this species. There is also a male in the same collection from Cameron Co., Tex., August 3, 1928, that appears to belong to biline- ata. It differs from the other specimens in a few minor characters. 2. Apiocera hanispex 0. S. (3, 4) This species appears to be the most widely distributed of those in Nortli America. The type has been studied and it agrees with the previous descriptions and the genitalia drawing given here (Plate VIII). Specimens have been seen from the following additional localities: Californu: 1 male. Yo.semite, AugiLSt 3. Hall and Hall coll., in J. ^^'ilcox collection; 1 female, Mt. Wilson, August 30; and 5 males, 1 female, Monrovia Canyon, August 16, 24. 29, 31 and September 1, C. H. Martin collector; 3 males, Beautivi.sta Canyon, July 5. C. H. Martin collector, and in his collection; 1 male, Pasadena, August 1. 1898, Grinnell (United States National Museum); 1 male, Tuolumne Co., Cal., July 29. 1926, E. R. Leach; 1 male. 2 females. San Diego Co., Cal., July, 1891. F. E. Blaisdell ; 3 males, Idyllwild, Cal., July 2 to 8, 1928, E. C. ^'an Dyke; 1 male, Oroville, Cal., July 15, 1926, H. H. Keifer 192 The University Science Bulletin (Cal. Acad. Sci.). Two males, 1 female, Xipomo, July 24, 1935; 1 male. Cajon Pass, August 1, 1935; 1 female, Indio, August 8, 1935; 2 females, San- Diego, August 7, 1935, J. Russell collector; 1 female, Idylhvild, August 3, 1935, E. I. Beamer collector; 3 males. 6 females, Cuyama Ranch, July 23, 1935,. J. Russell, Jack and R. H. Beamer, collectors. (In Snow Collection). Idaho: One male, one female, Lewistown, August 2, 1912 (U. S. N. M.) ; five males, three females, Lewistown, Idaho, July 14-22. 1925, C. L. Fox col- lector. Washington: One male, Eastern Washington (U. S. N. M.). Wyoming: Kahlotus, July 30, 1924, M. C. Lane, collector (J. Wilcox col- lection). British Columbia: One male, Oliver, July 24. 1923, P. N. Broom, col- lector (U. S. N. M.). Oregon : Three males, Adrian, July 22. 1934, C. H. Martin, Dorothy Martin. 3. Apiocera auger 0. S. (4) The Bembecine wasps, mentioned in the paper (4) as catching specimens of this species, have been determined by Richard Dow as Stictiella tennicornis (Fox). Drawings of the genitalia of A. augur are shown in Plate VIII. The following additional specimens of this species have been identified: Texas: Presidio, 2 females, July 29, 1928, 1 male, 1 female, August 16, 1929 (E. R. Tinkham collection) ; 1 male. Fort Stockton, July 18, 1927, L. A. Stephenson collector in Kansas University collection; 1 male, Brewster Co., June 24, 1929, J. B. Parks collector in J. Wilcox collection. New Mexico: 1 male, Mesilla Park, May 21, Cockerell (U. S. N. M.); 1 male, Eddy Co., N. Mex.. July 11. W. Benedict (Snow Coll.). A male and a female collected by F. M. Hull on Galveston Island, June, 1926, appear to belong to this species. The male genitalia differ only in some details which may be clue to position. The abdo- men is rubbed but seems to have had the typical pattern. Both sexes, however, have small spines on the dorsocaudal angle of the mesopleura. This is the most easterly record of this family. 4. Apioce7-a interrupta, n. sp. A pale, white pollenose species with white setae and with lateral rows of six abdominal spots which are straight on the lower and convex on the upper margin. Metapleurae and mesopleurae bare. Genitalia pale brown, similar to bilineata. Length, 19 nun. Male. Ground color, pale brown or yellow; mesonotum and last antennal joint dark brown or blackish. Pollen entirely white with perhaps a trace of a darker pattern on the mesonotum. All setae and pile white except for a little blackish or brownish pile on the underside of the male genitalia. A series of lateral spots commenc- Painter: The Family Apioceratidae 193 ing at the second segment and diminishing in size posteriorly are black on segments two to four and brownish on five to seven. These spots are convex above and straight in outline below, but vary some- what in size and shape. They are in approximately the same posi- tion as in bilineata. The genitalia are very similar to bilineata, dif- fering principally in the shape of the ninth sternite, less acute at the end and with a broader incision on the inner margin. Pulvilli about three fourths the length of the claws, wings hyaline, veins yellow. Female. Similar to male; retrose hairs on the last four abdominal segments white or yellowish. Spines of ovipositor brown. The black spots on abdominal segments two and three are subquadrate, and there is a trace of a brownish spot on the sides of segment four. Hototype. Male, Los Angeles, Cal., D. W. Coquillett collection. Allotype. Female, Los Angeles, Cal., D. W. Coquillett collection. Paratypes. Male, San Diego Co., Cal., D. W. Coquillett collec- tion. (In United States National Museum, Cat. No. 51432) 3 fe- males, Indio, Cal., August 5, 1935. (Jean Russell and Jack Beamer.) (In Snow collection.) 5. Apiocera aldrichi, n. sp. A robust, densely white pilose and pollenose species with the dor- sum of abdominal segments two, three and four jet black in ground color in the male. There is a subtriangular black spot in the female in the center of each side of segments two and three. Setae mostly black. Metapleura with a conspicuous tuft of hair in front of spiracle; dorsocaudal angle of mesopleura with three or four strong setae. Length, 23 mm. Male. Ground color of body and head black, last two joints of antennae, palpi, first and second tibiae, and abdomen beyond the fourth segment, brownish. Except for the genitalia and parts of the dorsum, the body is clothed with a thick mat of pollen consisting of minute curly hair, and in many places with erect thin hairs. The latter are especially abundant and long on the lower part of the head, coxae and first four abdominal segments. Dorsum of thorax brown pollenose with short brown pile and streaks of gray pollen which form the usual pattern. Setae of body and legs black ; of head white. Abdomen white pollenose; segments two, three, and four with quadrate black spots which are black pilose. In front of each of these, the margin is grayish, behind white. Separated from these 13—6037 194 The University Science Bulletin spots and on the same segments are smaller subquadrate blackish spots along the ventrolateral margins of the tergites. Dorsum of segments one and eight brownish. Genitalia brownish, pile black. Wing hyaline. Fejnale. Similar to male. The pattern on the thorax is less prominent and the body setae are partly white. The abdominal pile is not so long as in the male. Abdomen white pollenose; brownish in center of dorsum with an ill-defined subtriangular spot on each side of segments two and three. A similar spot is faintly visible on segment four. The quadrate spots on the ventrolateral margins of abdominal tergites two, three and four are faintly visible. Segments six, seven, and eight, and apex of five shining brown; clothed with retrose black hairs on fourth and following segments. Holotype male and allotype female, Yuma, Ariz., June 26, 1917, J. M. Aldrich, collector. (In United States Nat'l Museum, Cat. No. 51433.) Paratypes. Eight males, 1 female, same data as type; 1 male, Indio, Cal., June 6 (Dyar and Caudell) ; 1 female, Los Angeles, Cal. (collection D. W. Coquillett) ; 1 female, San Diego Co., Cal. (Coquillett collector) ; 1 male, S. E. San Bernardino Co., Cal.. June 15, 1930 (J. Wilcox collection) ; 2 males, Florence, Ariz., May 30, 1903 (collection of Acad. Nat. Sci. Phila.) ; 2 males and a broken female (not a paratype), Bill Williams Fort, Ariz., F. H. Snow (Kan. Univ.); 2 males, four females, Coachella, Cal., May 25, 1928, E. C. Van Dyke; 1 male, Phoenix, Ariz., July 17, 1932, H. Gentry (Cal. Acad. Sci.). Doctor Aldrich stated that most of his specimens were collected across the Colorado river from Yuma on the California side. 6. Apiocera caloris, n. sp. A more slender, smaller species than haruspex or aldrichi, but re- sembling them in general appearance and with different genitalia in the male. Metapleura with a conspicuous tuft of hair in front of the spiracle; dorsocaudal angle of mesopleura with three or four strong setae. All setae largely white. Length, 18 mm. Male. Ground color of body and head blackish-brown; basal points of antennae, palpi, tibiae, and apical two thirds of abdomen yellowish-brown. Pollen, all pile, and setae of body and head white; a few black setae on tarsi and apex of tibiae. Abdominal segments two, three, and four with central spots bare of pollen and hence appearing brownish or blackish. These spots on segments three and Painter: The Family Apioceratidae 195 four occupy most of the dorsum of the segments. On segment two the spot is of the same shape as on this segment of haruspex (4, fig. 6) except that the anterior border of white pollen is wider and the posterior border is not interrupted in the center. Genitalia brown, white pilose. Wings hyaline, veins brown. Female. Similar to male, all pile and hairs, except the retrose hairs on abdominal segments six and seven, white. Abdomen white pollenose a large subtriangle black spot on the center of each side of segments two and three. Holotype. Male and allotype female. Tinijas Altas, Southern Arizona, 1905, W. J. McGee, collector (in U. S. N. M. Cat. No. 51434). Paratypes. Two females, same data. All of these specimens have been in liquid (alcohol?) and one female appears to be teneral. The male genitalia are very distinct, however, and the body characters are different from any other species studied. Hence the species should be recognizable even though the specimens from which the description is drawn are poorly preserved and their coloration perhaps not normal. 7. Apiocera trimaculata, n. sp. A densely white pilose, white pollenose, species with three some- what diamond-shaped, black spots on abdominal segments two, three, and four. A tuft of hair on the metapleura in front of the spiracle, another which is not intermixed with setae on the meso- pleura in front of the wings. Setae except on the tarsi white. Length, 17 mm. Male. Ground color black, palpi yellow, tarsi and genitalia brown- ish. Clothed throughout with white pollen; a faint pattern of brownish pollen on the mesonotum. Pile white, especially dense on the abdomen, front coxae, and lower part of head. The pile is partly black on the black abdominal spots. These latter are as follows: segment two, with a central diamond-shaped spot with a triangular one attached on each side by the apex to the central spot; segment three, a central diamond-shaped spot occupying most of the dorsum of the segment; segment four, a smaller diamond-shaped spot. In addition, on the ventrolateral margins of segments three and four there is an indistinct ovoid black spot; a similar one shows very faintly on segment two. Genitalia white pollenose and pilose; notch on the tip of the ninth sternite a little more pronounced than in caloris, which it resembles. 196 The University Science Bulletin Holotype. Male, San D!ego Co., Cal, collection of D. W. Coquillett. (In United States National Museum, Cat. No. 51435.) The species is most closely related to caloris, but in genitalia and other characteristics it is distinct. 8. Apiocera convergens, n. sp. A moderately pilose species with two white pollenose stripes which converge on segments five and six and show conspicuously against the black and brown abdomen of the male. Metapleura bare, mes- opleura with thinly scattered hairs, genitalia and femora mahogany brown. Setae black. Length, 18 mm. Male. Ground color black; palpi yellowish, antennae, femora and posterior part of abdomen dark brown, tibiae light brown. Pile, white, the short hairs on thoracic and abdominal dorsum, and on genitalia, black. Pollen white, with grayish-brown patterns on the thoracic and abdominal dorsum. The abdominal pattern is compli- cated. .A slender central triangle with its base on segment two, and apex on segment five, has the base of each segment brownish, the apex of each one black, and is interrupted at the extreme apices of segments two and four by a narrow white band. This central triangle is separated from two rows of conspicuous black lateral spots by a white stripe on each side. These spots are subquadrate and become progressively larger from segment two to four, and smaller from segment five to seven. There is in addition faint black- ish stripes on the extreme ventrolateral margins of the tergites. The remainder of each segment is white pollenose. Pulvilli about as long as the claws. Wings hyaline, veins brown. Female. Similar to male, but abdominal pattern apparently con- fined to subquadrate black spots on the sides of segments two, three and four, (The specimens are greasy and rubbed.) Retrose hairs on last three abdominal segments, and those of the dorsum of thorax and abdomen black. Holotype. Male and allotype female, "Cal.," C. W. Riley collec- tion. (In United States National Museum, Cat. No. 51436.) Paratypes. Male and female, same data. The male specimens were labeled Apiocera haruspex 0. S. by Coquillett. 9. Apiocera clavator, n. sp. A small species resembling bilineata and interrupta in abdominal markings, but darker, and with entirely different genitalia. Meta- Painter: The Family Apioceratidae 197 pleura bare; mesopleura with a tuft of black setae in front of the wings. Pulvilli about two thirds as long as the claws. Setae of body mostly black. Length, 16 mm. Male. Ground color black or very dark brown, palpi yellow, tibiae and tarsi light brown. (Third joint of antennae missing.) Pollen white on head, and below the level of the wings; brownish- gray on mesonotum and abdominal dorsum, but lighter in color along the apices of the segments. Pile is white on the parts that are white pollenose, black on the mesonotum, abdominal dorsum, geni- talia, tibiae, and tarsi. Setae black, except white on head, femora, and coxae. The velvety black abdominal spots on each side of the second to seventh segments are subquadrate, but somewhat convex above. The one on the fourth is the largest; they diminish rapidly in size on the segments caudal to the fourth but to a less extent cephalad. There are faint oblong spots on the ventrolateral margins of tergites two to five, brownish in color and largest on segment four. Wings hyaline, veins brown. Genitalia much shorter than in the other species. Holotype. Male, state of Colima, Mexico, L. Conradt, collector. (In United States National Museum, Cat. No. 51437.) 10. Apiocera martinorum, n. sp. A moderately pilose species with five blackish spots on each of the segments two, three and four in the male. On segment four these tend to coalesce. Metapleura and mesopleura pilose, the pile on the former frequently reduced or absent. Genitalia and tibiae brown, setae mostly black. Length, 19 mm. Male. Ground color black; palpi yellowish, tibiae, tarsi, genitalia, and posterior margins of last three segments brown. Pile white, except the short hairs and pile on the metanotum, genitalia and the black spots on the abdomen which are black. The setae on the antennae, head, and prothorax white. Pollen white with grayish pattern on the thoracic and abdominal dorsum. On the abdomen the pattern is as follows: first segment brownish with two white spots on each side; second segment brownish at base, to which the four blackish spots are more or less connected; the remainder of the segment white; third segment with the four spots separated by white pollen; on the fourth segment the inner lateral spots are broader at the apex and almost unite with the others on this margin of the segment. The last three segments white pollenose with a central 198 The University Science Bulletin brown dorsal spot which may be due to being rubbed. Pulvilli about as long as claws. Wings hyaline, veins dark brown. Female. Similar to male, but abdominal pattern consisting of a subtriangular black spot on the center of each side of segments two and three, and an indistinct spot in the same place on four. There are also indistinct stripes on the extreme ventrolateral margins of the tergites. The first two antennal segments and femora brownish, the tibiae and last three abdominal segments light brown. Most of the pile and hair on the first four abdominal segments white. Holotype. Male, Adrian, Ore., July 22, 1932. Dwylee river sand dunes, Dorothy Martin, collector. (In C. H. Martin collection.) Allotype. Female, Adrian, Ore., July 21, 1934. C. H. Martin, collector. (In C. H. Martin collection.) Paratypes. Seven pairs, 21 females, 6Q males. Adrian, Ore., July 22, 24, 25, 31, Aug. 4, 11; Roswell Bench, Parma, Idaho. Aug. 4, 5, 1934. (Chas. H. and Dorothy Martin.) In the male, segment four of the abdomen is sometimes almost entirely black; there is variation in the size and shape of the black spots. The genitalia of the male differs from that of convergens in various proportions and in the presence of a distinct notch at the apex of the ninth tergite. In both sexes there is considerable varia- tion in the distribution of the black and white setae and in the amount of pile on the metapleura. The female differs from those of both haruspex and augur by having the black spots near the center of the sides of the segments rather than at the posterior angles. The female of convergens has subquadrate, rather than subtriangu- lar spots. This species is named for the collectors — Dorothy and Charles H. Martin. 11. Apiocera beanieri, n. sp. Abdominal pattern of male similar in pattern to that of haruspex from which it differs in characters of genitalia and in the presence of a tuft of hair on the metapleura in front of the spiracle and several setae on the dorso-caudal angle of mesopleura. Length, 22 mm. Male. Ground color of body and head black, antennae, palpi, tibiae, and abdomen beyond segment four dark brown. Body and head, except genitalia and parts of dorsum, clothed with a thick mat of pollen or minute curly hair. Palpi, front, prothorax, coxae, femora, and abdomen clothed with erect thin, white hairs. Dorsum of thorax grayish and brown pollenose, forming the usual pattern. Setae of head white, of coxae mostly white, of thorax mostly black, Painter: The Family Apioceratidae 199 of femora and tibiae black. Abdomen white pollenose; bare, black spots on the dorsum forming a pattern similar to that of haruspex; segment one brownish in the center and along the posterior margin ; segment two brownish on the anterior border, the black crossband expanded broadly in the center to the posterior border and also ex- panded laterally, the remainder of the dorsum snow white; segment three mostly black, the anteriolateral and posteriolateral margins brownish ; segment four black, the anterior margin narrowly brown- ish, the posterior white. Segments five, six, and seven mostly white. A quadrate black spot on the ventrolateral margins of tergites two, three, and four. Genitalia dark brown, pile black. Wings hyaline. Female. Almost identical with aldrichi. The single female avail- able is somewhat greasy. There appears to have been a black tri- angular spot on each side of segment four as well as on segments two and three. The retrose hairs on the fourth and following abdom- inal segments are mostly white where in aldrichi they are mostly black. Holotype. Male, Cuyama Ranch, Cahfornia, July 25, 1935 (R. H. Beamer), in Snow collection. Allotype. Female, Cuyama Ranch, California, July 25, 1935 (R. H. Beamer) , in Snow collection. Paratype. Male, Cuyama Ranch, California, July 25, 1935. (Jack Beamer) , Snow collection. This species is closely related to aldrichi, but has a different ab- dominal pattern in the male and different male genitalia. The species is named in honor of the collector. Dr. R. H. Beamer. 12. Apiocera notata, n. sp. Abdominal pattern of male similar to that of haruspex, from which it differs in having the genitalia black and lacking the notch on the ninth tergite. Length, 20 mm. Male. Ground color black to very dark brown; palpi, tibiae, and tarsi lighter brown. Head and lower parts of body and femora gray pollenose, thinly white pilose, dorsum of thorax and scutellum brown pollenose, short black pilose with streaks of grayish pollen forming the usual pattern. Setae of head and four front coxae mostly white, of remainder of body and legs black. Abdomen gray and brown pollenose with black bare spots which form a pattern similar to that of haruspex. First segment brown pollenose lighter on each side, darker in center, a tuft of white pile at the anterior corners and of black pile at the posterior corners; second segment brown pollenose 200 The University Science Bulletin on the anterior margin, white on the posterior margin, separated by three connected black triangles ; third segment with two brown spots along the anterior margin, the remainder black; fourth segment black with two small white spots on the anterior margin; fifth and sixth white pollenose, seventh mostly brown pollenose on the dorsum; laterally the usual three quadrate black spots, that on the fourth joined to the black band posteriorly. Most of the pile of the abdom- inal dorsum short, black. Genitalia blackish to dark brown, pile black. Wings hyaline. Female. Almost identical with the female of haruspex. In the single specimen at hand the triangular black spots on the abdominal segments are larger than usual in haruspex and on segments three and four are confluent with the black quadrate spots on the ventro- lateral margins of the tergites. Segments seven and eight are much darker in ground color than usual with haruspex, the knob at the apex of eight is black rather than brown. Holotype. Male, Campo, Cal., August 10, 1935 (E. I. Beamer), in the Snow collection. Allotype. Female, Campo, Cal., August 10, 1935 (Jean Russell), in Snow collection. Paratypes. Two males, same locality and date (E. I. Beamer and Jean Russell, collectors) , in Snow collection. This species is related to haruspex, but the genitalia of the male is very different. Cfiiscopui b^imeri Text Figure I. Ventral aspect of genitalia and lateral aspects of apices of the ninth tergites of Apiocera beameri and notata. Lateral aspects of antennae of Rhnphiuviidas xanthos, acton and episcopus. Painter: The Family Apioceratidae 201 LITERATURE CITED 1. CoQUiLLETT, D. W. A new Rhaphiomidas from California. West American Scientist 8:84-86. 1891. 2. A new genus of Diptera allied to Rhaphiomidas. Canad. Ent. 24:314-315. 1892. 3. OsTEN Sacken, C. R. Western Diptera. U. S. Geol. Survey Ter. Bui. 3: 281-284. 1877. 4. Painter, Reginald H. A review of the genus Apiocera Westwood from North America. Ent. Soc. Amer. Ann. 25:350-356, 1 pi. 1932. 5. TowNSEND, C. H. T. Notes on the Diptera of Baja California. Cal. Acad. Sci. Proc. Series 2, 4 :593-620. 1895. 6. New and little known Diptera from the Oregon Mountains and vicinity of New Mexico. Amer. Ent. Soc. Trans. 27:159-164. 1901. 202 The University Science Bulletin PLATE VIII Ventral aspect of genitalia of Apiocera aldrichi, caloris, haruspex, convergens, clavator, trimaculata, and bilincata and ventrolateral aspect of the ninth sternite of inten-upta and bilineata. Lateral aspects of the ninth tergites of bilineata, caloris, haruspex, augur, convergens, martinorum, clavator and aldrichi. Diagrams of the dorsal abdominal pattern of convergens and martinourm. All the genitalia drawings are made at the same magnification, except that of the tip of the ninth tergite of bilineata. Painter: The Family Apioceratidae 203 PLATE VIII bilineeta 9th. s. inlerrupta Sth. s aUnclii 9th,t clavator 9Th. t. martinorum 9fh,l eonverjenj 9|^ t converfens convergena THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 13. The Morphology of the Carolina Mantis* Philip Levereadlt, Department of Entomology, University of Kansas CONTENTS PAGE Introduction 206 Part 1. The Cr.anium 207 Epicranium : General morphology 207 Sclerities and structure 20S Cranial appendages and glotta: Antennae ' 210 Mandibles 211 Maxillae 212 Labium " 213 Glotta 2U Generalizations 215 Synonymic terms 215 Plates : IX 219 X 221 Part 2. The Thoracic Skeleton : The cervical 222 The prothoracic 223 The pterothoracic 226 The appendicular a. Wings 229 b. Legs 232 Generalizations 233 Synonymic terms 235 Plates : XI 239 XII 241 XIII 2i3 XIV 245 * This study would have been impossible for me had I not had the senerous aid of Dr. G. C Crampton of the State College of Massachusetts and of Mr. R. E. Snodgrass of the U. S. Bureau of Entomology. I am indebted to Mr. J. A. C. Rehn and Mr. A. N. Caudell for identifications. To Dr. R. V. Chamberlin of the University of Utah, and to Dr. H. B. Hungerford and his stsff of the University of Kansas, I am deeply indebted for facilities for my research, and for their encouragements and criticisms. In the matter of terminology I have received iiivaluab'e aid from Professors Sterling and Grant, of the Kansas University classical languages faculty. (205) 206 The University Science Bulletin Part 3. The Abdominal Skeleton: General morphology 246 Pregenital segments 246 Genital segments 248 a. Ovipositor 248 b. Male genitalia 252 Postgenital segments 253 Generalizations 254 Synonymic terms 254 Plates : XV 257 XVI 259 Abstract: This study is a detailed morphological consideration of the skele- ton of the Carolina mantis. It is not a topographical description, but an analysis of the skeletal structures in relation to function, origin, and develop- ment. The head skeleton has been reinterpreted. Some new terms are pro- posed, and others are given clearer meanings. The discussion is supplemented by eight plates of penned drawings. INTRODUCTION MOST of the recent work in insect morphology, especially that of this country, is of a comparative nature. The purpose of such work is the drawing of tentative concepts, a purpose of primary value for it furnishes us the broad outlines for the study of insect structures. In making an illustration the draftsman does not begin with the details of one portion and progress to another; he "blocks out" the fundamental lines first, then proceeds with the grosser features of particular areas, and finally considers the details. Com- parative analyses of insect structures give us the fundamental lines, but it should be remembered that these lines are tentative. Com- parative analyses, especially of adult features only, will not give us the final composition of insect morphology. It is with this observation in mind I offer this study of the adult skeleton of Stagmomantis Carolina Joh. I have made tentative identifications and interpretations of structures, based upon the con- sideration of currently acceptable morphological studies. Whether my identifications and interpretations, especially the latter, will re- tain their present status will depend upon further study of the Carolina mantis. I have begun this morphological study with the adult skeleton simply because we know more about the adult skeleton than of any of the other insect structures. Again, a knowledge of Levereault: The Carolina Mantis 207 the ultimate of such a basic system as the skeleton will aid in inter- preting the ontogeny, and the ontogenetic morphology of the Caro- lina mantis is the goal of my research. It will be noticed that I have not conformed closely to the use of terms and interpretations more or less generally adopted by the older morphologists. For the past few years I have become more and more certain that entomology is encumbered with many bad terms and interpretations, and fortunately, many can be dismissed readily because they are quite unnecessary. But recently I have presented publicly reasons for my objections to current usage, and since I shall convince no one of the validity of my objections if I conform to such usage, I must apply my convictions to my own research. I hope my aim for simplification and reasonableness does not produce too painful an effect upon the reader, but if it does, I suggest the reader turn to the lists of synonyms, at the end of each part of this paper, and write his preferences over mine. PART I— THE CRANIUM Epicranium All of the skeletal features comprising what is usually called "the head" are features of the cranium. These features are separable into two groups, the epicranial and the gnathocranial. The epicranium includes the prestomal flap (the upper lip) and the morphologically dorsal cuticular areas of the somites fused to form the insect head, whereas the gnathocranium includes the poststomal cranial append- ages and the ventral cuticular areas of the postoral cranial somites. The prestomal flap, or labrum, of the Carolina mantis is well de- veloped, though not strongly sclerotized. It extends from the sub- frontal sclerite (Sfr, figures 1 and 14) to the stomal or oral mem- brane. It has one main sclerite, the labrite (Lbt), a protecting and supporting plate to the inner labral membrane (Lb m, figures 4 and 14). The lower portion of the labral membrane has distinct sensory areas, a velvety median area and two lateral "peg" areas. Above these sensory "pegs" are two V-shaped sclerites, the so-called "tormae." The remainder of the epicranium, the sclerotized dorsa of the cephalic somites, is strongly developed along the transverse axis, giving the Carolina mantis a somewhat intelligent appearance. The large size of the compound eyes adds to the keen appearance, but these organs contribute more to the predatory effectiveness of the insect than to its psychical abilities. Movements from all directions 208 The University Science Bulletin can be caught by the eyes, even from the rear, as may be seen from figure 4. The corneas (E) of the compound eyes are separated from the median portion of the epicranium by cuticular infoldings which form the ocular ridges (or, figure 5). The major infolding of the epicranium is the submarginal infold- ing (see figure 3). This feature has not been recognized, heretofore, as a single, essentially continuous inflection, but as a number of separate infoldings; the subfrontal (sfr s, figure 3) or "epistomal," the subgenal (sge s) or "pleurostomal," the suboccipital (soc s) or "hypostomal," and the postoccipital (pocs). The Carolina mantis does not show to best advantage the continuous submarginal nature of these infoldings for the anterior "tentorial" pits are turned up partially and the subgenal infoldings are not evident. If we allow for these special modifications, though, we can see the essential con- tinuity by deriving the diagrammatic figure 3 from the actual speci- men shown in figure 5. The continuous nature of this epicranial feature is more obvious in the more generalized of orthopteroids, such as the cricket. I regard the submarginal inflection as a feature of first rank of the insect epicranium. It forms the chief supports and points of articulation. In addition to the submarginal infolding the Carolina mantis epicranium has the temporal (tms, figure 1) and the occipital (occ s, figure 4) infoldings. The temporals run into the occipital, and since the latter lacks the supraforaminal portion (shown by the dotted line in figure 4) the temporal and occipital infoldings appear to be a bilaterally paired feature rather than two separate features. All of these infoldings appear as two types of surficial features, externally as lines or "sutures," and internally as ridges. In using the term suture in reference to the epicranium the student should not confuse the external lines of the infoldings with the moulting line, the epicranial suture (eps, figure 1). This suture is a line of cuticular weakness, not the result of a cuticular infolding. In the Carolina mantis the epicranial suture extends from the occipital sclerite to the lateral ocelli (oc, figure 1). Its arms are not homologous to those of more generalized insects, but are morpho- logically posterior. In the generalized cranium the stem of the epicranial suture extends forward to the median ocellus, then branches about that organ into the "frontal" sutures, which extend to the anterior "tentorial" pits mesad of the antennal bases. These sutures, moulting lines and infolding lines, mark off areas of the epicranium. Some students see segmental significance in cer- Levereault: The Carolina Mantis 209 tain ones of these areas. I am skeptical of such interpretations for these sclerites are the results of a highly developed secondary con- dition. The generalized epicranium appears to be a sclerotized product of the fused dorsa of the cephalic somites. Cuticular in- flections, with their additional thickening and sclerotization, are features of support of this skull, and are features for the needs peculiar to the functions of the generalized insect skull. From this point of view the search for segmental definitions in the adult epi- cranium appears to be one entirely for the optimist. To me it seems more reasonable that the epicranial sclerites are more or less per- sistent areas because the specializations arise from a common funda- mental plan. The struts and braces may be altered variously, and consequently alter the sclerites, but they are essentially the same struts and braces. From this point of view the Carolina mantis epicranium is sub- divided into five sets of sclerites, single and paired, by the inflections and moulting line. On either side of the epicranial suture are the epicranial sclerites (Epc, figure 1). Laterad of these are the tem- poral sclerites (Tmp). Below the arms of the epicranial suture is the frontal sclerite (Frn), and laterad of the frontal area are the genal sclerites (Ge). In the Carolina mantis the preocular and hypocular* portions of the genal sclerites are greatly reduced, whereas the posterior portions (see figure 4) are rather large. The main sclerite of the posterior aspect of the Carolina mantis epi- cranium is the occipital (Occ). Some of these sclerites, the frontal, genal, and occipital, are sub- divided by the submarginal infolding; thus, from the frontal sclerite is separated the subfrontal (Sfr, figure 1) or ''clypeus," from the genal the subgenal (Sge — not definite in this mantid) or "pleuro- stoma"; from the occipital the suboccipital (Soc, figure 4) or "hypo- stoma," and the postoccipital (Poc). These terms and interpreta- tions will not be found in textbooks, but they should hardly be dismissed for such a reason. I have approached the interpretation of the Carolina mantis epicranium from a new point of view, and if it is worth while to reinterpret, it is also worth while to correct our terminology. Medical men no longer explain diseases in terms of biles and humors; with new concepts have come new terms. * The nrefixes sub and hypo have been used interchangeably, as their definitions permit ; but, for the sake of clarity I propose we limit sub for subdivisions of features, and hypo for features ventral in position. 14—6037 210 The University Science Bulletin There remains one more structure of the epicranium to discuss, the so-called "tentorium" or "endoskeleton." I am pleased to find that Dr. N. S. R. Maloeuf, of Cornell University, refuses to recog- nize any "endoskeleton" in insects. Now there are at least two with the same view. However, if we are to clean house we may as well be thorough, and if there is to be no endoskeleton I wish there would be no "tentorium." This structure (see figures 5 and 6) resembles more closely a brace than a tent; consequently, I believe transtrum, the Latin for a cross brace, preferable to tentorium. The transtrum of the Carolina mantis consists of two pairs of invaginations and one pair of weak evaginations. The anterior pair of invaginations forms the anterior transtral arms (At a). Usually these invaginations originate between the subfrontal and subgenal ridges, but in this peculiar insect they are turned up partially. From the anterior arms the weak, flexible dorsal arms (Dt a) arise and extend to the antennal ridges. The posterior por- tions of the anterior arms are partly fused to form the transtral body (BdT). This body unites with the posterior invaginations (Pta), which originate between the suboccipital and postoccipital ridges (see figure 16). The Cranial Appendages and Glotta The Antennae: These appendages of the Carolina mantis are simple and filiform. They consist of three segments, a basal (I of figure 2), an intermediate (II), and a distal (Flgm) which is sub- divided into a comparatively large number of subsegments. The appendage is an evagination of the cranial cuticula, and, therefore, is not inserted upon the epicranium. The sclerotized cuticula around the antennal base is submarginally infolded, and forms a finger- like process, the antennifer (antf), upon which the basal antennal segment pivots. The basal membrane permits considerable freedom of movement. The intermediate segment (II) is a plug-like segment which ar- ticulates with two points of the distal rim of the basal segment. No muscles extend from the third segment to the second, but if cleared antennae are examined under the compound microscope it can be seen that there is a very definite demarkation between the second and third segments, and that the "segments" of the flagellum are merely subdivisions of a superficial nature. The Gnathal Appendages: The antennae are preoral append- ages, and possibly not fundamentally metameric limbs. The gnathal Levereault: The Carolina Mantis 211 appendages are postoral and are unquestionably metameric limbs; what we know of their structure and ontogeny indicates they are fundamentally the limbs of the three postoral cranial somites. For the morphological interpretations of the gnathal appendages I follow Snodgrass' theories. The anterior gnathal appendages are the mandibles. These ap- pendages of the Carolina mantis are of the generalized orthopteroid type. They are rather conical in form (figures 7, 8, 10), and are heavily sclerotized. Figure 8 illustrates the manner of articulation of the right mandible with the epicranium. The anterior articula- tory point is the cup-shaped socket a, into which the knob a' fits. This knob is a sclerotized thickening of the subfrontal margin. The posterior articulatory point of the mandible is b, a rather large knob which fits into the socket b', a feature formed of the suboccipital ridge. About the points of articulation the connecting membranous cuticula is very narrow, not permitting much separation of the points of articulation. Between the anterior and posterior articulatory points, on the outer surface of the mandible, an infolding produces the internal shelf shown in figure 8, c. The purpose of this shelf is not apparent. The extensor muscle is attached to the tendon et, a cuticular exten- sion from the membrane just above the sclerotized edge of the man- dible. The flexor tendon, ft, is an elaborate cuticular extension of the membrane just above the median mandibular margin. The lower portion of this tendon is somewhat sclerotized. Immediately anterior to the base of the flexor tendon is a slender sclerite (3) to which a mandibular muscle is attached. The median end of this sclerite is in contact with the crescenteric sclerite (2) of the "hypopharynx." From the point of view of function the mesal surface of the man- dible is the most interesting. Figure 10 shows the structural features for mastication. Distally, on the right mandible, there are two rig- idly sclerotized processes (in p) which function as incisor teeth. These processes are sharp, and if their presence is kept in mind the finding of bits of even carabid beetle plates in the rectum of the mantis is not so astonishing. From the anterior incisor process a knifelike ridge extends halfway up the mandible. At this mid-point the ridge continues across the mesal surface, forming a sharp molar ridge. The Carolina mantis does not chew its victims too finely, and the form of the molar ridge explains this fact. The mandibles of the Carolina mantis are not bilaterally symmet- rical in respect to their masticatory features. The anterior portion 212 The University Science Bulletin of the molar ridge of the left mandible fits above the same structure of the right mandible, giving the apposition necessary for mastica- tion. In addition, the left mandible has three fully developed incisor processes rather than two. Posterior to the mandibles are the second gnathal appendages, the maxillae. From figures 9 and 13 it can be seen that the Carolina mantis maxilla is generalized and orthopteroid. The palp (Pip) is simple and five-segmented, with the basal segment in direct contact with the stipes (Stp). This plate is the largest of the appendage. It has a longitudinal inflection which furnishes a ridge (str, figure 13) for the attachment of a large muscle. Above the stipes is the cardo (Cd), the functional base of the Carolina mantis maxilla. An inflection (cds) traverses the center of the cardo, forming an internal ridge for muscular attachment and a process (cd p, figure 16) to which is attached the extensor tendon et (figure 13). The manner of articulation is shown in figure 16. This is a drawing of the right cardo from an angular view; the labels of the features should orient the reader. There are no definite basal maxillary articulatory points compara- ble to those of the mandible. The medial portion of the cardo is dis- tended into a cone-shaped process (d) which is in close contact with the suboccipital sclerite. The connecting membranous cuticula is- very narrow at this point of contact, and permits but little separa- tion. Other parts of the connecting membrane are ample, permitting much freedom of movement. The process of the cardinal inflection (cd p, figure 16) passes be- neath the suboccipital sclerite, and to its inner end is attached the extensor tendon. A tug by the extensor muscle will rotate and lift the distal portion of the cardo, and press point d against the suboc- cipital sclerite. The antagonists to the extensor pull against the car- dinal and stipital ridges, and in this function, too, point d is pressed against the suboccipital sclerite. In considering the endite lobes of the maxilla I feel again in a revising mood. The "galea" (Plx, figures 9 and 13) is a thumblike- endite, yet, galea is Latin for helmet or headpiece. How much more descriptive is the term pollex. "Lacinia" is Latin for the lappet of a garment. It might be we could stretch things a bit for the sake of priority and consider this lobe (Fcp) a lappet. However, it was with some difficulty that I first learned to identify this endite lobe as the "maxillary lacinia." If it had been named the maxillary forceps the Levereault: The Carolina Mantis 213 structure and its term would have had a more accurate meaning. I believe that students will find maxillary pollices and forcipes more significant. The anterior surface of the maxilla (figure 13) is not so interest- ing, morphologically, as the posterior surface. The mesal margin of the forceps is studded with stout spines and hairs. The pollex has scattered sensory hairs about its distal surface. Near the base of the forceps a stout tendon (ft) extends from the membranous cuticula. The third postoral appendages of the cranium are partly united to form a lower lip, the labium (figure 12). Figure 14 shows that both the prestomal flap and the labium are appropriately termed lips. The labium is in close contact with the epicranium at the lower ends of the postoccipital sclerite. There are no definite articulatory points ; the points e are separated from the postoccipital sclerite by narrow strips of membranous cuticula. At all other points the la- bium is separated from the remainder of the cranium by relatively large membranous areas. The points e are probably homologous to d of the maxillary cardo, but since we are not sure of the large basal labial plate (Mt) it is better to call it the mental sclerite. My use of mentum is different from that of the older students. They called this plate the "sub- mentum," and in insects with this chin-plate divided, they called the subdivision the "mentum," and the main part of the plate "submen- tum"! If we cannot assume safely that the mentum is the fused cardines of the labium we can at least be certain of the labial stipital areas (Stp, figure 12). The partly united stipites bear the simple three- segmented palps. The labial stipites of the Carolina mantis have been identified as the mentum. As in the maxillae the labial stipites each bear two endite lobes. We have admitted of the serial homologies of these lobes in the max- illae and labium, yet we have retained the old terms "glossae" and "paraglossae" for the labial lobes. On the first point, these endite lobes are hardly tongues in either form or function. Then, if these lobes are homologous why not recognize the homologies in our terms? I propose that we strive for simplicity and reasonableness in this complex enough study, and that the labial "glossa" become the labial forceps (Fcp, figure 12) and the "paraglossa" become the la- bial pollex (Plx). The labium of the Carolina mantis is not uniformly sclerotized. 214 The University Science Bulletin This fact may be made apparent by staining.* The basal portion of the mental plate is more heavily sclerotized than the lower or distal. There are two peculiar unpigmented spots (f) in the basal portion. The purpose of these spots is obscure; no muscles are attached to their inner surfaces. The stipites are not uniformly sclerotized. The median areas, out- lined with broken lines in figure 12, and the lateral pieces, are heav- ier. On the center of the basal margin of the stipites are two trian- gular areas, which are well sclerotized; they are bases for muscles. There are no sclerites on the anterior surface of the labium (figure 12 shows the posterior surface only). The Glotta: Between the gnathal appendages lies the venter of the cranium, the so-called "hypopharynx." Some years ago Snod- grass called our attention to the fact that what we formerly con- sidered as pharynx was not pharyngeal, but preoral. In figure 14 the mouth of the Carolina mantis is shown by Mth. The space be- tween the labrum, mandibles, and ''hypopharynx" is the preoral space (Prsp). We do not refute this observation of Snodgrass, but, regardless, we keep calling the inner surface of the labrum the "epi- pharynx," and the tongue-like cranial venter "hypopharynx." Ob- viously we can do without the former, and we should change the latter. The tongue, for it functions as a tongue in the generalized insect, has been called the lingua, but there is considerable prejudice against that Latin word. Possibly glotta, the Greek for tongue, will prove satisfactory. The glotta of the Carolina mantis is shown from the anterior sur- face in figure 11, and from the lateral aspect in relation to the la- brum and labium in figure 14. A velvety down of sensory hairs covers most of the anterior surface, which is relatively thick though not strongly sclerotized. The posterior surface is more sclerotized, but is thinner. At the base of the glotta are three pairs of sclerites; 1 is called the "suspensorial bar," ^ is a slender crescent, a base for muscular attachment, and 5 is a loop extending from 2 to the vestig- ial mandibular adductor. The salivary glands empty into a common duct (Sid, figure 14), which leads into a salivary pocket (SI p, figures 14 and 15), between the glotta and anterior surface of the labium. The saliva flows out of this pocket down the median groove of the posterior glottal sur- * If the skeleton is cleared of tissues with KOH anrl rinsed in distilled water, the sclerites can be stained by placing the skeleton in hot iron alum for a few minutes. The stain does not fade perceptibly for some time if the specimens are kept in eighty percent alcohol. Levereault: The Carolina Mantis 215 face, and collects in drops at the distal tip of the glotta. Figure 15 shows the details of the salivary pocket from an angular anterior view. The anterior surface is cut and folded over to show its rela- tionship to the posterior wall. The sides of this pocket are sup- ported by the sclerites 5 and 6. Generalizations The gnathocranium of the Carolina mantis is conservatively or- thopteroid and generalized. The maxillae, mandibles, and glotta differ little from Snodgrass' concepts of the generalized pterygotan forms. The salivary pocket differs from the generalized form in that it is quite large, and the labium is relatively lightly sclerotized. The epicranium has not retained the primitive form so well. The dorsal-most portion has been greatly elongated transversely, and flattened longitudinally. Temporal ridges have arisen, most prob- ably in response to the strains of the huge mandibular flexor muscles. An odd condition is the union of the temporal and occipital inflec- tions, with the suppression of the supraforaminal portion of the occipital. The subgenal inflections have been suppressed, possible because of the reduced condition below those enormous compound eyes. Synonymic Terms for the Cranium New terms Older equivalents Antennal segment I scape Antennal segment II pedicel Antennal segment flagellum remainder of "segments" Cranium h'-ad, head capsule Epicranial sclerites vertex, parietals Epicranial suture coronal plus frontal sutures Epicranium dorsal part of cranium plus prestomal flap Forceps in maxilla — lacinia ; in labium = glossa Glotta Iiypopharynx Ginathocranium Iiypopharynx plus gnathal appendages Labral membrane epipharynx Labrite "labrum" Labrum whole of prestomal flap Mental sclerite submentum plus mentum Occipital sclerite occiput plus postgenae Pollex in maxilla ;=. galea ; in labium = paraglossa Preoral space mouth cavity, cibarium Prestomal flap cuticula of prostomium Subfrontal infolding epistomal ridge Subfrontal sclerite clypeus Subgenal infolding pleurostomal ridge Subgenal sclerite pi urostoma, mandibularia Suboccipital infolding hypostomal ridge Suboccipital sclerite hypostoma Transtrum tentorium 216 The University Science Bulletin REFERENCES Crampton, G. C, 1923. Phylogenetic comparisons of maxillae throughout the orders of insects. Journ. N. Y. Ent. Soc. 31, pp. 77-106. 1925. External anatomy of the head and abdomen of the roach. Psyche, 32, pp. 105-200. 1932. Phylogenetic study of the head capsule. Bull. Brooklyn Ent. Soc, 27, pp. 19-55. SxoDGB.\ss, R. E. 1928. The morphology and evolution of the insect head. Smiths. Misc. Collect., 81, No. 3. 1931. The evolution of the insect head. Smiths. Rpt. 1931, pp. 443-489. 1935. Principles of insect morphology. McGraw-Hill Co., New York- City. W.\LKER E. M. 1933. Comparisons of the head of Grylloblatta with those of other orthopteroids. Anns. Ent. Soc. Am., 26, pp. 309-344. T.A.BLE OF AbBREVI.^TIOXS Mn m r A anterior in direction a a anterior articulatory points of mandible Mt . antf antennifer Mth At a anterior transtral arm oc . . at p anterior transtral pit Occ . b, b' posterior articulatory oc f . points of mandible ^^^ j. Bd T body of transtrum q ,. c inner shelf of mandible p Cd cardo Phy cd p cardinal process pj^ cd s cardinal suture Poc d basal point of cardo p^g g Dt a dorsal transtral arm pj. gp E coi-nea of compound eye pj ^ e ba.sal point of labium pt p Epc epicranial sclerite R, . . eps epicranial suture Sfr . et extensor tendon sfr s Fcp forceps Sge . Flgm flagellum of antenna sge s Frn frontal sclerite SI d ft flexor tendon SI p . Ge genal sclerite Soc Gl glotta soc r in p incisor process soc s L ; left Stp Lab labium st r . Lbm lahnun St s . Lb m labral membrane Tmp LI m labial membrane tms . mandible molar ridge mental sclerite mouth ocellus occipital sclerite occipital foramen occipital ridge ocular or corneal ridge posterior pharynx pollex postoccipital sclerite postoccipital suture preoral space posterior transtral arm posterior transtral pit right subfrontal sclerite subfrontal suture subgenal sclerite subgenal suture salivary duct salivary pocket suboccipital sclerite suboccipital ridge suboccipital suture stipes stipital ridge slii)ital suture temporal sclerite temporal suture 218 The University Science Bulletin EXPLANATION OF FIGURES PLATE IX Figure 1. Anterior view of epicranium. 2. Ventral surface of antennal base. 3. Diagram of submarginal inflection, derived from figure 5. 4. Posterior view of epicranium. 5. Angular view of epicranium with upper anterior portion cut off. 6. Dorsal view of transtrum. Le\'ereault: The Carolina Mantis 219 PLATE IX Tmp Epc FIG. 2 Tmp Epc FIG. 6 220 The University Science Bulletin PLATE X FlOURE 7. Anterior surface of right mandible. 8. Articulation of mandible with epicranium from mesal plane. 9. Posterior surface of right maxilla. 10. Mesal surface of right mandible. 11. Anterior surface of glotta. 12. Posterior surface of labium. 13. Anterior surface of right maxilla. 14. Lateral surface of glotta, with labrum and labium in section. 15. Angular view of salivary pocket. 16. Articulation of right cardo. Levereault: The Carolina Mantis PLATE X 221 Ji cd Occ Cd Urn FIG. 13 FIG. 15 Soc FIG. 14 FIG. 16 222 The University Science Bulletin PART 2— THE THORACIC SKELETON The Cervical The neck of the Carolina mantis is an elaborate mechanism inter- posed between the head and prothorax. It is relatively elaborate be- cause few insects have this body region so highly developed for the movement of the head. The Carolina mantis need not be envious of the mammal for its flexible neck. Figures 7 and 10 (Plate XI) indicate the external form of this mantid's neck skeleton ; figure 7 is a ventral view, and 10 is an angu- lar dorsal view from the left side. An interesting oddity of this neck skeleton is its suspended position. Usually the orthopteroid neck is inserted within the prothorax, but in this insect it lies outside and below the prothorax. ' The cervical sclerites are large, and rather numerous. A pair of dorsal sclerites (d Cv, figure 10) intervenes between the postoccipital sclerite of the cranium and the dorsal longitudinal muscles passing through the prothorax. These muscles are attached to the tendons 2, which are cuticular extensions from the cervical sclerites. Laterad of these sclerites are the lightly sclerotized areas 3. The function of these areas is not obvious for no muscles are attached to them, and, their degree of sclerotization is too slight for protection to this vul- nerable region. The lateral cervical sclerites (1 Cv^, 1 Cv,), the important features from the standpoint of function, are large and well sclerotized plates. The anterior knoblike points of the first lateral cervicals lie adja- cent to the indentations of the postoccipital sclerite below the label Poc of figure 4, plate IX. These points are the fulcra upon which the cranium nods up and down. Posterior to the first lateral cervical sclerites are the second lateral cervicals (ICv,). These sclerites are folded ventrally and toward each other, instead of being in line with the first lateral cervicals, the generalized condition. Two small sclerites (4), probably parts of the posterior margins of the second lateral cervicals, intervene be- tween the latter and the cervical rim of the prothorax. The dorsal points of these sclerites are adjacent to the juncture of the protho- racic "episterna" and precoxal bridges. Between the lateral cervi- cals are the riblike ventral cervical sclerites (v Cv^, v Cv,, figure 7). These cervicals are well sclerotized, but they are not connected with the musculature. Levereault: The Carolina Mantis 223 In his textbook Snodgrass said, "The true morphology of the cer- vix is still obscure — ." This observation should not be disregarded, for nothing is quite so valueless as speculation upon morphological identities from scant anatomical knowledge. I am deferring the con- sideration of the morphology of the Carolina mantis neck skeleton until I study its embryology. The Prothoracic As a general characteristic the Orthoptera have the prothorax rather distinct from the other two thoracic segments, and the man- tids are of special interest in this respect, for certain species have this somite extremely elongated. In the Carolina mantis the pro- thorax is not only a distinct body segment, but because of its length it is also an important functional unit, a lever for this insect's pe- culiar predatory features. The skeleton of the Carolina mantis prothorax is formed mostly of the tergum and sternum (see figure 1, plate XI). The pleural structures contribute to the coxal and supracoxal regions, but they are in a reduced condition in contrast to the greatly elongated ter- gum and sternum. Such a long cuticular unit needs strengthening features, especially if it is to be subjected to the actions of the neck, foreleg and posterior prothoracic muscles of the Carolina mantis. These strengthening features are in the tergum. This sclerotic plate, the tergum, forms a hemicylindrical arch over the relatively flat sternum. In addition to the intrinsic transverse strength of this form, a transverse cuticular infolding (1 of figure 2) braces the tergal arch against the strains of the large foreleg muscles. The hemicylindrical arch has also an intrinsic longitudinal rigidity, but this intrinsic resistance is intensified by three features, a median carina and the two peculiarly folded lateral edges of the tergum. The median carina is postcoxal in position. It is sharply folded, thickened, and well sclerotized. The lateral tergal edges function as V-shaped longitudinal beams. The sternum is extensively fused to the arching tergum, particu- larly in the postcoxal region of the prothorax. In the precoxal por- tion the pleural sclerites intervene between the sternum and tergum. Figure 6 shows the relationship of tergum, pleurites, and sternum in the precoxal region, in transverse aspect. From the posterior edges of figures 2 and 3 the relationship of tergum to sternum in the post- coxal region may be seen. 224 The University Science Bulletin The sternites of the generalized pterygotan thoracic sternum arc well developed in the prothoracic sternum of the Carolina mantis, though they are not sharply demarked. The anterior-most, the basisternite (Bst, figures 1, 3, 4) , is long and relatively narrow. It is separable from the precoxal portions of the pleura only by slight cuticular inflections (shown by broken lines in figure 3). The pos- terior margin of the basisternite is continuous with the "episterna." Between the coxae is a splinterlike sclerite, which may be part of the basisternite, or of the furcasternite. Possibly the study of the development of the Carolina mantis will indicate the identity, but for the present time this identity is of little importance. The impor- tant feature of this sternum is the elongated furcasternite (Fst, fig- ure 1), the sternite which contributes most to the ventral surface of the prothorax. This sclerite of the Carolina mantis has been identi- fied as the spinasternite, but from figure 4 it can be seen that the furcal invaginations (fa) arise within the anterior margin of this sclerite. Posterior to these invaginations the furcasternite is defi- nitely a single sclerite to the points of the separation of the tergum from the sternum (see figure 1). At these points of separation there are two peculiar notches in the edges of the sternum (5, figure 9). In the African Cameroon mantid Theopompella heterochroa Gerst., the sternum, at points similar to those of the Carolina mantis, is deeply notched. In addition, in the Cameroon mantid, there is a definite line between the notches, about which the sclerotization is weaker than in either the furcasternite or spinasternite. I regard this condition as intermediate between the fused condition of the Carolina mantis, and the primitive state in which these sclerites were separate. Posterior to the furcasternite is the spinasternite (Spst, figures 1, 9). Fundamentally the spinasternite is a ventral intersegmental element. In the case of the Carolina mantis it seems that a portion of the prothoracic sternum proper has become sclerotized and con- tinuous with the intersegmental element. This element is restricted, most probably, to the area about the spinal process (Sp p, figure 9). The pleura of the prothorax are peculiarly modified and over- lapped, but they are essentially modifications of the generalized pterygotan thoracic pleurum. However, before considering the mod- ifications of this mantid's prothoracic pleura, my definition and interpretation of the generalized pleurum should be made clear. First, I prefer to Latinize the Greek pleuron, for Greek diminutives are not so easily adapted to our terminology. On the second point, Levereault: The Carolina Mantis 225 I prefer to think of the pleurum as the sclerotizations between the tergum and sternum, not as the whole side or lateral wall of the tho- racic segment. The lateral wall of the thoracic segment of pterygotans is braced by two ribs. As to the origins of these ribs I favor Snodgrass' sub- coxal theory. This theory appears overelaborate at first glance, but if the student study the fundamental morphology of the insect abdo- men the subcoxal origin of pleurites becomes a most plausible one. The basal subdivision of the primary coxa is the pleural arch of the adult pterygotan, the important support of the thoracic lateral wall. The second subdivision is the pleurellar, or "trochantinal" arch. These two subdivisions are not separate plates in the mature pterygotan, but are partially fused to form the functional thoracic pleurum. In most Orthoptera an anterior portion of the pleurellar arch remains relatively independent of the pleural arch. This inde- pendent sclerite is called the "trochantin," a term of doubtful sig- nificance in this case. Since it is a part of the pleurellum, I think it should be called the pleurellite. The united pleurellum and pleural arch are infolded dorsoven- trally. This infolding is usually thickened and strongly sclerotized to form the brace between tergum and sternum. It divides the pleural arch into an anterior prepleurite ("episternum") and a pos- terior postpleurite ("epimeron"). Often the ventral portion of the prepleurite is separated off, and termed the precoxal bridge, or "latero-sternite." The prothoracic pleurum of the Carolina mantis is an odd modifi- cation of this basic plan. The prepleurite (Prpl, figures 2, 4, 5, 8) is relatively well developed, though by no means comparably to the tergum or sternum. It extends from the pleural infolding (pi r, fig- ure 8) to the cervical rim of the prothorax. Its lower, or ventral, portion is marked off by slight inflections as a precoxal bridge (Prcb). The postpleurite is not readily identified in this segment. From figure 4 we can see that this pleurite (Pspl) which is generally pos- terior to the pleural infolding, is, in the Carolina mantis, reflected forward over the prepleurite. However, as shown in figure 5, the postpleurite is reduced, and not clearly demarked from the tergum. In this mantid's prothorax the pleurellar arch is remarkably de- veloped. It is an important part of the functional pleurum, as may be seen from figures 5 and 8. In figure 5 the anterior portion of the 15—6037 226 The University Science Bulletin pleurellum (Pll) is shown detached from the prepleurite. A better idea of the relationship of the pleurellum to the pleural arch may be gained from figure 8. The line labeled 6 is the line of separation of the two pleural plates. It continues across the pleural apophysis (pi a), and if the lower part is pulled away from the upper portion of the pleurum, the longer part of the pleural apophysis remains with the pleurellum, leaving the pleurum as on the right side of the speci- men drawn in figure 3. The anterior portion of the prothoracic pleurellum is divided into two pieces, a basal piece which is in close contact with the prepleu- rite, comparable to the ventral portions of the pterothoracic pre- pleurites proper (not the precoxal bridges), and a distal piece, the movable pleurellite. The tip of this pleurellite forms an accessory coxal articulatory point. The main coxal articulatory point is the knobbed ventral tip of the pleural ridge (cxf, figures 5 and 8). Posterior to the pleural infold- ing the pleurellum is oddly folded and distorted. It is divided into two sclerites, the knobbed 7 of figure 8, and the folded 8. The latter sclerite is a base for muscular attachment. Usually this part of the pleurellum is fused to the postpleurite, and, the condition of the Carolina mantis is appropriately labeled peculiar. In addition to the peculiar modifications of the pleurites them- selves, the prothoracic pleura of the Carolina mantis are overlapped by the precoxal portion of the tergum. Figure 6 is a transverse sec- tion of the precoxal region, illustrating the relative development of tergum, pleura, and sternum. The Pterothoracic The mesothoracic and metathoracic segments of the Carolina mantis form a body unit, the pterothorax. It is a unit as distinct as the prothorax, and one as important, for it is the locomotor center of this insect's body. The prothoracic legs are so modified for grasping they are of little use for walking. This function is left to the ptero- thoracic legs. These two pairs of legs are practically the sole means of locomotion of this insect, since the wings and their associated structures are hardly adequate for effective flight, especially those of the female. Although the pterothoracic segments are strongly modified to form a running mechanism, the basic plan of the skeleton is essentially that of the properly functioning, generalized pterothorax. The terga (figures 12 and 15) are large, relatively well-sclerotized plates. Levereault: The Carolina Mantis 227 Their anterior portions are crossed by the dorsal intersegmental in- foldings. Externally these infoldings are indicated by the sutures isg s of figure 12, and internally by the ridges isg r of figure 15. Anterior to these intersegmental inflections of the terga are the pretergite? (Prt). Though these sclerites are integral parts of the pterothoracic terga they are not of the same somite origin as the terga with which they are associated, but of the preceding somites. The pretergite of the mesothoracic tergum is a part of the protho- rax.* In the Carolina mantis the pretergite of the mesothoracic ter- gum is a single sclerite, and an odd one. The lateral edges are sub- marginally infolded, and from the anterior ends of these infoldings two stoutly sclerotized, tubular invaginations (1 of figures 12 and 15) extend into the body cavity. The pretergite of the metathoracic tergum is not single, but of four pieces. The posterior-most piece forms a border to the tergum. Its lateral edges are not infolded, and the cup-shaped processes 2 (figure 15) are the counterparts of the tubular processes of the mesothoracic pretergite. Anterior to the border of the metathoracic tergum are the other three pretergal sclerites in the intertergal membrane (figure 15) . The posterior margins of the pterothoracic terga of this mantid are demarked from the central shields by slight inflections. These scle- rites, the posttergites (Pst), are folded, the morphologically posterior margins lying beneath the inflections. This manner of folding forms protecting plates to the intertergal membranes. The shield-shaped portion of each tergum is crossed by two V-shaped inflections. The anterior inflection of the mesothoracic tergum forms a pair of short ridges (ats r, figure 15), and the poste- rior inflection a pair of long, closely approximated ridges (pssr). These two pairs of ridges are struts supporting the tergum longitu- dinally. They do not make the tergum absolutely rigid, for there is a line of flexure between them. This flexure is a part of the flexing line of the tergum, a line between the two-wing fulcra. The meta- thoracic tergum has the same inflections as the mesothoracic, though they are not so strongly developed. The surface areas of the terga these inflections mark off are used frequently by taxonomists for characters. The taxonomists have a set of terms for these areas, which, no doubt, will prove to be per- sistent names, but which nevertheless are not suitable from the mor- phologist's point of view. A part of a tergum is a tergite, so ''prescu- » For more details of this condition the reader should study Chapter IV of Snodgrass' "Principles of Insect Morphology." 228 The University Science Bulletin turn," ^'scutum," and "scutellum" hardly fit into our interpretations and terminology. The pterothoracic terga are structurally adequate for flight, but the pleura are not; they are better suited for the running type of insect. These two pairs of lateral supports are poorly consolidated in the Carolina mantis, for nearly half of the lateral pterothoracic walls are membranous. In addition to this poor degree of consolida- tion, the dorsal portions of the pleura are inclined anteriorly. The pleural infoldings form the ridges pi r (figure 14), the braces separating the terga and sterna. The lower portions of these infold- ings are extended into large flat apophyses (pi a). These projections are in close contact with the furcal apophyses (f a, figure 16). The ventral-most points of the pleural infoldings are knobbed to form the main coxal articulatory points (cxf, figure 14) ; and, the dorsal- most points form the wing fulcra, the alifers (alf). Anterior to the pleural infoldings are the prepleurites (Prpl, fig- ures 11 and 14), the largest pleurites of this insect. The anterior ventral portions of these pterothoracic sclerites are not separated definitely as precoxal bridges (Pre b) as in the prothorax. From each pterothoracic prepleurite are separated two epipleurites (1 Ep, 2 Ep) . Posterior to the pleural infoldings are the long, narrow post- pleurites (Pspl). From each of these pleurites one epipleurite (3 Ep) is separated. Of the pleurellum only the movable pleurellite (Pit) is distinct from the functional pleurum. Points 3 and 4 (fig- ures 11 and 14) of the pleurellites are accessory coxal articulatory points. In contrast to the pleura the pterothoracic sterna are relatively well consolidated though they do permit lateral flexure at the inter- sternal region. If their posterior portions are disregarded, the form of these sterna is rather suggestive. The basisternites (Est) are shields not only in function, but decidedly so in shape. As indicated in figure 14, the anterior border of the mesothoracic basisternite is folded back, forming a protection for the preceding intersternal membrane. An interesting feature of the mesothoracic basisternite is the V-shaped inflection (5, figures 13 and 16). This inflection gives the sternite a rigidity not possible from the degree of scleroti- zation alone. In the metathoracic basisternite a vaguely V-shaped carina (6) corresponds in function to the mesothoracic sternal in- flection. Posterior to the basisternites are the furcasternites (Fst). In the mesothoracic sternum the furcasternite is a continuation of the basi- Levereault: The Carolina Mantis 229 sternital inflection. It forms a semicircular arch, through the sagit- tal plane, between the pterothoracic sternal shields. At the apex of the arch a pair of invaginations forms two stout processes, the furcal apophyses (fa, figure 16). Posterior to these apophyses the fur- ca'sternite is flattened, and terminates with its spadelike margin in close contact with the metathoracic sternum. The furcasternite of the metathoracic sternum is similar to that of the mesothoracic in form, but differs in the greater development of its anterior portion and in the reduction of its posterior portion. Of the thoracic ventral intersegmental elements there are two well developed spinal processes in the Carolina mantis, and possibly there is a vestigial third. The anterior spinal process has been dis- cussed in the section on the prothorax. The posterior spinal process (Sp p of figure 16) is fused to the anterior border of the metathoracic basisternite. It is usually claimed that no intersegmental sclerite occurs between the metathoracic and first abdominal sterna, but, in the Carolina mantis a suspicious looking, buUbous sclerite lies just posterior to the metathoracic furcasternite. It may be merely a part of the furcasternite, but the temptation to call it a vestigial ventral intersegmental sclerite is a strong one. Possibly the study of the development of this insect will furnish something more substantial than a temptation. The Appendicular The wings are relatively large, but they are not well modified for effective flight structures, especially those of the female. The male's wings are long and have sufficient surface to maintain his slender body in the air, though he is capable of only a fluttering manner of flight, but the female's wings are relatively shorter than the male's in spite of her heavier body. As in all winged Orthoptera, the mesothoracic wings of the Caro- lina mantis differ from the metathoracic in that they are narrower and more sclerotized. Figure 18 represents the right forewing from its dorsal surface. The costa (C) extends along the entire anterior margin, forming a stiff rib for this margin. Subcosta (Sc) is usually unbranched in the Carolina mantis, though some forewings have an irregularly branched subcosta. Radius (R) is a strongly sclerotized vein, usually two-branched, but often subradius (the "radial sector") is irregularly branched. Medius (M) is weakly sclerotized at its base, but distally it is better developed. Its anterior branch is as often dichotomously branched as it is simple; as a matter of 230 The University Science Bulletin observation, an individual may have one wing with M^^^ &s a single piece, and the other Mi and M2 separated. The first ''cubitus" (1 Cu) branches closely to its base, with the anterior section (1 Cua) branching dichotomously twice more. The second "cubitus" (2 Cu) is a single vein, which is sometimes fused with 1 Cub, and sometimes terminates without fusing, but it rarely touches the margin. The vannal veins (IV, 2V, 3V) have a common base which articulates with the vannal axillary. Most of the specimens examined had only three vannals, but occasionally a weak fourth vein may be seen. In the wing region between the vannal area and the tergum, the jugal region (Ju r), are usually four weak veins. The forewings fold back and cover the hind wings, overlapping each other but slightly. Beneath the protecting forewings lie the flexible, fanlike metathoracic wings. These appendages do not over- lap each other, but lie side by side, folding first in the region between 1 CUb and 2 Cu, and beneath these remigial areas lie the vannal areas folded in the manner of plaited fans. The longitudinal veins of the metathoracic wing differ from their counterparts of the forewings (see figure 19) in that medius is un- branched, there are usually nine vannals, with the first vannal (IV) many branched, and with the jugal region much reduced. In both pairs of wings numerous cross veins, some X-shaped, some Y-shaped, but most of them simple, intervene between the longitudinal veins. A close study of the wing bases is essential to an accurate inter- pretation of the venation, for there is a definite relationship, at least in the more generalized insects, between the longitudinal veins and the wing-base sclerites, the axillaries. Because of this general re- lationship a change in the specific naming of the axillaries is here proposed: Proposed terms Snodgrass' terms Crampton's terms Subcostal axillary 1st axillary Notale Radial axillary 2d axillary Proximediale Medial axillaries Median plates Intermediale Distimediale Vannal axillary 3d axillary Basanale 4th axillary Adanale The subcostal axillary is generally in close contact with the dorsal base of the subcostal vein, the radial axillary with the dorsal base of the radius, the medial axillaries, though variable, are in close contact with the medius or cubitus, or both, whereas the vannal axillary articulates with the bases of at least the anterior vannal Levereault: The Carolina Mantis 231 veins. The fourth axillary, or adanale, is not often present, and when it is present it is obviously a secondary constriction of the tergal edge. This general relationship is not a new observation, so the proposed changes of names should not appear too radical, and they may be beneficial to the elementary student. Apparently every generalization must have its exceptions, and in this instance the mesothoracic subcostal axillary (Sc ax, figure 17) articulates with the base of radius instead of the subcostal base. This part of the Carolina mantis has been rather modified, for the subcostal base is detached from the subcosta and has fused with the radial base (Rb). However, the identity of the subcostal axillary can be affirmed in spite of the modification by its position as inter- mediary between the tergum and the radial axillary (Rax). The latter sclerite can alwaj^s be identified, since it rests upon the alifer of the pleurum and is usually in close contact with the radial base. In the mesothorax the subcostal base (Sc b), the medial axillaries (M ax) , and the dorsal sclerite of the radial axillary are all con- tinuous with the radial base, separable from one another only by slight inflections. The vannal axillary (V ax) is continuous with the posterior medial axillary, though it is strongly deflected from the surface level of the medial axillary. In the metathorax (see figure 24) the condition of the subcostal and vannal axillaries is more gen- eralized than that of their counterparts in the mesothorax, but the subcostal base has merged with the radial base, with which the dorsal sclerite of the radial axillary and the medial axillaries are also continuous. From the posterior medial axillary a prong extends into the otherwise separated vannal axillary. It should be remembered that there are two cuticular surfaces close together at the wing base, the dorsal surface (dwm, figures 20 and 24), which is a lateral extension of the thoracic dorsum; and the ventral surface (vwm) , an extension of the dorsolateral thoracic wall. The "tegula" (tg), the subcostal and medial axillaries, and the vein bases are distinctly sclerotizations of the dorsal surface. The other sclerites are more or less closely associated with sclerites of the ventral surface. In figure 20, the ventral surface of the mesothoracic wing base, the subcostal has a ventral base (Sc b). A smaller sclerite lies basad of this subcostal base. Of more sig- nificance, however, is the ventral sclerite of the radial axillary. This axillary is a compound sclerotization, for in cross-section it can be seen that the dorsal and ventral sclerites of the axillary are partially fused. Internally the prong of the ventral sclerite is 232 The University Science Bulletin thickened, and shaped like the engineer's bearing cap, and fits over the alifer of the pleurum. The vannal axillaiy also has a ventral sclerite, though it does not fuse with the dorsal part. For locomotion and the procuring of food the Carolina mantis depends almost entirely upon its legs. For the procuring of food the prothoracic legs are more effective than are the pterothoracic for locomotion. The mantis is an awkward insect at any gait. The pterothoracic legs, however, are the more generalized (see figure 27). The coxa (Cx) of these legs is more or less conical in form. It has a basal infolding (be r figure 29) to reinforce the basal rim. Two points on the anterior portion of this basal rim are especially thickened for articulation; point a (figure 29) is the chief articulatory point which abuts against the coxifer of the pleural infolding, and point b is the accessory point which is adjacent to the tip of the pleurellite. The two articulations, the primary and accessory, differ struc- turally. Figure 21 is a semidiagrammatic representation of the primary articulation. To the left of the figure is external to the skeleton, and to the right is internal. The Y-shaped piece is the coxifer (cxf ) in section, and the folded piece below the coxifer is the basal rim of the prothoracic coxa, in section. The intervening mem- brane is labelled m. From this drawing it may be seen that the thickened point of the basal rim abuts against the coxifer externally. The accessory articulation is more simple. The tip of the pleurellite and point b (figure 29) are in close contact, separated only by a narrow strip of thickened membranous cuticula. Distally the coxa articulates with the base of the femorella (the "trochanter"), at two points. Structurally these articulations are different from the pleuro-coxal. From the disticoxal rim two sclero- tized prongs point inwardly at right angles to the coxal wall, and lie adjacently to two sclerotized prongs of the basifemorellar rim which point outwardly at right angles to the femorellar wall, so that with the slender strips of the intervening membranous cuticula, simple hinges are formed. It should be noted that these points are at right angles to the pleuro-coxal points. The femorella (Fml: "trochanter" is the only misfit in the termi- nology of the insect leg, and it can be discarded easily) is a small segment functioning as a plumber's "elbow-joint" for the Carolina mantis leg. It is very closely joined to the femur, thereby prevented from moving appreciably upon the femur, but oddly enough there Levereault: The Carolina Mantis 233 are rudimentary articulatory points of the prong-hinge type between the two segments, and these points are at right angles to the coxo- femorellar articulations. The pterothoracic femur (Fm) is a long slender tube, articulated to the tibia (Tb) by two prong-hinges. To the tibia the tarsus (Tar) is articulated by a single point. The tarsus is rather gen- eralized, being subdivided into five distinct tarsal subsegments (I-V). The sixth subdivision is represented by the claws (cl) and the ■unguitractor mechanism (figures 22 and 25). The pulvillus is not present in the Carolina mantis tarsus. The prothoracic legs differ strikingly from the pterothoracic (see figure 23). They move parallel to the sagittal plane rather than at an angle to it, as do the pterothoracic legs. The coxa (Cx) is greatly elongated and cylindrical in form. Three more or less ser- rated carinae strengthen this long cylinder. The coxofemorellar articulations of the prothoracic legs are in the same plane as the pleuro-coxal, an odd condition. Ordinarily these two sets of articu- lations are at right angles to each other. The femorella (Fml) is rather large, no doubt necessarily, for the femur (Fm) is a very large leg segment. Both femur and tibia (Tb) are studded with strongly sclerotized spines. Whatever prey is not daunted by these spines is effectively held by the hornlike elongation of the distal portion of the tibia. The tarsus of these legs has remained rather generalized. Its long slender form dangling from such a highly modified leg appears, to say the least, incon- gruous. Generalizations The thoracic skeleton of the Carolina mantis is the most highly modified part of its body. Two striking modifications characterize this skeletal region, one which might be called a predatory modi- fication, and the other, less obvious but very fundamental, a modi- fication of the generalized plan into a running type of thoracic mechanism. This latter modification is evident in the pterothoracic segments, especially in the form of the pleura. These supporting ribs are reduced, poorly consolidated, and inclined anteriorly. It is not necessary to be familiar with the pterothoracic musculature to surmise that the pterothoracic skelton functions only as a base for the legs of these segments, though, if the ungainliness of the Carolina mantis is considered, it seems that such a strong modification should produce better results. 234 The University Science Bulletin The pterothoracic appendages are rather generalized. The legs are very simple and slender, and it may be that the slender form of the femora is a negative factor in the development of speed in this insect. Though very generalized, the pterothoracic legs have not retained the puvillus of the tarsus. The wings are typically orthopteran, the forewings being more sclerotized than the hind wings, and the latter being large, fanlike structures. Another orthopteran characteristic of these wings is the great degree of in dividual variation in the venation. For all practical purposes the wings of the Carolina mantis are merely accessory appendages, in- herited but of little value. The insect depends upon its four slender pterothoracic legs for locomotion. The most interesting modification of the Carolina mantis is the predatory modification, the elongation of the prothoracic skeleton, and the efficient-looking, grasping forelegs. The tergum and ster- num are greatly elongated and fused together. The pleura are greatly reduced and overlapped and are, functionally, but minor parts of the skeletal unit. I call this modification a predatory one, for it has changed a generally modest body segment into a long lever which permits full play to those grasping legs. In connection with this predatory modification, the complex neck structures of this mantid should not be overlooked. The neck permits the Carolina mantis to move its head as freely as can most mammals. Such a flexible mechanism increases the predatory effectiveness of those sharp, tearing mandibles. The best-known characteristic of the mantid is the peculiar form of the prothoracic legs. These legs are held in a prayerful attitude; however, if popular observation were not so superficial, and the sharp look of the mantid's face noticed, the insect would have been named, originally, less deceptively the preying mantis. These pro- thoracic legs are so highly modified for the grasping function they are quite useless for other functions. They move parallel to the sagittal plane, rather than at an angle to the plane. The tibial, femoral, and coxal articulations are all strongly developed, indicat- ing something of the strength of these legs. An odd and interesting specialization is the hornlike extension of the tibia, a process which prevents even the most armored victim from scjuirming free of the vice formed by the tibia and femur. Levereault: The Carolina Mantis 235 Synonymic Terms for the Thoracic Skeleton yew terms Older equivalents Alifer pleural wing process Anterior scutal infolding prescutal infolding Dorsal intersegmental infolding antecostal infolding Epipleurites: 1 Ep, 2 Ep. basalares (of episternum) 3 Ep subalare (of epimeron) Femorella trochanter Furcal apophysis furcal arm Jugal region posterior part of anal wing region Lateral wall (of segment) pleiiron Medius media Pleural apophysis pleural arm Pleurellite trochantin Pleurelliim trochantinal arch Pleurite a sclerite of pleuron Pleurum trochantinal plus pleural arches Posterior scutal infolding scutellar infolding Postpleurite epimeron Posttergite posterior reduplication Prepleurite episternum Pretergite acrotergite, precosta Sternum all of ventral sclerotization Subradius radial sector Tarsal subsegment tarsal segment Tergum notum Vannal (vein) anal REFERENCES Crampton, G. C. 1909. A contribution to the comparative morphology of the thoracic sclerit«s. Proc. Acad. Nat. Sci. Phila., Jan., 1909, pp. 3-54. 1914. Notes on the thoracic sclerites of insects. Ent. News, 25, pp. 15-25. 1914. Ground plan of a typical thoracic segment in winged insects. Zool. Anz., Bd. XLIV, 2, pp. 56, 67. 1915. Suggestions for the standardization of technical terms in ento- mology. Anns. Ent. Soc. Amer., 8, pp. 74-78. 1916. Phylogenetic origin and nature of wings according to the para- noial theory. Jour. N. Y. Ent. Soc, 24, pp. 1-39. 1917. Nature of the neck region in insects. Anns. Ent. Soc. Amer., 10, pp. 187-197. 1917. Phylogenetic study of the lateral head, neck, and prothoracic regions. Ent. News, 28, pp. 398-412. 1918. Thoracic sclerites of immature pterygotans. Proc. Ent. Soc. Wash., 20, pp. 39-62. 1923. Preliminary notes on the terminology applied to parts of an in- sect's leg. Can. Ent., 55, pp. 126-132. 1926. Comparison of neck and prothoracic sclerites throughout the orders. Trans. Amer. Ent. Soc, 52, pp. 199-248. 1927. Thoracic sclerites of Periplaneta americana. Psyche, 34, pp. 60-72. Levereault, P. 1935. The insect tarsus. Kansas Univ. Sci. Bull., 22, pp. 521-525. 236 The University Science Bulletin Maloeuf, N. S. R. 1935. The role of muscular contraction in the production of configurations in the insect skeleton. Jour. Morph., 58, pp. 41-85. Martin, J. F. 1916. Thoracic and cervical sclerites of insects. Anns. Ent. Sos. Amer., 9, pp. 35-83. Snodgrass, R. E. 1909. The thorax of insects. Proc. U. S. Nat. Mus., 36, pp. 511-595. 1927. Morphology and mechanism of the insect thorax. Smiths, Misc. Coll., 81, No. 3. 1929. Thoracic mechanism of a grasshopper. No. 2. 1929. 1935. Smiths. Misc. Coll., 82, How insects fly. Smiths, Rept., 1929, pp. 383-421. Principles of insect morphology. McGraw-Hill Co., New York City. Table of Abbreviations a main coxal articulatory Pre b precoxal bridge point alf alifer ats r anterior-scutal ridge atss anterior-scutal suture b accessory coxal artic- ulatary point be r basicoxal ridge Bst basisternite C costa C b costal base Cer f cervical foramen cl claws Cx coxa ex f coxal foramen cxf coxifer d Cv dorsal cervical sclerite dwn dorsal wing membrane fa f ureal apophysis Fm femur Fml femorella Fst furcasternite isg r intersegmental ridge isg s intersegmental suture Jur jugal region iCvi— Cv2.. lateral cervical sclerite M mediua m membrane M ax medial axillary PI iileurum pi a i^leural apophysis PU pleurcUum pi r pleural ridge pi s pleural suture Pit ■ pleurellite Prpl prepleurite Prt pretergite Pspl postpleurite pss r posterior-scutal ridge pss s posterior-scutal suture Pst posttergite R radius R ax radial axillary Rb radial base Rs subradius Sc subcosta Sc ax subcostal axillary Scb , subcostal base Spp , spinal process Spr spiracle Spst spinasternite St . sternum T . tergimi Tar . tarsus Tb . tibia tg . "tegula" tg r . tergal ridge ugt . unguitractor V ax . vannal axillary V Cvi — CV2 . ventral cervical sclerite vwin ... ventral wing membrane lAbSt ... . first abdominal sternum 1 Cu, 2 Cu . . first and second "cubitus" lEp, 2Ep. . epipleurites of prepleu- rite 3 Ep . epipleurite of postpleu- rite IV, 2V, etc ., vannal veins 238 The University Science Bulletin PLATE XI FlOURK 1. Angular lateral view of prothoracic skeleton. 2. Inner surfaces of pleura and tergum in supracoxal region. 3. External surface of anterior portion of sternum. 4. Inner surfaces of pleura and anterior portion of sternum. 5. Outer surfaces of left pleurum, shown outstretched. 6. Transverse section of precoxal region. 7. Ventral view of neck. 8. Inner surfaces of right pleurum and tergum. 9. Inner surface of posterior portion of sternum. 10. Angular dorsal view of neck. Levereault: The Carolina Mantis 239 PLATE XI Fig. I Spst Fig. 2 Fig. 3 Fig. 4 dCv -ICv. Fig. 8 Fig. 9 Fig. 10 240 The University Science Bulletin PLATE XII Figure 11. IJuter surfaces of mesothoracic and metathoracic pleura. 12. Outer surfaces of mesothoracic and metathoracic terga. 13. Outer surfaces of mesothoracic and metathoracic sterna. Levereault: The Carolina Mantis 241 PLATE XII 2 Ep alf 2 Ep alf 3Ep iEp\ Fig. II Prcb Fig. 13 16—6037 242 The University Science Bulletin PLATE XIII Figure 14. Inner surfaces of mesothoracic and metathoracic pleura. 15. Inner surfaces of mesothoracic and metathoracic terga. 16. Inner surfaces of mesothoracic and metathoracic sterna. Levereault: The Carolina Mantis 243 PLATE XIII 2 Ep alf 3 £p FiG. 14 FIG. 15 244 The University Science Bulletin PLATE XIV Figure 17. Dorsal view of right forewing base. 18. Dorsal surface of right forewing. 19. Dorsal surface of right hind wing. 20. Ventral view of right forewing base. 21. Section through main articulation of prothoracic coxa and pleurum. 22. Dorsal surface of tip of tarsus. 23. Lateral view of left prothoracic leg. 24. Dorsal view of right hind wing base. 25. Ventral view of tip of tarsus. 26. Ventral view of right hind wing base. 27. Anterolateral view of left middle leg. 28. Inner basal structure of left foreleg. 29. Inner basal structure of right middle leg. Levereault: The Carolina Mantis 245 Cb PLATE XIV iCu M R Sc C Fig. 26 Fig. 29 246 The University Science Bulletin PART 3— THE ABDOMINAL SKELETON In the discussion of the thorax I made the statement that if the morphology of the insect abdomen is understood, the subcoxal origin of the pleurites appears to be a plausible theory of origin. Not only the probable origin of the pleurites becomes apprehensible through an understanding of the morphology of the insect abdomen, but the fundamental plan of the sclerotization and metamerism of the entire insect trunk becomes evident. It was after his studies of insect abdomens that Snodgrass perceived most clearly the fundamental plan of the insect skeleton. Every teacher of entomology should read the first seven pages of his "Morphology of the Insect Abdomen; Part II. The Genital Ducts and the Ovipositor."* These few pages will do more to unsettle the antediluvian concepts of insect structure still prevalent, in spite of Crampton's and Snodgrass' researches of the past twenty years, than any amount of analysis and criticism from a younger student. Pregenital It is in studying the abdomen that it becomes evident the skeleton of an insect somite has two primary sclerotizations, a tergum and a sternum. Between these two plates are the lateral, or podial regions, from which the paired metameric limbs are developed. Whatever sclerotizations may occur in these podial regions are essentially sclerotizations of the limbs, of either a primary or secondary nature. However, the sclerotization of an adult abdominal segment of such an insect as the Carolina mantis could not be interpreted as having simply a tergum and sternum, with the metameric limbs secondarily suppressed and represented by the membranes between the dorsal and ventral plates. In the first place, the abdominal skelton has the secondary condition of segmentation, so both terga and sterna are not of a simple intrasomitic composition. In the second place, most of the abdominal limbs are present, though they do not appear as limbs. In the second to the seventh (inclusive) segments of this mantid (see figure 1, Plate XV) the limbs are flattened, and are integral parts of the functional abdominal sterna. How much of the functional sternum is true or primary sternum, and how much limb base, it is not possible to determine from the adult condition. In Thysanura most of the functional sternum is morphologically limb base, the true sternum appearing as a small medial triangle between the two laterally disposed limb bases. * Smiths. Misc. Coll., 89, No. 8. Levereault: The Carolina Mantis 247 Since the terga and sterna are the two primary sclerotizations, and are separated from each other by the podial regions, the insect trunk is fundamentally divided, longitudinally, into four surficial areas by two pairs of lines. According to Snodgrass the membranous infolding below the spiracles (d-11, figure 1) indicates the position of the dorsolateral line in the abdomen. The ventrolateral line is not so clearly indicated in the adult, because of the complete fusion of limb bases to the sterna. Transversely the Carolina mantis abdomen is divided into at least ten somites. After a careful study it is obvious there are eleven abdominal somites in this insect, the eleventh being a reduced seg- ment usually held retracted within the larger tenth. These somites can be grouped conveniently into three sets. The first seven are pregenital, the eighth and ninth are genital, and the tenth and eleventh are postgenital. The skeleton of the pregenital segment (second to sixth, inclusive) is more generalized than that of the other segments. The tergum (IIT-VIT, figure 1) is a broad shield to the somitic dorsum. Its structure indicates a simple expression of secondary segmentation. The intersegmental infolding (isg r, figure 11, Plate XVI) traverses the anterior margin of the tergum. The pretergite (Prt) is but a narrow border, yet quite distinct. The lateral edges of the terga of this insect contain the spiracles. These edges are made distinct from the remainder of the terga by longitudinal carinae, and they might be distinguished from the dorsal portions of the terga as paratergites (part, figure 1). The sternum, in spite of its complex composition, appears to be as simple a plate as the tergum, in the pregenital seg- ment. It has a narrow presternite, and a slight intersegmental in- folding. The lateral portions, the limb bases, are indistinguishable from the primary sternum. The relationships of these features, in the transverse plane, is shown in figure 10. The tergum (T) forms the upper half of the oval outline. Its paratergites (part) are usually bent inwardly. The sternum (St) forms the bottom half of the oval outline. Between these two plates are the lateral mem- branes. The dorsal portion of the podial area is indicated by the dorsolateral line (d-11). When the female is gravid, and the eggs are large, the membranes become exposed and the paratergites are partially straightened. Figure 1 shows the female abdomen partly distended. Just prior to oviposition the abdomen is so distended the oval outline is changed into a nearly circular one. 248 The University Science Bulletin In the gravid condition the female abdomen is distended not only transversely, but also longitudinally. Figure 11 is a sagittal section through the fourth segment. The intertergal (itgm) and the inter- sternal (istm) membranes are rather ample, and if they are un- folded, the length of the abdomen is noticeably increased. I have referred to the second to the sixth, inclusively, as the more generalized of the pregenital segments of the Carolina mantis. A second glance at figure 1 will disclose that the first and seventh segments differ from those between them. The tergum of the first segment is closely associated with the metathoracic tergum, its pre- tergite being in direct contact with the thoracic plate. The dorsal intersegmental infolding furnishes points of attachment for what re- mains of the dorsal longitudinal muscles. Another point of differ- ence in the first tergum is its lack of paratergites. As in most Orthoptera, the first abdominal sternum is greatly reduced (see figure 16, 1 AbSt). In the abdomen of the female Carolina mantis the sternum of the seventh segment is elongated and greatly enlarged to form a ventral cover for the ovipositor (see figures 1 and 3). There are some spe- cial terms for this ovipositor cover, but since some orthopterans have the eighth sternum as the ovipositor cover, it is better to discard terms that might indicate a homodynamous condition. Nothing much is gained by having such terms as "subgenital plate," "hypan- drium," etc. I do not believe any entomologist is so busy that re- ferring to the ovipositor cover as the seventh or eighth abdominal sternum, whichever it may be, is an imposition upon him, and the long designation can be abbreviated for labeling purposes. The Genital Within this seventh abdominal sternum is an ovipositor mech- anism which is the source of much controversy in regard to its morphology. One group of entomologists has interpreted the ptery- gotan ovipositor by comparing the more generalized pterygotan mechanisms with those of the Lepismatidae and Machilidae. The interpretation derived from this comparison appears creditable, and seems to explain the origins and forms of the pterygotan ovipositors. However, R. I. Nel, when he was at the Imperial College of Science and Technology, London, did some work on the development of the genital ducts and genitalia of certain orthopteran females, and he has questioned the value of comparative analyses of the adult structures. Levereault: The Carolina Mantis 249 Nel thought that the ventral collineae (the "valves," "valviilae") of Blattella are serially homologous with its dorsal collineae ("dorsal valves"), and that its medial collineae ("inner valves") are endite lobes, or apophyses of the gonopods of the ninth segment. No such apophyses, in Blattella, are developed from the gonopods of the eighth segment. This interpretation differs from Snodgrass' in that Snodgrass identifies the ventral collineae as apophyses of the gono- pods of the eighth segment, serially homologous to the medial collineae. According to Snodgrass the gonopod proper of the eighth segment is reduced to form the collineafer (the "valvifer"). This controversy arises from two opinions, one that the value of comparative anatomy is greater than ontogenetic evidence, and the other that comparative analyses may be deceptive and that onto- genetic studies must form the basis for interpretation. To quote from Snodgrass,* ". . . it is quite a different matter to prove that these parts represent true segmental appendages. Even the fact that the first rudiments of the valvulae appear in some insects on the embryo in line with the vestigal appendages on the pregenital seg- ments is not necessarily evidence that they are homodynamous with the latter, since secondary structures arising in the same relative positions as the true limbs would be very likely to assume the same form in early stages of growth. The best evidence of the origin of the ovipositor from the limb structures is furnished by the Thy- sanura." But, Nel insisted,! "It has been generally accepted up to now — that in pterygote insects the anterior ovipositor valves (the "ventral valves") of segment 8 are serially homologous with the inner ovi- positor valves of segment 9. My observations on Locustana, Cole- mania, and Blattella indicate that it needs revision. In these insects the ovipositor valves are first represented by two pairs of lobe-like outgrowths on the eighth and ninth sterna. The manner and place of origin of these rudiments leaves no doubt that they are serially homologous. Later the lobes on the ninth give rise to a pair of outgrowths on their mesal margins. The lateral structures of the ninth develop into the lateral ovipositor valves ("dorsal valves"), the mesal pair into the inner valves. No mesal outgrowths are differentiated from the lobes of the eighth sternum and they develop into the anterior ovipositor valves ("ventral valves"). * Page 48 (bottom), "The Abdominal Mechanisms of a Grasshopper," Smiths. Misc. Coll., 94, No. 6. t Page 49 (middle), "Studies on the Development of Genitalia and Gemtal Ducts m In- sects," Quart. Journ. Micr. Sci., 73 n. s., pp. 25-86. 250 The University Science Bulletin In connection with the possible relationships of the pterygotan ovipositor with that of the machilid, Nel wrote (page 50, middle), "A comparison of the origin of the ovipositor lobes in young Orthop- tera and in the young stages of the machilid Petrobius carpenteri Bagnall is instructive, and clearly shows that the anterior ovipositor valves (ventrals) of Orthoptera do not correspond morphologically with the ovipositor lobes of Machilidae but with the gonocoxites (limb-bases) of these insects." To which Snodgrass said (page 54, one third), "It may still be difficult to prove that the first valvulae (ventrals) are gonapophyses homodynamous with the second val- vulae ('inner'), and not coxal processes corresponding with the third valvulae (dorsals) ; but the identical relations of the first and second valvulae to their respective valifers in most insects, and the fact that these valvulae constitute the usual blades in the shaft of the ovi- positor, to which the third valvulae are mere ensheathing lobes, leaves little basis for questioning the apparent and generally accepted homologies of the ovipositor components." Thus, we have two sets of interpretations, and the proponent of each appears to be certain of his ground. One of them must be in error, and, though I refuse to commit myself definitely in view of the meager evidence we have on hand, I am inclined to look upon ontogenetic evidence with more favor than upon comparative analyses of adult structures. Until ontogenetic work is proven in- adequate for purposes of interpretation of insect structures, I shall interpret the ovipositor of the Carolina mantis from Nel's side of the fence. The gonopods of the eighth and ninth abdominal segments of the Carolina mantis are shown from lateral views in figures 6 and 7, and from a ventral view in figure 2. The gonopods of the eighth segment form the ventral collineae ( v Coll : derived from collineare, to direct in a straight line. Though this designation may be too specific if applied literally, we cannot use director, from directus, since it is too common. At least collinea is preferable to "valve"). These processes are oddly shaped; they are bowed at their bases. There are essentially two sclerotizations in each ventral collinea, a basal, the collineafer (collfg), and a central sclerotization which encircles the main part of the process. The tip is membranous. The gonopods of the ninth segment are more complex in structure. The main part of the gonopod forms the dorsal collinea (dColl) and the collineafer (collfg). The medial apophysis of this gonopod forms the medial collinea (m Coll, figures 2 and 7). The dorsal collinea is Levereault: The Carolina Mantis 251 better sclerotized than the ventral, but the medial collinea is en- tirely membranous. Figure 7 shows the relationship of the medial to the dorsal collinea from the mesal aspect. In this figure the medial collinea is separated from the dorsal, but the broken lines indicate points of contact. In figure 4 the relationship of these collineae, at their bases, is indicated. An odd feature is the articu- lation between the two processes. The collineafer of the dorsal collinea is a triarticulated sclerite. Its dorsal point (1, figures 6 and 5) is in contact with the distal point of the fused paratergites of the eighth and ninth terga (part, figure 6). This articulation is the fulcral point from which the ventral collinea is slipped back and forth beneath the dorsal collinea. Point 2 is the articulation permitting movement between the col- lineafer of the ventral gonopod. Point S is the articulation between the dorsal collinea and its collineafer. The internal surfaces of these structures are shown in figure 5. The terga of the genital segments are noticeably smaller than the preceding pregenital terga. The paratergites of both the eighth and ninth terga are greatly elongated, strongly sclerotized, and fused to- gether to form suspension points for the collinear apparatus. It will be noted that no spiracles are in the paratergites of the ninth tergum. The sterna of the genital segments are as reduced as the paratergites are developed. What I identify as the eighth sternum is a small triangular plate between the bases of the anterior pair of gonopods (VIII St, figures 2 and 5). The sternum of the ninth segment ap- pears to be split into two pieces, one which lies between the bases of the dorsal collineae (IXSta), and the other between the medial collineae (IXStb, figure 5). My interpretation of the eighth sternum of the female Carolina mantis probably will be questioned since that sclerite is posterior to the gonopore of this insect. However, is it not more reasonable to call the sclerite between the bases of the ventral collineae the ster- num of the eighth rather than to place that sternum morphologically anterior to the gonopore because it is said the gonopore is posterior to the eighth sternum? Nel's developmental studies, as well as those of Denny, Peytoureau, Bordas, Chopard, Wille, Vogel, and Ford, leave little doubt as to the position of the gonopore in female man- tids and blattids ; it is in the seventh intersternal membrane. Figure 8 is a sagittal scheme of the distal portion of the female abdomen of the Carolina mantis. From this drawing we may see that the seventh sternum (VII St) is a long plate divided into two 252 The University Science Bulletin regions by an infolding. The distal region may be the fused append- ages of the seventh segment, or it may be merely a secondary con- striction of the functional sternum. Above the seventh sternum is the folded and thickened intersternal membrane. The thicker por- tions of this membrane are somewhat sclerotized, and it is this sclerotization that has been identified as the eighth sternum of the Carolina mantis. The morphologically posterior part of the inter- sternal membrane is enlarged and invaginated to form a genital chamber (GC). The gonopore (Gpr) opens in the posterior part of the floor of this genital chamber. In the dorsal wall of the genital chamber is the eighth sternum (VIII St). Posterior to this sclerite is the opening to the spermathecal duct (Spm d). Between the two sclerites of the ninth sternum (IX St) empties the products of the colleterial glands (Colgl). As to the morphology of the spermatheca Nel pointed out that it is not always homologous, that "the term spermatheca has a 'func- tional' rather than a 'morphological' significance." However, he does concur that the spermatheca of generalized Orthoptera is an ectodermal invagination of the eighth abdominal somite. Nel identifies the colleterial glands as invaginations of the ninth sternum. At least about these two invaginations there is no marked contro- versy. Singh-Pruthi thought the spermatheca an invagination of the ninth segment, primitively, but he worked with beetles and Nel has placed many questions in his way. If it is wise to leave the interpretation of the morphology of the neck until after I study the development of the Carolina mantis, it would be wiser still to refrain from interpreting the male genitalia of this insect. The interpretation of these organs will require a careful study of the development of not only this mantid, but com- parative ontogenetic studies of its relatives and of lower forms. Figure 9, Plate XVI, is an angular lateral view, from the left side, of the distal abdominal structures of the Carolina mantis male. The eighth pair of spiracles (Spr) are contained in the paratergites of the eighth tergum. The sternum of this segment, in the male, is a large, well-sclerotized plate. Posterior to the eighth is the greatly elongated ninth sternum. The form of this sternum indicates its composition; the basal portion is the primary sternum, and the distal portion is of the fused limb bases of this segment. The telopodites of these limb bases are retained as the styles (Sty), in the adult. Above this scoop-shaped ninth sternum are the male genitalia. Though they appear to be highly complicated, these organs can be Levereault: The Carolina Mantis 253 separated into three definite groups, or lobes. These lobes are drawn separated from each other in figures 12 and 16. The medial lobe (ML) is ventral in position, in relation to the other two. It carries the gonopore (Gpr, figure 12) in its membranous dorsal surface. Its ventral surface is well sclerotized and the distal point of this sclero- tization is drawn into a recurved hook. The right lobe (R L) is the dorsal most in position; it is the least developed of the lobes. The outer edge is supported by a peaked sclerite, and ventrally there is a transverse band. In contrast to this simple lobe is the complicated lobe to the left and above the medial lobe (LL). Its ventral sclerites terminate in two curved hooks. A curious membranous evagination, resembling vaguely a bird head, projects from the posterior portion of the dorsal membrane of the left lobe. In the young male mantid three lobes emerge from the posterior region of the ninth venter. Two of these lobes are lateral in position, and the third is ventral in position and carries the gonopore. This ventral lobe becomes the medial of the adult male, the left lateral forms the complicated left lobe, and the right lateral lobe becomes the simple right lobe of the adult male. The two laterally disposed lobes have been identified as the apophyses of the gonopods of the ninth segment. Snodgrass, however, finds nothing in these two lobes to suggest they have any relation to the gonopod bases. The last word concerning the orthopteran male genitalia has not been said. The postgenital segments appear to be simple, yet they are not so simple that the interpretation of their morphology is free from controversy. Concerning the tenth segment there is little question. Some would have the cerci as the appendages of this somite, but in embryological studies it has been commonly observed that the limb rudiments of the tenth abdominal somite are suppressed before hatching, and that the cerci are the metameric appendages of the eleventh segment. The association of the cerci with the tenth tergum is a secondary condition. The sternum of the tenth segment is not apparent in the Carolina mentis. There is considerable doubt about the morphology of the eleventh segment. In this mantid the eleventh segment is not part of the tenth, but is a distinct structure held retracted within the tenth. From figure 14 the relation of the eleventh to the tenth segment may be seen from the sagittal plane. The tergum of the eleventh (XI T, figure 15) is a small, triangular, weakly sclerotized plate lying beneath the stronger tenth tergum. On either side of the anus (A) are the two paraprocts (Papt). These sclerites have been called 254 The University Science Bulletin the split halves of the eleventh sternum, and, some have maintained that the anterior portions are parts of the tenth sternum and the posterior parts are of the eleventh sternum. Crampton (1929) considered the paraprocts to be the limb bases of the cerci. I rather suspect this is true. In the Carolina mantis, at least, it seems that the functional bases of the cerci (see figures 13 and 15) are not the morphological bases. The articulation of the cerci to the tenth tergum is definitely a secondary association. It is hoped that a careful study of the ontogeny of the mantis, with the aid of newer technique methods, will indicate the solution of many of these fundamental questions about the Carolina mantis. Generalizations The abdominal skeleton of the Carolina mantis is very generalized for that of a pterygotan insect. Most of the terga have the edges distinct as paratergites. In the eighth and ninth terga of the female the paratergites are strongly developed to form suspension points for the collinear apparatus. The sterna are simple in appearance; the first sternum is c^uite reduced, but distinct. In the female the seventh sternum is enlarged to form an ovipositor cover. In the male the form of the ninth sternum indicates its composition of primary sternum and metameric appendages. The male genitalia are peculiarly twisted lobes, three in number, and undoubtedly the derivatives of the three lobes of the immature male mantid. The eleventh segment is reduced, but distinct; it is retracted within the tenth. The cerci are relatively long, hairy, and annulated. The female ovipositor is a simple pterygotan mechanism. The female gonopore opens upon the seventh intersternal membrane, and the male gonopore upon the medial lobe of the genitalia. Synonymic Terms for the Abdominal Skeleton New terms Older equivalents Collineafer valvifer Dorsal coUinea dorsal, or third valve Genital segment eiglith or ninth abdominal segment Intersegniental infolding antecosta Lati ral membrane pleural membrane Medial collinea inner, or second valve Podial area limb-base area Postgenital segment tenth or eleventh abdominal segment Pregenital segment any of the first seven abdominal segments Presternite acrosternite Pretergite acrotergite Sternum sternite Tergum tergite Ventral collinea ventral, or first valve Levereault: The Carolina Mantis 255 REFERENCES Crampton, G. C. 1917. Phylogenetic study of the terminal abdominal seg- ments of some female apter>'gotans and lower pterygotans. Jour. N. Y Ent. Soc, 25, pp. 225-237. 1918. Phylogenetic study of terminal abdominal structures of male Apterygotans, Orthoptera, etc. Bull. Brooklyn Ent. Soc, 13, pp. 49-68. 1920. Terminal abdominal structures of Mastotermes darwinensis. Trans. Ent. Soc. London, 1920, pp. 137-145. 1920. Remarks on basic plan of terminal abdominal structures of males of winged insects. Can. Ent., 52, pp. 178-183. 1923. A comparison of the terminal abdominal structures of the female Mastotermes darwinensis with Periplaneta americana. Bull. Brooklyn Ent. Soc, 18, pp. 85-93. 1925. External anatomy of the head and abdomen of Periplaneta ameri- raria. Psyche, 32, pp. 195-220. 1927. The abdominal structures of Grylloblatta. Pan-Pacif. Ent., 3, pp. 115-135. 1929. Terminal abdominal structures of female insects, compared throughout the orders. Jour. N. Y. Ent. Soc, 37, pp. 453-496. Ford, Norma. 1923. A comparative study of the abdominal musculature of orthopteroid insects. Trans. Roy. Canad. Inst., 14, pp. 207-319. Nel, R. I. 1929. Studies on the development of genitalia and genital ducts in in.sects. Quart. Jour. Micr. Sci., 73, pp. 25-85. Walker, E. M. 1919. Terminal abdominal structures of orthopteroid insects. Anns. Ent. Soc. Amer., 12, pp. 267-316. 1922. Terminal structures of orthopteroid insects. Anns. Ent. Soc. Amer., 15, pp. 1-76. Wheeler, W. M. 1893. A contribution to insect embryology. Jour. Morph., 8, pp. 1-160. Table of Abbrevl\tions A anus L L left lobe art articulation m Coll medial collinea Cer ccrcus ML medial lobe Col d colleterial duct part paratergite collf coUineafer Parpt paraproct c Ovd common oviduct R L right lobe d Col! dorsal collinea Spm d spermathecal duct d 1 1 dorsolateral line Spm p spermathecal pore Ej d ejaculatory duct Spmt spermatheca G C genital chamber Spr spiracle Gpr gonopore St sternum ist m intersternal membrane Sty style itg m intertergal membrane T tergum Lb limb-base v Col ventral collinea 2,i)C) The University Science Bulletin PLATE XV FlfiDRE 1. Lateral view of female abdomen. 2. Ventral view of ovipositor. 3. Angular lateral view of terminal segments of female. 4. Base of right gonopod of ninth segment. 5. Base of ovipositor from inner view. 6. Lateral view of ovipositor. 7. Mesal surfaces of gonopod of ninth segment. 8. Sagittal section through terminal segments of female. Levereal'lt: The Carolina Mantis PLATE XV iT ffT" 'fJ '^^ ,. '^^ 7rT 257 cOvd FIG. 7 Fig. 8 17—6037 258 The University Science Bulletin PLATE XVI Figure 9. Angular lateral view of male terminal segments. 10. Cross-section of pregenital segment. 11. Sagittal section of pregenital segment. 12. Dorsal view of male genitalia, separated. 13. Base of left cercus. 14. Sagittal section through postgenital segments. 15. Ventral view of eleventh segment. 16. Ventral view of male genitalia separated. Levereault: The Carolina IVIantis PLATE XVI 259 wlT izT Fig. 9 ^~%5fi^'i\ " ^ ^ part Fig. 14 ^"^^ Parpt rCer Fig. 10 Fig. II Cer Fig. 13 Fig. 16 Fig. 15 THE UNIVEESITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 14. Species of Erythroneura of the Comes Group (Homoptera-Cicadellidae) R. H. BEAMER, Department of Entomology, University of Kansas Abstract : North American species of Erythroneura (grape leaf hoppers) of the Comes group are reviewed. Seventy species and varieties are included. One was unrecognizable: {E. octonotata Walsh), seventeen are placed in synonymy: E. vitis var. flava Robinson and E. vitis var. veniista McA. {=E. v'tlis (Harris) n. syn.) ; E. fusco-flava Beamer {=E. bistrata McA. n. syn.) ; E. ruhranotum Rob. (= bistrata var. stricta McA. n. syn.); E. tricincta var. rubravitla Rob. {:= diva McA. n. syn.); E. tricincta var. erasa McA. {=E. calycidu McA. n. syn.); E. breakeyi Johnson (^E. pontifex McA. n. syn.); E. mallochi McA. i=E. rosa Rob. n. syn.); E. portea Rob. {^=E. reflecta McA. n. syn.) ; E. atlenuata Johnston {=z E. nuclata McA. n. syn.) ; E. tvdella Rob. and E. scripta Rjob. {^E. delicata var. accepta McA. n. syn.); E. omata Osb. {=E. corni Rob. n. syn.); E. comes var. snffusa McA. {=:E. bidens McA. n. syn.) ; E. irrorata Rob. (== E. rubra (Gill.) n. syn.) ; E. Cherokee Rob. and nigroscuta Johnson (=E. compta McA. n. syn.); and the following are described as new: E. prima, E. jestiva, E. tacita, E. ziczac var. walshi. All existing types have been examined. Drawings of male genitalia, keys to groups, species and varieties are included. Types of the new species are in the Snow collection, University of Kan.sas, Lawrence, Kan. INTRODUCTION IN this paper I have attempted to bring together all of the North American species of Erythroneura of the Comes group as de- fined by Wm. Robinson, University of Kansas Sci. BuL, vol. XVI, March, 1926. Seventy species and varieties are treated. One of this number was not recognizable, eighteen are placed as synonyms and four are described as new. All existing types have been studied, most of the males dissected and the internal genitalia figured. It is this group to which most of our economic grape leaf hoppers (261) 262 The University Science Bulletin belong. The cultivated grape is without doubt the greatest sufferer, with the majority of the species and varieties listed feeding on this host. Virginia creeper, Boston ivy, red bud, willow and various species of dogwood are also hosts to some of the forms. GROUPS OF ERYTHRONEURA The genus Erythroneura was divided into six groups by W. L. McAtee, Trans. Amer. Ent. Soc, vol. XLVI, 1920. Wm. Robinson in Univ. of Kan. Sci. Bui., vol. XVI, No. 3, March, 1926, further revised the genus into five groups, giving them names instead of numbers, as did McAtee. After the dissection of many thousands of specimens I propose a further revision into four groups. It was found that the Scutelleris and Maculata groups of Robinson vary from one to the other and the genitalia were more or less of one type. The Scutelleris group is therefore dropped. It must always be borne in mind when using wing veins as diagnostic characters that they are variable. In general, however, they follow the char- acters pointed out in the keys. Key to Groups of Erythroneura 1. M-Cu cross vein present 2 M-Cu cross vein usually absent, media and cubitus forming a more or less continuous line 3 2. Base of cell M* curved ; two posteriorly diverging vittae usually present on vertex and pronotum obliqua group Base of cell M4 angulate ; vertex and pronotum not marked as above ; cross veins usually white vulnerata group 3. Base of cell M4 oblique; apex of cell R3 without black spot. .. .maculata group Base of cell M4 square; apex of cell R3 with black spot or cloud. . . .comes group Key to the Species and Varieties of Erythroneura of the Comes Group 1. Color markings of dorsum one or more transverse bands or, solidly colored 2 Color markings of dorsum not transverse bands nor solidly colored 16 2. (1) Dorsum infuscated throughout (1) infuscata (Gill.), p. 265 Dorsum cross-banded (not so definite in E. bistrata) 3 3. (2) Dorsum with one crossband (2) amanda McA., p. 265 Dorsum with more than one crossband 4 4. (3) Anterior crossband covering base of tegmina 5 Anterior crossband not covering base of tegmina 8 5. (4) Usually a semblance of inverted V on vertex; last abdominal segment of male more or less infuscated 6 Markings of vertex usually confined to a wide basal spot ; last ventral segment of male stramineous 7 6. (5) Markings of dorsum distinct bands (3) vitis Harris, p. 266 Markings of dorsum almost solid ; light areas confined to a circle of spots vitis var. corona McA., p. 267 7. (5) Color of dorsum often almost solid, all bands more or less connected (4) bistrata McA.. p. 267 Color of dorsum in distinct crossbands bistrata var. stricta McA., p. 268 8. (4) Anterior crossband involving base of scutellum 9 Anterior crossband not involving base of scutellum 13 Beamer: Species of Erythroneura 263 9. (8) Anterior crossband covering all of pronotum except small part of anterior margin 10 Anterior crossband leaving most of disc of pronotum bare 12 10. (9) Crossbands one and two of different colors rosa var. repitita McA., p. 282 Crossband one and two same color 11 11. (10) Crossband one and two sanguineous to black (5) tricincta Fitch, p. 268 Crossbands one and two bright red (6) diva McA., p. 269 12. (9) All crossbands continuous (7) calyrula McA., p. 270 Crossbands broken calycula var. noncincta John., p. 270 13. (8) Crossband one covering most of pronotum 14 Crossband one leaving most of disc of pronotum uncovered 15 14. (13) Cro.ssband one sanguineous to dusky (8) integra McA., p. 271 Crossband one bright red diva var. complemcnta McA., p. 269 15. (13) Crospband continuous across hind margin of pronotum. .(9) cymbiuvi McA., p. 271 Crossband broadly interrupted on hind margin cymbium var. disjuncta McA., p. 272 16. (1) Angulate vittae continuous from humerus to crossveins 17 Angulate vittae not continuous from humerus to crossveins 40 17. (16) Without definite colored spot on apex of clavus 18 With a definite colored spot; on apex of clavus 20 18. (17) Marking of vertex a broad stripe (1) fraxa Rob., p. 272 Markings of vertex definitely V-shaped 19 19. (18) Elytral markings narrow, of uniform width, red or orange (11) acuticephala Rob., p. 272 Elytral markings broad, of varying widths, brownish. .. .(12) elegans McA., p. 273 20. (17) Basal two thirds of clavus bright red compta var. rufomaculata McA., p. 292 Basal two thirds of clavus not bright red 21 21. (20) Dorsum from base of scutellum to crossveins occupied by a light diamond - shaped spot (13) cancellata McA., p. 273 Dorsum not occupied by a large light spot 22 22. (21) Pronotum usually rrdbrown to black (lateral margins light in E. aclys McAtee) 23 Pronotum not redbrown to black 25 23. (22) Lateral margins of pronotum light (14) aclys McA., p. 274 Lateral margins of pronotum dark 24 24. (23) Vertex reddish brown (15) prosata Johnson, p. 275 Vertex light with red marks (16) ziczac Walsh, p. 275 25. (22) Basal angles of scutellum usually with large black spots 26 Basal angles of scutellum without definite back spots 27 26. (25) Vertex sharp pointed, tube of aedeagus projecting, posterior point of style almost as long as foot ...(17) elegantula Osb., p. 277 Vertex bkmt, apex of aedeagus truncate (18) coloradensis (Gill.), p. 277 27. (25) Red markings of clavus overlaid with velvety black, (14) palimpsesta Mc.\., p. 278 Red markings of clavus not overlaid with black 28 28. (27) Principal markings of vertex two black fingerlike marks, (2) pontifex McA., p. 279 Principal markings of vertex not two black fingerlike marks 29 29. (28) Median longitudinal red vitta on vertex (21) kanwakae Rob., p. 279 Markings of vertex not a median vitta SO 30. (29 Main dorsal vittae more or less darkened; posterior point of style much shorter than foot (except in reflecta) 31 Main dorsal vittae orange; posterior point of style longer than foot (ex- cept in ancora) 37 31. (30) Markings of dorsum lighter, scarcely any dark color 32 Markings of dorsum distinctly dark colored 33 32. (31) Processes of aedeagus coiled near middle (Ariz.) ....(22) anjracta Beamer, p. 280 Processes of aedeagus not coiled near middle (23) vitifex Fitch, p. 28 33. (31) Color markings of vertex usually enclosing three round white dots 34 Color markings of vertex not enclosing three round white dots 35 34. (33) Basal angles of scut^-llum usually dark; outer processes of pygofer hook about one fourth as long as inner, sides of aedeagus rounded (24) caetra McA.., p. 281 264 The University Science Bulletin Basal angles of scutellum usually not so dark; outer processes of pygofer almost absent ; aedeagus in dorsoventral view with sides almost parallel (25) rosa Rob., p. 282 35. (33) Zigzag vittae of dorsum even sided; processes of aedeagus in lateral view not narrowed suddenly 36 Zigzag vittae of dorsum enlarged over disc of elytra, usually forming a com- plicated pattern ; processes of aedeagus in lateral view rapidly narrowed near middle (26) reflecta McA., p. 283 36. (35) Zigzag vittae of dorsum narrower and more definite; pygofer hook U-shaped with two processes ziczac var. walshi, n. var., p. 276 Zigzag vittae of dorsum more suffused ; pygofer hook with three proc- esses (27) prima n. sp., p. 283 37. (30) Usually dorsal markings very narrow ; aedeagus in dorsoventral view broad, bell-shaped, processes scarcely diverging (28) nudata McA., p. 284 Dorsal markings usually heavier; aedeagus not as above 38 38. (37) Aedeagus in dorsoventral view narrow, many times longer than wide 39 Aedeagus in dorsoventral view broad, not much longer than wide (29) beameri Rob., p. 285 39. (38) Posterior point of style longer than foot (30) vaga Johnson, p. 285 Posterior point of style much shorter than foot (31) aiicora Beamer, p. 285 40. (16) Markings of vertex usually more than an inverted V 41 Markings of vertex usually not more than an inverted orange V or a median stripe 50 41. (40) Markings of clavus usually consisting of three spots (E. omaska may have just two) 42 Markings of clavus consisting of two spots 44 42. (41) Markings of dorsum heavy; milky colored background not so evident; shaft of aedeagus in dorsoventral view narrow, tip bifid.. (32) omaska Rob., p. 286 Markings of dorsum very delicate; general ground color niilky white; aedeagus in dorsoventral view broad 43 43. (42) Median spot of clavus without dark spot (33) delicata McA., p. 286 Median spot of clavus with dark spot delicata var. accepta McA. p. 287 44. (41) Vertex with dark median line 45 Vertex with light median line 48 45. (44) Vertex and pronotum often suffused with brown 40 Vertex and pronotum not suffused with brown 47 46. (45) Processes of pygofer hook almost touching at apex (34) comi Rob., p. 287 Outer process of pygofer hook slightly more than half as long as inner; tip of latter enlarged (35) bidens McA., p. 287 47. (45) Aedeagus in dorsoventral view one half as broad as long, (36) rubrella McA., p. 288 Aedeagus in dorsoventral view very narrow, many times longer than wide (37) ontari Rob., p. 289 48. (44) Body very dark, showing through elytra 49 Body light, not darkening upper surface (38) aza Rob., p. 289 49. (48) Markings of dorsum broad; posterior point of style short, (39) rubra (Gill.), p. 290 Markings narrow; posterior point of style as long as foot, (40) f estiva n. sp., p. 290 50. (40) Usually some dark markings on scutellum and on anchor-shaped spot of clavus (41) compta McA., p. 291 No dark markings on scutellum and on anchor-shaped spot of clavus 51 51. (50) Process of aedeagus in lateral view with notch or projecting point near middle (42) comes (Say), p. 292 Process of aedeagus without such a notch 52 52. (51) Markings of vertex a median longitudinal vittae (43) tacita n. sp., p. 293 Markmgs of vertex usually V-shaped 53 53. (52) Aedeagus in dorsoventral view broad, bell-shaped; outer process of pygofer hook not over half as long as inner nudata McA., p. 284 Aedeagus in dorsoventral view broad but strongly constricted on basal half ; outer process of pygofer hook almost as long as inner. .(44) gilensis Beamer, p. 293 Beamer: Species of Erythroneura 265 1. Erythroneura infuscata (Gillette) Typhlocyba comes var. infuscata Gillette, C. P. Am. Typhlocybinae, 1898, p. 764. The only specimen mentioned in the original description was type number 3451 in the United States National Museum. This descrip- tion, a very short, terse one, does not fit this type 3451. W. L. McAtee, Trans. Am. Ent. Soc, vol. XLVI, 1920, redescribed the type ^as follows: "Smoky brown to black above, in some cases underlaid by deep dull red, a little paler at rondure of vertex, especially on sides, pronotum sometimes faintly paler on disc, and scutellum sometimes with a pale median vitta; tegmina with more or less hyaline spots at base, middle and apex of inner margin of clavus, middle of radial margin of fourth apical cell and on costal margin of second apical cell; costal plaque opaque whitish, tinged with red, an almost equal area of dull red just posterior, and costal margin and to some extent the crossveins pencilled with dull red. Face and legs pale yellow, abdominal incisures, etc., sometimes pale yellow." Genitalia. Pygofer hook U-shaped, outer process about half as long as inner. Foot of style medium, heel small; base almost straight; anterior point about half as long as width of foot; posterior point shorter and sharper. Aedeagus in dorsoventral view broad, tube projecting sharply, lateral margins narrowing rapidly toward base. This beautiful dusky species is quite common throughout the United States east of the Rocky Mountains. 2. Erythroneura amanda McA. Erythroneura comes var. amanda McAtee, W. L., Trans. Am. Ent. Soc., XLVI, p. 319, 1924. This striking species is easily separated from all others in the Comes group by the large, almost triangular, reddish brown spot which occupies most of clavus, corium between clavus and costal plaque and apex of scutellum. The male genitalia show a very close relation with E. cotnpta McA., from which it differs only in minor details. If the external markings were like that species it would be difficult indeed to separate them. Genitalia. Pygofer hook U-shaped. Foot of style large; heel short and sharp; base straight; anterior point short, sharp; pos- terior point longer than foot, almost parallel-sided. Aedeagus in ventral view rather broad, diamond-shaped, with each process bifid. 266 The University Science Bulletin This species differs from compta in having the base of the processes of the aedeagus somewhat heavier and in the sides of the pygofer hook being longer. Specimens are at hand from Douglas, Atchison, Cherokee and Wyandotte counties, Kansas. 3. Erythroneura vitis (Harris) Tettigonia vitis Harris, T. W., Encyclopedia Americana, VIII, p. 43, 1831. Erythroneura vitis var. flava Robinson, Can., Ent., vol. LVI, No. 3, p. 62; 1924. Erythroneura vitis var. venusta McAtee, W. L., Trans. Am. Ent. Soc., XLVI, p. 319, 1920. The original description is as follows: 'Tt may be called tetti- gonia vitis (Harris). It is, in its perfect state, nearly one tenth of an inch long; of a straw color, with two broad, scarlet bands across the wing cases, one at the base and the other on the middle, and the tips of the wing cases are blackish." General ground color yellowish white. Vertex marked with pair of reddish, longitudinal arcs, probably remnant of orange in- verted V. Pronotum of type male, red on lateral margins, much lighter on median two thirds with darker Y-shaped mark at mid- dle. Most specimens are uniformly dark reddish-brown through- out most of surface. Scutellum of type buff with two longitudinal pink stripes and pink tip. Elytra with base and median band bright red. Anterior cross band wider than scutellum on costal margin, narrower next scutellum. Median band not quite cover- ing tip of claws. Costal plaque with large black spot, cross veins more or less red, beyond infuscated. Venter more or less stramin- eous, mesosternum usually darker and last ventral segment of male darkened. Genitalia. Style with foot short; heel moderate; base curved; anterior point sharp, slightly longer than posterior; posterior point narrower and shorter, about half as long as base. Aedeagus small, tip broad, sharply narrowed on basal two thirds. Pair of heavy almost parallel processes, longer than aedeagus, almost straight. Type in collection of Boston Society of Natural History. This species is usually easily separated from closely related forms by the semblance of an inverted V on the vertex and the darkened last ventral segment of the male. Through the courtesy of the board of directors and Mr. Richard Dow, of the Boston Society of Natural History, I was able to study the male genitalia and a water color of the type. This is a common species on grape throughout the United States east of the Rocky Mountains. Beamer: Species of Erythroneura 267 Erythroneura vitis var. corona McAtee Erythroneura litis var. corona, McAtee, W. L., Trans. Am. Ent. Soc, 46, p. 305, 1920. Original description. "Similar to variety vitis Harris, but middle crossband on tegmina connected by red lines along longitudinal veins to anterior crossband and to ramose marking on cross veins, so that the saddle mark appears to be surrounded by a wreath of pale spots." A study of male genitalia of specimens with the above markings shows no marked difference from vitis Harris. The markings are so characteristic, however, that the varietal name is retained. Type, female, Plummer's Island, Maryland, in collection of Mc- Atee. Since McAtee did not designate an allotype a male from Windsor, Vermont, 21, VIII, 1934, R. H. Beamer, is here so desig- nated. This variety is even more plentiful on grape than E. vitis over the same territory. The same dark last ventral segment in the male and the inverted V of the vertex, as in E. vitis, separate this species from the E. bistrata forms and the enlarged middle band encircled with a ring of white spots sets it off from E. vitis. 4. Erythroneura bistrata McAtee Erythroneura vitis var. bistrata McAtee, W. L., Trans. Am. Ent. Soc., XLVI, p. 305, 1920. Erythroneura fusco-clava Beamer, R. H., Can. Ent. 1927, p. 31, LIX. Original description. "This variety illustrates the extreme degree of erythrization of this species I have seen; the vertex may be pale yellow, but usually it has the normal pair of vittae strongly de- veloped, and it may be almost entirely red; the pronotum is red with touches of pale yellow on anterior border, with a dot in middle and traces of pale yellow at about two points on the hind margin ; scutellum red with a pale median vitta enlarged at apex; tegmen often entirely red with the exception of a pale spot at inner base of clavus and another involving apex of clavus and adjoining corium; there may be other traces, however, of both anterior and posterior pale bands, and the costal plaque, except for its posterior blackish marking, and an equal sized area behind it and the bases of the apical cells are pale; undersurface and legs chiefly pale yellow." Dissection of the male type revealed a type of genitalia suf- ficiently different from E. vitis (Harris) to be worthy of specific rank. Genitalia. Pygofer hook typically U-shaped. Foot of style of medium length, heel definite; base almost straight; anterior point 268 The University Science Bulletin short and sharp, about as long as foot is wide inside points; pos- terior point about two thirds as long as foot, slightly wider at base than anterior, sides almost straight, converging to sharp point. Aedeagus in dorsoventral view more or less diamond-shape, widest near middle and tapering to rather narrow apex and slightly wider base. Aedeagal processes rather slender, reaching almost to tip of shaft, then bending at right angles for about one third their length. Type male in the collection of W. L. McAtee. As Mr. McAtee states in the description, this species is quite va- riable in color pattern. The vertex markings, however, are usually just rectangular spots covering most of base, leaving outer third, at least, unmarked. This and lack of the darkened last ventral seg- ment in the male easily separates this form and E. bistrata var. stricta McAtee from E. vitis (Harris) and its variety. It occurs in large numbers in some localities, but not so plentiful usually as the variety stricta. Erythroneura bistrata var. stricta McAtee Erythroneura vitis var. stricta McAtee, W. L., Trans. Am. Ent. Soc, XLVI, p. 305; 1920. Erythroneura nibranotum Rcbin-son, Wm., Can. Ent., vol. LVI, No. 3, p. 60; 1924. Original description. "Ground color whitish hyaline to pale yel- low; vertex chiefly pale, sometimes with a few touches of red, pro- notum and scutellum chiefly pale, sometimes with a few touches of red, pronotum and scutellum chiefly red to dusky, the latter some- times pale medianly and at tip; tegmen with three crossbands, the anterior red, narrow, not exceeding scutellum, the middle one nar- rower than in the other varieties, usually nearly quadrilateral in shape ; the posterior much as in first variety, pale spaces between the bands broader than in the other varieties; usually half or more of costal plaque bluish or blackish, often overlaid by opaque white; lower surface stramineous to pale yellow." Dissections of males of this variety exhibit the same type of geni- talia as E. bistrata, so it is given as a variety of that species. The vertexal marks and the color of the last ventral segment of the male are the same as in that species. This is a very common variety, often occurring in great numbers on grape. Distribution same as for E. bistrata. 5. Erythroneura tricincta Fitch Erythroneura tricincta Fitch, Asa, N. Y. State Cab., p. 63, 1851. Original description. "Three-banded Erythroneura, E. tricincta. Pale yellow, with three broad bands, the anterior velvet-black, oc- Beamer: Species of Erythroneura 269 ciipying the thorax and basal half of the scutel; the middle bright ferruginous ending outwardly in black, forward of the middle of the elytra, the posterior dusky brown, on the apex. Length, 0.12. No. 823, female." Genitalia. Py gofer hook U-shaped, outer process about one third as long as inner. Foot of style medium ; heel medium ; base slightly curved; anterior point broad, almost as long as toe is wide; posterior point less than half as long, sharp. Aedeagus in dorsoventral view broad, tube projecting, processes longer than shaft, in lateral view bent around tip of shaft. This is the outstanding internal character in separating this form from E. calycula McA. Even though a very short description, it is very good and one should have no trouble in determining this insect. It is very com- mon throughout the eastern half of the United States and Canada. 6. Erythroneura diva McAtee Erythroneura tricincta var. diva McAtee, W. L., Trans. Am. Ent. Soc, XLVI, p. 308; 1920. Erythroneura tricincta var. rubravitta Robinson, Wm., Can. Ent., vol. LVI, No. 7, p. 156; 1924. Original description. "Crossbands one and two chiefly bright red; crossband one covering most of pronotum and base of scutellum, where its color sometimes is brownish yellow instead of red; cross- band two, bright red to costal plaque which is chiefly blackish, more or less overlaid by opaque white; longitudinal color markings sul- phur yellow." Genitalia. Pygofer hook U-shaped, outer process almost as long as inner. Foot of style medium; heel almost absent; base straight; anterior point broad, almost as long as toe is wide; posterior point sharp and short, not more than one third as long as anterior. Aedea- gus in dorsoventral view broad, almost circular, processes about as long as shaft with tooth near middle on outside margin. It is this tooth that is the distinctive internal character of this species. Since this species was described from a holotype female, a male paratypc, Plummer's Island, Md., Jan 23, 1916, W. L. McAtee, de- scribed above, is here designated allotype. A fairly common species throughout the eastern half of the United States. Erythroneura diva var. complementa McA. Erythroneura tricincta var. complemcnta McAtee, W. L., 111. Nat. Hist. Surv., vol. XVI, Art. Ill, p. 135; 1926. Original description. "Crossbands one and two bright red, three dusky, differing from var. diva McA. in crossband one being con- fined to pronotum ; subsidiary markings yellow. Length, 2.75 mm. 270 The University Science Bulletin "Holotype. Female, Ocean Springs, Miss., Aug. 4, 1921, C. J. Drake." Numerous specimens of this beautiful insect from Orange county, Texas, have been studied Vi^ith no real difference becoming apparent between it and E. diva McA. other than the darker base of the scutellum in diva. In my estimation it is but the summer form of this species. Specimens are at hand from Orange county, Texas, Inverness, Hilliard, Wildwood and Fort Meade, Fla. 7. Erythroneura calycida McAtee Erythroneura tricincta var. calycida McAtee, W. L., Trans. Am. Ent. Soc, XLVI, p. 308, 1920. Erythroneura tricincta var. erasa McAtee, W. L., Trans. Am. Ent. Soc., XLVI, p. 309, 1920. Original description. "Ground color ivory, a broad U-shaped dusky to black band on base of scutellum and sides of pronotum, the often concolorous eyes extending it on vertex; crossband two san- guineous to dusky, black on costa; longitudinal color streaks red." This is a beautiful, quite definitely marked species and the color of the first crossband, as described by McAtee, will separate it from all others. Genitalia. Pygofer hook U-shaped, outer process about one third as long as inner. Foot medium; heel medium; base almost straight; anterior point wide, as long as toe; posterior point sharp, not over one third as long as the anterior. Aedeagus in dorsoventral view broad, tube very slightly projecting, processes about as long as shaft not curved around the tip in lateral view. Common on grape throughout the eastern half of the United States and Canada. Erythroneura calycida var. nondncta Johnson Erythroneura tricincta var. nondncta Johnson, Dorothy M., Ohio Jr. Sci. No. 4, p. 261, 1934. Original description. ''There are no true crossbands in this vari- ety which is closely related to Erythroneura tricincta var. calycula McAtee, as the basal angles of scutellum and sides of pronotum be- hind the black eyes are dark. There are dark reddish triangular spots, bordering side of costal plaques, and not attaining clavi which are immaculate. The background is creamy white and faint yel- low-colored streaks follow the longitudinal veins of the elytra. The cross veins are narrowly black, heavily margined anteriorly, except the fourth, with bright red; apical cells dusky." Beamer: Species of Erythroneura 271 This is a variety of doubtful validity although a few specimens are at hand from Louisiana. It occurs very rarely. 8. Erythroneura Integra McAtee Erythroneura tricincta var. Integra McAtee, W. L., Trans. Am. Ent. Soc., XLVT, p. 309, 1920. Original description. "Crossband one, sanguineous to dusky, con- fined to pronotum, most of which it covers; crossband two bright red to sanguineous, bluish to black on costa; longitudinal color markings red." A very distinctive species marked as described above. Genitalia. Pygofer hook U-shaped, outer process very short, about one fourth as long as inner. Foot of style medium; heel small; base curved; anterior point broad, about as long as width of foot in narrowest place; posterior point narrow, sharp, about one fourth as long as anterior. Aedeagus in dorsoventral view almost circular, tube projecting, processes about as long as shaft, slightly diverging on outer half. A male paratype, Plummer's Island, Md., March 18, 1917, W. L. McAtee, described above, is here designated allotype. Holotype fe- male, allotype, and paratypes in McAtee collection. Fairly com- mon throughout the eastern half of the United States. 9. Erythroneura cymbium McAtee Erythroneura tricincta var. cymbium McAtee, W. L., Trans. Am. Ent. Soc., XLVI, p. 310, 1920. Original description. "Crossband one, dusky to black, confined to pronotum, profoundly emarginate anteriorly, leaving a large part of disk uncolored; crossband two, sanguineous to red, bluish to black on costa; longitudinal color streaks yellow." Genitalia. Pygofer hook U-shaped, outer process almost as long as inner. Foot of style medium; heel marked; base curved; ante- rior point broad at base, almost as long as width of foot; posterior point, short, sharp, about one third as long as anterior. Aedeagus in dorsoventral view almost circular, tube slightly projecting, proc- esses about as long as shaft, slightly diverging on outer third. A common species over the eastern half of the United States. Holotype female, Pennsylvania, in collection of W. L. McAtee. Allotype male, described above. White county, Illinois, March 31, 1929, in Snow Entomological Collection. 272 The University Science Bulletin Erythroneura cymbium var. disjuncta McAtee Erythronexira tricincta var. disjuncta McAtee, W. L., Trans. Am. Ent. Soc, XLVI, p. 310, 1920. Original description. "Like the last, cymbium, but crossband (if it may be so called) one, broadly interrupted in the middle, cover- ing only sides of pronotum." A variety of doubtful value, but sometimes occurring. A few specimens are at hand from Louisiana and Illinois. 10. Erythroneura fraxa Rob. Erythroneura fraxa Robinson, W. M., Can. Ent., vol. LVI, No. 12, p. 292; 1924. This beautiful red marked species described by Robinson from a single female has since been found in numbers. A male specimen from Anderson county, Kansas, November 26, 1927, is figured as the allotype. Genitalia. Pygofer hook of typical U-shape with points of U rather shorter than usual. Style with small foot; base almost straight; anterior point short, and sharp; posterior point about half as long as base and quite sharp. Aedeagus in ventral view rather bluntly diamond-shaped with slender processes roundingly separated. This species resembles E. kanwakae Rob., but is larger, vertex blunter, orange spot of vertex in form of round spot rather than parallel-sided stripe, spot on tip of clavus more just an indication of a spot, seemingly about to disappear, pygofer hook with but two prongs and aedeagus in dorsoventral view more diamond-shaped than rectangular. Numerous specimens at hand from Kansas, Arkansas, and Illinois. 11. Erythroneura acuticephala Robinson Erythroneura acuticephala Robinson, Wm., Can. Ent., vol. LVI, No. 3, p. 61 ; 1924. This bright orange-marked little species resembles E. kanwakae Rob., but may be easily separated by the absence of most dark marking on abdomen, by posterior point of style as long as foot and without extra process in pygofer hook. Qenitalia. Pygofer hook typical U-shaped. Foot of style small; heel prominent; base curved, anterior point fairly long and thick; posterior point more than half as long as base, very thin and sharp. Aedeagus in ventral view truncate with a small projection at either corner, processes small, about as long as shaft, diverging to about width of shaft. Beamer: Species of Erythroneura 273 Specimens at hand from Douglas, Anderson, and Cherokee counties, Kansas, and Ramsey county, Minnesota. 12. Erythroneura elegans McAtee Erythroneura comes van elegans McAtee, W. L., Trans. Am. Ent. Soc, XL VI, p. 305; 1920. General ground color yellowish white marked with orange or red- dish-brown. Vertex with inverted U-shaped mark, arms enlarged at base, continued on pronotum as very heavy Y-shaped mark. Pronotum with an additional vitta back of each eye. Scutellum mostly dark reddish with lighter spot near apex. Elytra with six light areas, two more or less pentagonal mesal spots, smaller at tip of scutellum, other on apex of clavus, one angular dash from costa to claval suture between humerus and plaque and the other between the plaque and cross veins. Cross veins pink or red. Apical cells more or less infuscated, darker spot in costal plaque and apex of cell Rg, Mesosternum dark. Genitalia. Pygofer hook U-shaped, inner process about one- fourth longer than outer. Foot of style small; heel prominent; base almost straight; anterior point two thirds as long as width of foot at narrowest place; posterior point slightly shorter, both sharp. Aedeagus in dorsoventral view more or less three pronged, processes about as long as shaft, almost parallel and about straight in lateral view. Redescribed from the male holotype, Plummer's Island, Md., Jan. 11, 1914; W. L. McAtee. In his collection. This species externally resembles E. ziczac Walsh, but may be sep- arated from that species by the lack of an orange spot on apex of clavus and by the colored zigzag vitta of dorsum being of varied widths. A beautiful species distributed throughout most of the United States. 13. Erythroneura cancellata McAtee Erythroneura comes var. can-cellata McAtee, W. L., Trans. Am. Ent. Soc, XLVI, p 320; 1920. This is one of the most striking species in the genus with its beau- tiful lighter colored diamond-shaped spot occupying a large portion of dorsum of elytra, surrounded by the darker wine color. Genitalia. Pygofer with typical comes U-shaped hook. Style with small foot; base curved; anterior point short and heavy; pos- 18—6037 274 The University Science Bulletin terior point short and sharp. Aedeagus in lateral view almost cir- cular with processes diverging one third distance before tip. Although not as plentiful as some, specimens of this species are not uncommon in the states east of the Rocky Mountains. This species was described from 2 females from Plummer's Island, Md., one designated type, the other paratype and in the collection of W. L. McAtee. A male from Anderson county, Kansas, described above is therefore designated allotype. Miss Dorothy M. Johnson's statement "female (holotype) Adams county, IX-1-31, E. P. B." Ohio State Univ. Bui. 31 Vol. 39, 1935, p. 109, is evidently an error. 14. Erythroneura aclys McAtee Erythroneura aclys McAtee, W. L., Trans. Am. Ent. Soc, vol. XLVI, p. 290, 1920. Original description. "Base of fourth apical cell angulate, vertex rather pointed; color pattern consisting of a broad, brown to black stripe extending whole length of insect, margined by pale yellow. The dorsal stripe nearly black on scutellum and anterior parts sharply cut off from a pale yellow margin about the width of eye; on tegmina the stripe is smoky brown, greatly expanded and irregu- larly set off from the pale yellow costal margin, percurrent to second apical cell which is about the width of costal plaque; the latter has a slight opaque whitish coating, is margined interiorly and anteriorly by reddish, and underlaid posteriorly by blackish; there are touches of red upon apex of clavus and first cross vein and more or less hyaline spots at extremities of apical third of clavus, on the corium nearby and in middle of fourth apical cell. The legs, pleura and face vary from flesh-color to pale yellow, and the venter is pale yellow with a median series of slaty spots, the posterior ones extending en- tirely across last abdominal segment and across base of genitalia; dorsum slaty." Genitalia. Pygofer hook U-shaped, outer process about two thirds as long as inner. Foot of style medium; heel small; base straight; anterior point almost as long as width of foot; posterior point about as long as base. Aedeagus in dorsoventral view, broad, almost quad- rangular, tube projecting, lateral margins with long sharp projecting apices. This is a very common species throughout the United States east of the Rocky Mountains. Beamer: Species of Erythroneura 275 15. Erythroneura prosata Johnson Erythroneura infuscata var. prosata Johnson, Dorothy M., Ohio Biol. Survey Bui. 31, p. 108, 1935. Resembling E. ziczac Walsh, but with much more dark coloring on vertex pronotum and elytra, by posterior point of style being as long as base of foot, sharp, and by processes diverging radically on outer third. General color, blood-brown. Vertex sometimes with red inverted V evident, often with median longitudinal stripe and two lateral spots light. Thorax also with Y-shaped median and lateral red dashes more or less evident. Scutellum usually with a median longitudinal light rectangular spot, often spool-shaped. Clavus with small median light spot on mesal margin, another smaller spot slightly outside clavai sutm'e and toward cross veins and an indi- cation of a light spot on apex, the three forming a semi-circle. Corium with oblique oblong area anterior to costal plaque; costal plaque, except dark oblique spot in outer end and two spots beyond plaque, light. Light areas usually bounded with red. Cross veins ligliter, apical cells dark with clear areas sometimes present. Meso- sternum dark. Genitalia. Pygofer hook U-shaped, outer process about half as long as inner. Foot of style medium; heel prominent; base straight; anterior point half as long as width of foot, sharp; posterior point as long as base, straight. Aedeagus in dorsoventral view broad, tube protruding, shaft sharply constricted on basal half, processes sharply diverging on outer half. Named from three specimens from Pickaway county, and Coal Grove, Ohio. Numerous specimens are at hand from Wabash county, Illinois. 16. Erythroneura ziczac Walsh Erythroneura ziczac Walsh, B. D., Prairie Farmer 10, No. 10, 1862, p. 149. The original description is as follows: 'Tale yellowish. Front of head in a right angle, the apex rounded. Eyes dusky; ocelli pale; vertex with two pale sanguineous vittae, generally subobsolete. Thorax blood-brown, yellowish in front, often with a yellowish vitta; scutel blood-brown with a yellowish vitta, occasionally en- tirely yellowish. Abdomen often blood-brown, except at base and tip; tip of ovipositor and of tarsi dusky. Elytra pale yellowish; on the middle of the costa an oblique rhomboidal black spot, on the 276 The University Science Bulletin costal tip a black dot, and on the interior margin, forming a tri- angle with the other two spots, a black spot; a blood-brown, irregular broad strip, covering the humerus, thence running to the interior margin, thence in a zigzag direction to the rhomboidal spot, thence to the spot on the interior margin, thence not quite attaining the terminal dot. Wings hyaline; tips of costal veins, often dusky. Length to tip of wings a little over one tenth inch. Occurs abun- dantly on the grapevine." A careful study of Walsh's description with specimens at hand reveals the fact that what has commonly been called E. ziczac is not true ziczac at all but the light form of this species which I am describing as E. ziczac var. walshi. True E. ziczac is much darker throughout with all of the pronotum "blood-brown" except a small rectangular area on anterior margin. Since Walsh's types are lost a male and female from Wabash county, Illinois, March 31, 1929, are here selected as neotypes. Other specimens are at hand from Illinois, Ohio, Indiana, New York, and Vineland, Ontario. (Plate XIX, fig. 16.) Erythroneura ziczac var. walshi n. var. Resembling E. ziczac Walsh but with lighter coloring, narrower markings, and pronotum never dark colored on basal two thirds. Color. General ground color yellowish white, markings red to dusky. Vertex with inverted V-shaped red mark, sometimes closed basally. Pronotum with Y-shaped and lateral bands red to dusky. Scutellum with basal angles yellow bordered with red, apex red. Tegmen with red to dusky zigzag stripe arising on humerus, ending at base of cell M4. Red spot on apex of clavus. Cross vein red. Apical cells more or less dusky. Holotype male Jefferson county, Nebraska, Beamer and Hunger- ford. Allotype female, Reno county, Kansas, August 13, 1924, Beamer and Lawson, and the following paratypes, 10 males and 6 females same data as allotype; 7 males and 11 females, Douglas county, Kansas; 5 males and 2 females, Atchison county, Kansas; 3 males and 6 females, Jordan, Ontario, Aug. 1922, Wm. Robinson; 3 males and 3 females, Vineland, Ontario, Wm. Robinson; 1 female Clayton county, Iowa, 1930, R. H. Beamer. Examination of the genitalia of many specimens failed to reveal any characteristic differences between this and E. ziczac Walsh. They are, however, easily distinquished by the absence of the dark coloring on the posterior two thirds of the pronotum. Beamer: Species of Erythroneura 277 17. Erythroneura elegantula Osborn Erythroneura elegantula Osborn, Herbert, An. Carnegie Mus., Pitts., 1928, p. 289. Resembling Erythroneura coloradensis (Gill.), but with much more pointed vertex, widely separated arms of the inverted V of the vertex, rounded tip of the aedeagus in ventral view and poste- rior point of style being longer than anterior. General ground color ivory with principal markings red or orange. Vertex with narrow-sided inverted V with arms widely separated, sometimes touching eyes. Pronotum with median Y-shaped mark sometimes not touching either margin and a dash back of each eye. Scutellum usually with basal angles black but often just yellow, apex usually red or orange. Clavus with typical anchor-shaped spot and smaller one on apex. Corium with dash on base, an irregular sided vittae surrounding costal plaque, extending to claval suture before apex and usually fading out about base of cell M^. Black dash in posterior end of costal plaque, small round black spot in apex of cell R3 and larger one in base of cell R^. Abdomen dark, especially the dorsum, which often gives the elytra a darker appear- ance. Genitalia. Pygofer hook U-shaped, one arm much shorter than other. Style with small foot; heel small but prominent, base curved, anterior point about twice as wide as posterior and about as long, posterior point sharper, about as long as base. Aedeagus in ventral view more or less globular, somewhat pointed at apex, usual pair of processes rather short and almost parallel. Reclescribed from the types in Osborn Collection labeled "Canal Zone, Panama." This is one of the commonest and most destructive species found on grape in California. Specimens are also at hand from Colorado and Kansas. 18. Erythroneura coloradensis (Gill.) Typhlocyba vitifex var. coloradensis Gillette, C. P., Bui. 19, Colo. Agr. Exp. Sta., May, 1892, p. 16, fig. 8. Redescribed from the types in U. S. Nat. Mus. General ground color semihyaline to yellowish white. Markings orange. Vertex with inverted orange V not touching eyes. Pronotum with median V, heavy basally, arms short; lateral dashes back of eyes, not reach- ing posterior margin. Scutellum with basal angles black surrounded by yellow tip tinged with orange. Clavus with heavy basal anchor spot and another at tip. Corium with oblique dash near base and an irregular sided vitta arising on costa at anterior end of costal 278 The University Science Bulletin plaque, almost surrounding it, touching claval suture near apex of anchor spot, following vein Cu to end in cross veins. Black spot in posterior end of costal plaque, another in apex of cell R3, a third in base of cell M^, cross veins pink, apex of elytra more or less infus- cated. Venter stramineous, face marked with pink, mesosternum and abdomen more or less infuscated. Genitalia. Style with short foot; heel inconspicuous; anterior point longer than foot, wider than posterior ; posterior point shorter and sharper almost as long as base. Tip of aedeagus in dorsoven- tral view tiTincate, processes reaching tip, slightly diverging. Pygo- fer hook typically U-shaped. Hololectotype male No. 3447, U. S. Nat. Mus., allolectotype, fe- male, No. 3447, U. S. Nat. Mus. This is a very common species in the central United States on grape and Virginia creeper. It is easily separated from other species by the blunt vertex and the two round black spots in basal angle of scutellum. 19. Erythroneura palimpsesta McAtee Erythroneura comes var. -palimpsesta McAtee, W. L., 111. Nat. Hist. Survey, Bui., vol. XV, Art. II, p. 43, 1924. Original description. "Like E. conies var. vitifex Fitch except that the red vitta on anterior half of clavus and the adjacent short vitta on corium are overlaid or replaced by black ; the lateral vittae on pronotum and the sides of the scutellum also may be dusky to black, and the oblique dusky band of antennae, pro- and mesopleuro, base and apex of genitalia and broad vittae on dorsum of abdomen also may be dusky to black. Length, 3 mm. "Holotype male, and two females, one of which is allotype. Forest City, III, April 3, 1917." This species is easily separated from all others, as McAtee points out by the black coloring overlying the red in the basal anchor- shaped mark of the clavus as well as some of the spots of the corium. The red mark of the vertex differs too in that it is oval or sometimes almost round. Genitalia. Pygofer hook U-shaped, outer process about one fourth shorter than inner. Foot of style large; heel prominent, projecting; base curved; anterior point short, blunt; posterior point longer than foot, narrow, almost parallel sided. Aedeagus in dorsoventral view bell-shaped, apex truncate, processes as long as shaft diverging on outer half. Beamer: Species of Erythroneura 279 This species is quite rare. Specimens are at hand from Illinois and Kansas. Types in the Collection of the Illinois Natural History- Survey, Urbana, 111. 20. Erythroneura pontifex McAtee Erythroneura conies var. pontifex McAtee, W. L., 111. Nat. Hist. Sur. Bui., vol. 16, p. 136, 1926. Erythroneura breakeyi Johnson, Dorothy M., Ohio Jr. Sci., No. 4, p. 261, 1934. Original description. "Like E. comes var. refiecta McAtee (Bui. III. State Nat. Hist. Sur., 15, Art. II, April, 1924, p. 43, Md., Va., 111., la., Kan.), but with two black finger-shaped vittae on vertex over- lying an inverted heart-shaped brownish marking; a marking some- what similar to latter can be seen through the disk of pronotum. Length, 3 mm. Holotype female, Dubois, 111., May 24, 1917. Deposited in the collection of the Illinois State Natural History Survey." General ground color white, markings reddish fumose to black. Vertex with inverted V with arms usually connected basally, mostly black. Pronotum with infuscated red Y and a short red dash back of each eye. Scutellum Avith basal angles yellow margined with red, apex red. Zigzag markings of elytra more or less infuscated. Cross veins red. Apical cells more or less fumose. Genitalia. Py gofer hook U-shaped, outer process about half as long as inner. Foot of style large; heel prominent; base almost straight; anterior point very short; posterior point longer than foot, sides almost parallel. Aedeagus in dorsoventral view with very broad lateral wings, canal protruding, processes longer than shaft, bent almost at right angles out on outer third. Processes in lateral view without abrupt narrowing as in E. refiecta McAtee. Since this species was named from a female holotype, the male described above, Wabash county, Illinois, March 31, 1929, is desig- nated allotype. This species resembles E. refiecta McAtee, but is easily separated from that species by the black on the red inverted V of the vertex. Not a very common species, but found throughout the eastern half of the United States. 21. Erythroneura kanwakae Robinson Erythroneura kanwakae Robinson, Wm., Can. Ent., vol. LVI, No. 12, p. 292; 1924. This species closely resembles E. fraxa Rob., but may be sepa- rated from it by being smaller with head much sharper pointed, 280 The University Science Bulletin with much more dark coloring on venter of abdomen, aedeagus almost rectangular in dorsal view, posterior point very short and pygofer hook with three processes instead of two. Genitalia. Pygofer hook U-shaped with an additional process somewhat longer than shorter process and slightly shorter than the longest. This character is unique in this species. Foot of style very short; base curved; both points short and not very sharp. Aedeagus about rectangular in ventral view, sides almost parallel, processes present, about as long as shaft, slightly divergent at tip. Specimens are at hand from Kansas, Nebraska, and Iowa. 22. Erythroneura anfracta Beamer Erythroneura anfracta Beamer, R. H., Ann. Ent. Soc. Am., vol. XXII, p. 123; 1929. Most easily confused with E. vitifex Fitch, from which it may be separated by the fumose coloring in the red markings, by each process of aedeagus having a distinct spiral near base, sometimes incomplete and by aedeagus in ventral view having narrower lateral processes. Common in Arizona on grape. 23. Erythroneura vitifex Fitch Erythroneura vitifex Fitch, Asa. Rep., N. Y. Ag. Soc. 10, pp. 392-393, 1856. Original description, in part: "Yellowish white, the wing covers with oblique confluent blood red bands and a short oblique black line on the middle of their outer margin; thorax commonly with three red stripes, the middle one forked anteriorly and confluent with two red stripes on the middle of their outer margin; thorax commonly with three red stripes, the middle one forked anteriorly and confluent with two red stripes on the crown of the head. Length, 0.12. AVhen the wing covers are closed they appear red with a cream-colored spot shaped like a heart anteriorly, and on their middle a large diamond-shaped spot with a small red spot in its center. It resembles an individual of the comes of Say, having the red spots so enlarged as to all run together, but the marks upon the thorax are totally different from those of that species . . ." General ground color yellowish white marked with orange. Ver- tex with orange inverted V, continued across pronotum as Y. Pronotum with an additional stripe on lateral margin. Scutellum with basal angles and tip orange. Elytra with rather broad zigzag vitta from humerus to cross veins, more or less surrounding costal plaque. An additional oblique dash on costa near base. Cross Beamer: Species of Erythroneura 281 veins red. Apical cells more or less infuscated. Black dash in posterior end of costal plaque, black spot in base of cell M^ and apex of cell R3. Mesosternum black. Genitalia. Pygofer hook U-shaped outer process about one third as long as inner. Foot of style small; heel not prominent; base more or less curved; anterior point broad, about as long as foot in narrowest place; posterior point short and sharp, not more than half as long as anterior. Aedeagus in dorsoventral view heart- shaped, processes longer than shaft, often diverging. Since the types of this species are lost the male, Vineland, On- tario, Aug., 1922, described above is designated neotype. E. vitifex Fitch closely resembles E. heameri Rob., but may be separated by the dark mesosternum and by the short posterior point of foot of style. This is a damaging hopper on grape throughout the eastern half of North America. 24. Erythroneura caetra McAtee Erythroneura ligata var. caetra McAtee, W. L., Pioc. Biol. Soc, Wash., vol. 37, p. 133, Dec, 1924. E. caetra McA. resembles E. Rosa Rob. quite closely, but occurs west of the Rocky Mountains, the color of pronotum usually is not evenly distributed, and outer process of pygofer hook is about one third as long as inner. General ground color semihyaline to yellowish white with dusky and reddish markings. Vertex yellowish with some darker shading, nothing very definite. Pronotum with semblance of U-shaped darker mark; large angular brown spots tinged with red back of each eye. Scutellum with tip orange, basal angles dark brown. Clavi with usual anchor-shaped basal mark very large, tip tinged with pink, remainder dusky, elongated pink spot on tip. Coria with irregular- sided pink mark beginning opposite pink tip of anchor-shaped mark on clavi, reaching to plaque, following around it almost to costa at its posterior end, ending on Cu before base at cell M^. Black or smoky stripe running almost length of R3, crossing base of M2 and ending in darker spot in base of M4. Black spot in posterior end of costal plaque. Tip of front with black spot, mesosternum shining black, dorsum of abdomen and most of ventral sclerites black. Genitalia. Pygofer hook semi-U-shaped, the outer process much shorter than in typical comes groups, about one fifth as long as inner. Foot of style rather small; heel medium; anterior point long, sharp. 282 The University Science Bulletin less than right angle; posterior point very short, obtuse angled. Aedeagus in ventral view flattened, almost parallel sided, constricted just before tip, ventral processes usually short, almost straight, not reaching nearly to tip. In lateral view shaft almost straight, broad, almost half as wide as long, evenly hollowed out at tip, ventrally notched. A male, described above. La Fayette, Cal, July 14, 1933, Mrs. R. H. Beamer, is here designated allotype. The species was de- scribed from a single female in the collection of E. D. Ball. Other specimens are at hand from Idaho, Oregon, and British Columbia. It is commonly found on willow. According to Doctor Ball's notes this is the host of the type. It is possible this is a synonym of E. rosa Rob. as they are very close indeed, but since the ranges are different and there seems to be some slight morphological differences I prefer to consider them distinct. 25. Erythroneura rosa Robinson Erythroneura rosa Robinson, Wni., Can. Ent., vol. LVI, No. 3, p. 58, March, 1924. Erythroneura mallochi McAtee, W. L., Bui. 111. Nat. Hist. Sur., vol. XV, Art. II, p. 41, April, 1924. This species is quite variable in coloring. It may usually be dis- tinguished from E. caetra McA. by having markings of the clavus evenly distributed, by having a broader aedeagus in ventral view and the short point of pygofer hook almost absent. Genitalia. Pygofer hook sickle-shaped, outer point very short. Foot of style short; heel almost absent; anterior point about half as long as width of toe, sharp; posterior point almost absent. Aede- agus in dorsoventral view rectangular, lateral margins almost par- allel, processes shorter than shaft, diverging sharply near tip. This is a very common species on willow in the United States east of the Rocky Mountains. Erythroneura rosa var. repetita McAtee Erythroneura repetita McAtee, W. L., Bui. 111. Nat. Hist. Survey, vol. XVI, Art. Ill, p. 131, Aug. 10, 1926. Specimens of this variety may easily be separated from those of E. rosa Rob. by having the color of the dorsum in the form of three crossbands. The anterior crossband is usually darker while the middle one is often quite red as in E. vitis (Harris). Numerous specimens of this variety were swept from willow in Scott county, Arkansas, August, 1928, along with a lot of other variations, includ- ing typical E. rosa Rob. Beamer: Species of Erythroneura 283 26. Erythroneura reflecta McAtee Erythroneura comes var. reflecta McAtee, W. L., 111. Nat. Hist. Sur. Bui., vol. 1.5, p. 43, April, 1924. Erythroneura portea Robinson, Wm., Can. Ent., vol. 56, No. 7, p. 154, 1924. Original description. "Much like E. comes var. rubra Gillette in connection with which it was mentioned in my key, but the pale markings are more extensive, and the red ones, instead of being a solid jasper-red as in that form, are dilute bluish-red with carmine edgings; the fact that all of the red vittae are of this compound character gives the variety the appearance of having the most com- plex pattern of any of the comes varieties. "Type female, Plummer's Island, Md., etc." General ground color yellowish-white, with the following mark- ings: Vertex with inverted V-shaped, reddish vitta. Pronotum with a slaty-brown area on disk with two reddish arms projecting forward to meet the vitta on vertex; a lateral vitta behind each eye. Scutel- lum with basal angles yellow surrounded with red, apex yellow. Tegmina, a reddish vitta arising in base of clavus, soon broadening and proceeding across corium to costal plaque, where it becomes as wide as length of plaque, then proceeding inward and dividing into two arms, one of which touches tip of clavus and the other the base of cell M^. This vitta is sometimes paler within and margined in dark red; sometimes bluish within dark red margin; tip of clavus pale red. Cross veins crimson, apical cells more or less infuscated. Genitalia. Py gofer hook U-shaped, outer process about half as long as inner. Foot of style large; heel prominent; base straight; anterior point short, slightly less than a right angle; posterior point longer than foot, narrow, almost parallel sided. Aedeagus in dor- soventral view broad, tube slightly projecting, processes longer than shaft, diverging sharply on outer half, in lateral view contracting sharply near middle. This is a very common species throughout the eastern half of the United States and Canada. It is quite variable in color. Especially is the summer form different from the overwintering. It is often quite light on anterior half while the elytra have the markings con- centrated into a large circular area, which is often bright red. The contracted process of the aedeagus seen in lateral view separates this species from all others. 27. Erythroneura prima n. sp. Resembling E. rubra (Gill), but color on vertex in two longitu- dinal vitta, color on elytra in even sided zigzag stripes, pygofer hook 284 The University Science Bulletin with three processes, and processes of aedeagiis with recurved lateral hooks. General ground color yellowish-white marked with brownish-red. Vertex with two longitudinal narrow vittae, in winter specimens this may be one bar. Pronotum with heavy median Y-shaped spot with large irregular-shaped spot back of each eye. Scutellum with basal angles reddish. Clavus reddish-brown except small oval spot at tip of scutellum and another smaller one two thirds distance to apex. Corium with zigzag vitta arising on costa at anterior end of costal plaque, widening to broadly touch claval suture near apex of anchor spot and ending in cross veins near base of cell M^. Cross veins bright red. Apical cells more or less dusky. Venter dark. Genitalia. Pygofer hook U-shaped with three processes in grad- uated series, outer the shortest. Style of foot small; heel evident; base curved; anterior point about one third as long as toe is wide, about as wide at base as length; posterior point just about a right angle, scarcely evident. Aedeagus in dorsoventral view spatulate; processes shorter than shaft, apices sharp, diverging with recurved hook on outer margin. Holotype, male, allotype female, 1 male, and 6 female paratypes, Bretton Woods, N. H., August 31, 1934, R. H. Beamer. In Snow Entomological Collection. 28. Erythroneura nudata McAtee Erythroneura comes var. nudata McAtee, W. L., Trans. Am. Ent. .Soc, XLVI, p. 316, 1920. Erythroneura attenuata Johnson, Dorothy M., Ohio Jr. Sci., vol. XXXIV, No. 4, July, 1934. Original description. "Color of scutellum and anterior parts pale yellowish with irregular pellucid areas, instead of the color vittae and spots usually present in other varieties; tegmina whitish hya- line, apical cells somewhat fumose; costal placjue rather opaque white, an oblique black streak across it posteriorly, and black dots in bases of fourth apical, and apices of second apical cells. Placed as a variety of comes because of the presence of dark spots in apices of second apical cells, altho so far as coloration goes no intergrading specimens have been seen." This species was described from teneral specimens and has only been recognizable through the internal male genitalia. General ground color yellowish-white marked with red or orange. Vertex with inverted V quite narrow, continued across pronotum as a thin Y. Lateral margins of pronotum with a narrow vitta. Beamer: Species of Erythroneura 285 Scutellum with basal angles yellow, often bordered with red, tip orange. Elytra with typical orange markings of E. comes. Apical cells more or less infuscated. Black line in posterior end of costal plaque, spot in base of cell M^ and apex of cell R3. Genitalia. Pygofer hook U-shaped, inner process more than twice as long as outer. Foot of style large; heel medium; base straight; anterior point about as long as shortest width of foot; posterior point much longer than foot, about as wide at base as foot at narrowest place. Aedeagus in dorsoventral view quite broad, bell- shaped, processes about as long as shaft, heavy, slightly divergent. This species is close to E. beameri Rob. but may be separated from it by the broad bell-shaped aedeagus and heavier, straighter processes. A common species throughout the eastern half of the United States, 29. Erythroneura beameri Robinson Erythroneura beameri Rwhinson, Win., Can. Ent., vol. LVI, No. 3, p. 61; 1924. Resembles Erythroneura vitifex Fitch, but may be separated by its light mesosternum, and by the very long posterior point of style. A very common species in the United States east of the Rocky Mountains. 30. Erythroneura vaga Johnson Erythroneura vage Johnson, Dorotliy M., Ohio Jr. Sci., No. 4, p. 260, 1934. This species is close to E. comes (Say) or E. beameri Rob., but may usually be separated from those species by the extra lateral orange marks on the vertex and quite easily by the long slender shaft of aedeagus with bifid tip. Named originally from two specimens. Numerous specimens are at hand from Louisiana, Arkansas, Illinois and Kansas. Unlike the drawing of the type most specimens have an addi- tional spine beneath the base of the aedeagal shaft. 31. Erythroneura ancora Beamer Erythroneura ancora Beamer, R. H., Ann. Ent. Soc. Am., vol. XXII, p. 122 ; 1929. This species is near E. omaska Rob., but may be separated by color on vertex confined to inverted V, dark marking near cross veins confined to black spot in base of cell M4 and aedeagus without divided tip. It has not been taken since the type series from Cul- berson county, Texa.*. 286 The University Science Bulletin 32. Erythroneura omaska Robinson Erythroneura omaska Robinson, Wni., Can. Ent., vol. LVI, No. 3, p. 61 ; 1924. This species is often milky white as in E. delicata McAtee and may be confused with this species, but is usually easily separated by the broader pink or orange markings. Genitalia. Pygofer hook U-shaped, both arms practically same size. Foot of style small; heel prominent; base curved; anterior point almost as long as width of foot at narrowest place; posterior point about half as long. Aedeagus long and slender, bifid on outer fifth, pair of slender lateral processes near middle, in lateral view bent dorsally on apical fifth. This is a very common species throughout the United States east of the Rocky Mountains. Specimens are at hand from Kansas, Illinois, Arkansas and Louisiana. 33. Erythroneura delicata McAtee Erythroneura comes var. delicata McAtee, W. L., Trans. Am. Ent. Soc, XLVI, p. 317, 1920. Erythroneura lacta Robinson, Wm., Can. Ent., LVI, No. 3, p. 62; 1924. Original description. "Ground color milky white, with red color markings of the com,es type, the spots usually small and well sep- arated." This beautiful species is easily distinguished by its milk-white general color with delicate pink markings. Poorly marked speci- mens of E. omaska Rob. might be confused with it but may be sep- arated by the male genitalia and much broader pink markings. Vertex with thin inverted pink U-shaped mark with curving sides. Pronotum with narrow median Y-shaped spot and small dash on each lateral margin pink. Scutellum with basal angles yellow, tip pink. Clavus with anchor-shaped basal spot broken into a small basal dash and larger apical portion, another small dash near apex. Corium with oblique dash near base, a vitta arising on costa at anterior end of costal plaque, following its inner margin with off- shoot to meet claval suture near its middle, angling to end in base of cell M^. Cross veins pink. Apical cells more or less infuscated. Venter yellowish white with mesosternum and last ventral segment of male fuscous. Genitalia. Pygofer hook U-shaped, inner arm more slender and about one third longer than outer. Foot of style small ; heel small ; base curved; anterior point medium sized, outer face convex; pos- terior point very small, one third as wide as anterior and very short Beamer: Species of Erythroneura 287 and sharp. Aedeagus in dorsoventral view, for the most part broad, canal produced in middle, lateral margins with sharp, forward pro- jections about one third distance from apex, usual processes almost parallel and not reaching to apex. Holotype female, Plummer's Island, Md., July 26, 1914; W. L. McAtee. Since no allotype was made a male paratype, described above, Plummer's Island, Md., Oct. 26, 1913, W. L. McAtee, is so designated. Types in collection of W. L. McAtee. This beautiful milk-white species is very common in the United States east of the Rocky Mountains. Erythroneura delicata var. accepta McAtee Erythroneura comes var, accepta McAtee, W. L., Trans. Am. Ent. Soc, XLVI, p. 317 ; 1920. ErythroTieura scripta Robinson, Wm., Can. Ent., vol. LVI, No. 12, p. 290; 1924. Erythroneura tudella Robinson, Wm., Can. Ent., vol. LVI, No. 12, p. 291; 1924. Like Erythroneura delicata McAtee, but with a dark mark in the orange spot near middle of clavus. It occurs over the same range but is not so common. 84. Erythroneura corni Robinson Erythroneura conn Robinson, Wm., Can. Ent., Vol. LVI, No. 3, p. 60 ; March 1924. Erythroneura omata Osborn, H., Ohio Biol. Surv. Bui., XIV, p. 364, 1928. This is the common species on cornus in central and southern United States. It may usually be distinguished by the red marks of vertex and pronotum being suffused with fuscous and by the rather indefiniteness of the markings in general. The slender aedea- gus in ventral view, the more definite points on the foot of style and the forks of the pygofer hook touching or almost touching are also characteristic. A very common species east of Rocky Mountains. 35. Erythroneura bidens McAtee Erythroneura comes var. bidens McAtee, W. L., Fla. Ent., vol. VIII, p. 39; December, 1924. Erythrone%ira comes var. suffusa McAtee, W. L., Fla. Ent., vol. VIII, December 24, 1924, p. 89. Original description. "Like E. comes var. rubra Gillette, except that the upper surface of pronotum save a semielliptical space on each anterior angle, and a semicircular spot in middle of front margin, is black. Length, 2.8 mm. Holotype male, Virginia near Stubbleficld Fall, on Finns virginiana, October 23, 1921, J. R. Mal- loch." 288 The University Science Bulletin Specimens from Glen Echo, Mel, and others near Washington, D. C, collected by P. W. Oman, on comparison with the holotype, a female instead of a male, as the original description states, are unquestionably this species. General ground color semihyaline to whitish with orange and fuscous marks. Vertex red, with a pair of mesal longitudinal white vittae and an additional row of white spots on each side. Pronotum usually fuscous, but sometimes orange except three white spots on anterior margin. Scutellum orange to fuscous. Clavus with anchor-shaped mark and another elongated area before tip orange. Corium with irregular-sided vitta on costa, surrounding costal plaque, approaching claval suture at three points and touching cross veins on costal and mesal margin. Apical cells dark. Venter dark. Legs lighter. Genitalia. Py gofer hook large, long side with tip enlarged and bent out at about right angles, about twice as long as short side. Style with medium foot, medium heel and rather short anterior and posterior point. Aedeagus with shaft narrow, rather long, par- allel-sided, in lateral view bent dorsally on outer third. This species is distinguished from the others found on Cornus sp. by the thickened outwardly angled long point of pygofer hook. The dark pronotum will distinguish most specimens of bidens, but I have a few from Illinois that lack this color. 36. Erythroneura rubrella McAtee Erythroneiira comes var. rubrella McAtee, W. L., Trans. Am. Ent. Soc, XLVI, p. 316, Aug. 26, 1920. General ground color milky white with the dark body color vis- ible; markings bright red. Vertex red except two median longitu- dinal vittae separated by narrow red line, two round spots near each eye and marginal band around tip of vertex which are yellowish white. Pronotum with broad Y-shaped median mark and marginal band red. Scutellum with basal angles yellow, remainder red except inverted white T-shaped mark. Clavus with usual red basal anchor- shaped mark and another elongated spot near apex. Corium with red irregular sided vittae arising near base more or less surrounding plaque, ending in cross veins near tip of Cu. Cross veins slightly brighter red. Apical cells more or less infuscated. Black spot in posterior end of costal plaque. Venter dark. Genitalia. Pygofer hook double; inner fork heavy at base, twice as long as outer, curved out on outer half at right angles to basal ; outer fork narrower, leaving base at an angle, forming gentle curve Beamer: Species of Erythroneura 289 toward inner fork. Style with median foot; toe broad; anterior and posterior points usually small but not so well marked as in E. corni Rob. Aedeagus in dorsal view short and thick, half as broad as long, evenly narrowed on outer half to sharp tip, quite spinose; in lateral view short and thick, tip slightly curved dorsally. Type dissected and numerous specimens studied from British Columbia, Arkansas, Mississippi, Illinois. This is structurally very distinct from all the other forms. The very broad aedeagus is quite different from the long slender struc- ture found in the others. 37. Erythroneura ontari Robinson Erythroneura ontari Robinson, Wm., Can. Ent., vol. LVI, p. 60, March, 1924. This species resembles E. corni Rob., but may usually be dis- tinguished by the narrower markings on vertex and pronotum which lack the fuscous suffusion. The pronotal Y-shaped mark is usually more evident. The pygofer hook will separate it from the other species except E. rubrella, from which it can be distinguished by the very narrow aedeagus in dorsal view. This is a widespread species both in the United States and Canada. 38. Erythroneura aza Robinson Erythroneura aza Robinson, Wm., Can. Ent., vol. 56, No. 12, p. 291; 1924. Original description. "Ground color of vertex, pronotum and scu- tellum ivory, of tcgmina white, with following marks in orange; vertex, a narrow median inverted U-shaped vitta; pronotum, a broad median Y-shaped vitta and a broad lateral vitta behind each eye; scutellum, a broad spot in each basal angle and a small spot at tip. Tegmina lightly washed with orange throughout except at inner basal angle of clavus and in base of corium, where the orange is deepened to nearly red. Cross veins and adjacent parts of longi- tudinal veins crimson. A black streak caudad of costal plaque; apical cells whitish at base, remainder dusky, especially near the base. A black spot in apex of cells R3 and base of cell M^." This species resembles E. delicata McAtee, but may be separated from that species by a more hyaline appearance, the red or orange markings larger and heavier and the posterior point of style half as long as base. Genitalia. Pygofer hook typically U-shaped. Foot of style medium sized; heel medium; base straight; anterior point about as long as width of foot at narrowest place, outer face convex, rather wide at base; posterior point sharp and narrow, about half as long 19^6037 290 The University Science Bulletin as base. Aedeagus in dorsoventral view almost three pronged, two laterals almost retrorse at tip. Known from the types from Lawrence, Kansas, and three addi- tional male specimens, one each from Douglas county and Leaven- worth county, Kansas, and one from Natchitoches, La. 39. Erythroneura rubra (Gill) Typhlocyba comes var. rubra Gillette, C. P., Am. Typholocybinae, 1898, p. 764. Erythroneura irrorata Robinson, Wm., Can. Ent., vol. LVI, No. 7, p. 154; 1924. This red-marked species resembles the dogwood series and is often found confused with them in collections. It may easily be separated from them by always having the mesal longitudinal line of vertex light colored instead of dark as in the dogwood series. Widely distributed throughout the eastern two thirds of the United States and Canada. Type in the United States National Museum. 40. Erythroneura festiva n. sp. Resembling E. rubra Gill., but can be separated by the smaller size, the much narrower color marks, by anterior point of style as long as foot and aedeagus in dorsoventral view quadrangular, tube not protruding. Ground color semihyaline to whitish with coppery colored marks more or less clouded with fumose. Vertex copper colored except mesal, longitudinal keyhole-shaped spot, circular spot either side of this near base and a narrow spot on lateral margin between apex and eye. Pronotum with median Y-shaped spot with very short base, and two lateral stripes coppery. Scutellum red, except basal angles yellow and mesal white bar with end knobbed near apex; clavus with basal anchor-shaped mark and apical spot red. Co- rium with red basal dash and irregular-sided mark surrounding costal plaque, touching claval suture in two places and ending at cross veins near base of cell M^. Costal plaque with black basal dash. Apical cells more or less infuscated. Veins whitish. Venter quite dark. Genitalia. Foot of style small; posterior point as long as base; anterior short and sharp. Aedeagus in ventral view almost rec- tangular, apically incised either side of orifice with pair of ventral processes extending slightly beyond apex. Pygofer hook of usual comes type. Holotype male and allotype female, White county, Illinois, March 31, 1929, R. H. Beamer. Beamer: Species of Erythroneura 291 Paratypes as follows: 11 males and 3 females, Colfax, La., De- cember 23, 1931, R. H. Beamer; 1 male and 1 female, Clearwater, La., December 21, 1931, R. H. Beamer; 4 males and 2 females, Cherokee county, Kansas, December 17, 1930, R. H. Beamer; 1, Fouke, Ark., December 21, 1931, R. H. Beamer; 1 female, Coal Creek, Tenn., August 27, 1930, R. H. Beamer; numerous males and females from White, Clay, and Gallatin counties, Illinois, collected in March, 1929, by P. W. Oman and R. H. Beamer. 41. Erythroneura compta McAtee Erythroneura comes var. compta McAtee, W. L., Trans. Am. Ent. Soc, XLVI, p. 318; 1920. Erythroneura cherokee Robinson, Wm., Canl. Ent., vol. LVI, p. 154; 1924. Eryihroixeura nigroscuta Johnson, Dorothy M., Ohio Jr. of Sci., vol. XXXIV, No. 4, p. 258; 1934. Resembling E. delicata var. accepta McAtee, but pink markings heavier, white background not so striking, scutellum usually with dark markings, posterior point of style longer than foot and proc- esses of aedeagus bifid. General ground color yellowish-white. Red inverted V of vertex rather narrow, branched even with anterior edge of eye, encircling light spots between eyes and main arms of V, continued across pro- notum to unite on posterior margin. Pronotum with an additional red strip on lateral margin. Scutellum mostly dark, extreme lateral angles and spot near center light. Clavus with usual anchor-shaped spot at base and smaller one at tip red, basal spot more or less blackened at tip. Corium with red oblique dash on costa midway to costal plaque, another red stripe arising on costa at anterior end of plaque, almost surrounding it, connected to the claval suture between its two spots and ending in the base of cell M4. Cross veins red, margined on outside with hyaline, which is in turn mar- gined with fuscous. Black spot in posterior margin of costal plaque and apex of cell R3. Ventral surface stramineous more or less suf- fused with pink. Genitalia. Pygofer hook U-shaped with outer fork about same thickness as inner but about a third shorter. Foot of style of me- dium size; heel prominent; base almost straight; posterior point about one half as long again as base of foot; anterior point short and sharp, about half as long as toe is wide. Aedeagus in dorso- ventral view bell-shaped, processes bifid almost entire length, reach- ing to tip of shaft. Tijpe. Female, Plummer's Island, Md., March 28, 1915. For some reason McAtee did not name an allotype, therefore the male 292 The University Science Bulletin paratype, Plummer's Island, Md., Dec. 21, 1915, described above, is so designated. This is a very abundant species in the United States on grape east of the Rocky Mountains. Erythroneura compta var. rujomaculata McAtee Erythroneura comes var. rufomacidtata McAtee, W. L., Bui. 111. Nat. Hist. Sur., April, 1924, p. 43. Dissections of the male genitalia of this form reveal no differences from E. compta. Summer specimens of the right age usually have the red marking as indicated in the original description. 42. Erythroneura comes (Say) Tettigonia comes Say, Thomas, Ins. Jour. Ac. Nat. Sci. Phila., LV, p. 343; 1825. General ground color yellowish-white, marked with orange. Ver- tex with inverted U-shaped mark with lateral extensions enclosing white spots next each eye. U mark continued on pronotum as Y- shaped vitta reaching posterior margin, an additional vitta on each lateral margin. Scutellum with basal angles yellow often bordered with orange, tip orange. Clavus with usual basal anchor-shaped spot and a much smaller one at tip. Corium with angular dash on costal margin near base, another vitta arising on costa at anterior end of costal plaque enlarging to meet claval suture near its middle, almost surrounding plaque, then angling to end in base of cell M^. Cross veins usually pink, apical cells more or less infuscated. Black spots in posterior end of costal plaque, base of cell M^ and in apex of R3. Genitalia. Pygofer hook of the U type, outer process about two thirds as long as inner. Style with large foot; heel marked; base straight; anterior point short, fairly sharp; posterior point longer than foot, sides almost parallel, usually slightly sinuate. Aedeagus in dorsoventral view oval, very broad, apex almost straight; proc- esses longer than shaft, diverging on outer third with a distinct notch or tooth on their margin near middle. This notch is the distinctive character of this species. Since the types of this species are lost the male specimen described above, Leavenworth county, Kansas, April 11, 1930, is designated neotype. This is a very common species on grape throughout the United States east of the Rocky Mountains. Beamer: Species of Erythroneura 293 43. Erythroneura tacita n. sp. Resembling Erythroneura nudata McA. in the narrow markings, but larger; posterior point of style almost absent; pygofer hook single and aedeagus very long and narrow with bifid tip. General ground color whitish, marked with orange. Vertex with median longitudinal band often enclosing two long white areas, con- tinued across pronotum enclosing a white arc on anterior margin. Pronotum with lateral orange bar not reaching posterior margin. Clavus with usual anchor-shaped mark and small mesal spot just before tip. Corium with orange dash at base, another surrounding costal plaque connected to claval suture slightly beyond middle and ending on Cu before cross veins. Cross veins red. Black spot in posterior end of costal plaque and another at base of cell M^. Venter stramineous, mesosternum dark. Genitalia. Pygofer hook single, U-shaped, heavy at base. Foot of medium size; almost no heel or posterior point; anterior point long, less than right angle. Aedeagus in ventral view with flattened bifid tip, pair of heavy diverging processes at base, and between basal processes and shaft proper a curved, flattened, enlarged area. Holotype, male, allotype, female, 15 male and 10 female para- types. Republic, Wash., August 6, 1931, R. H. Beamer; 6 female paratypes, Oliver, B. C, August 6, 1931, R. H. Beamer; 1 male, Meritt, B. C, and 1 male, Dixie, Ore.; 17 females and 11 males, Poudre river, Colo., Aug. 22, 1931, R. H. Beamer. The specimens were swept from Comus sp. 44. Erythroneura gilensis Beamer Erythroneura gilensis Beamer, R. H., Annals Ent. Soc. Am., vol. 22, p. 123, 1929. This species is very close to E. beameri Rob., but may be sepa- rated from it by its larger size, by narrower, more broken markings and by the processes of aedeagus not diverging at so great an angle. A fairly common species on grape in Arizona. Erythroneura octonotata Walsh Erythroneura octonotata Walsh, B. D., Prair. Farm., Sept. 6, 1862, p. 149. The original description follows: "Whitish. Head as in the pre- ceding, {ziczac.) Thorax a little clouded with fuscous. Abdom- inal and ventral joints, in mature specimens dusky except at tip. Tips of tarsi dusky. Elytra whitish subhyaline, with the same three spots as the preceding, and in addition, one on the inner 294 The University Science Bulletin margin not far from the base; on the cross veins an irregular fus- cous band, and on the disk a small brown cloud, often obsolete. Length, a little over one tenth inch." I have not been able to place the species that Walsh must have had when he wrote the above description. It fits almost equally well either of the two species which are known in this paper as Erythroneura compta McAtee and E. delicata var. accepta McAtee, but in both of these there are always orange marks on the tegmen and in the first, black marks on the scutellum. When one considers the description of the color of E. ziczac which immediately pre- cedes that of E. octonotata, I feel convinced that he would not have omitted these marks. Index to Species accepta 287 aclys 274 Bcuticephala 272 amanda 265 ancora 285 anfracta 280 attenuata 284 aza 289 beameri 285 bidens 287 bistrata 267 breakeyi 279 caetra 281 calycula 270 cancellata 273 Cherokee 291 coloradensis 277 comes 292 complementa 269 compta 291 corni 287 corona 267 cymbium 271 delicata 286 disjuncta 272 diva 269 elegans 273 elegantula 277 erasa 270 festiva 290 flava 266 fraxa 272 gilensis 293 infuscata 265 irrorata 290 integra 271 kanwakae 279 lacta 286 nigroscuta 291 noncincta 270 nudata 284 octonotata 293 omaska 286 ontari 289 omata 287 palimpsesta 278 pontifex 279 portea 283 prima 283 prosata 275 reflecta 283 repetita 282 rosa 282 rubra 290 rubranotum 268 rubravittata 269 rubrella 288 ruf omaculata 292 scripta 287 stricta 268 suffusa 287 tacita 293 tricincta 268 tudella 287 vaga 285 venusta 266 vitifex 280 vitis 266 walshi 276 ziczac 275 296 The University Science Bulletin PLATE XVII Wing venation of the four groups of Erythroneura. Beamer: Species of Erythroneura 297 VULNERATA GROUP OBLIQUA GROUP MACULATA GROUP COMES GROUP 298 The University Science Bulletin PLATE XVIII 1. E. infuscata (Gill.) Dorsoventral view of aedeagus, foot of style, pygofer hook and lateral view of aedeagus. 2. E. amanda. 3. E. vilis Harris. 4. E. bistrata McAtee. 5. E. tricincta Fitch. 6. E. diva McAtee. 7. E. calycula McAtee. 8. E. Integra McAtee. 9. E. cymhium McAtee. Beamer: Species of Erythroneura 299 PLATE XVIII 300 The University Science Bulletin PLATE XIX 10. E. jraxa Robinson. Dorsoventral view of aedeagus, foot of style and pygofer hook. Lateral view of aedeagus. 11. E. acuticephala Robinson. 12. E. elegans McAtee. 13. E. cancellata McAtee. 14. E. aclys McAtee. 15. E. prosata Johnson. 16. E. ziczac Walsh. 17. E. elegantula Osborn. 18. E. coloradensis (Gill.). Beamer: Species of Erythroneura 301 PLATE XIX 302 The University Science Bulletin PLATE XX 19. E. palimpsesta McAtee. Dorsolateral view of aedeagus, foot of style and pygofer hook. Lateral view of aedeagus. 20. E. pontijex McAtee. 21. E. kanwakae Robinson. 22. E. anfracta Beamer. 23. E. vitifex Fitch. 24. E. caetra McAtee. 25. E. rosa Robinson. 26. E. reflecta McAtee. 27. E. prima n. sp. Beamer: Species of Erythroneura 303 PLATE XX 304 The University Science Bulletin PLATE XXI 28. E. nudata McAtee. Dorsolateral view of aedeagus, foot of style and pygofer hook. Lateral view of aedeagus. 29. E. beameri Robinson. 30. E. vaga Johnson. 31. E. ancora Beamer. 32. E. omaska Robinson. 33. E. delicata McAtee. 34. E. comi Robinson. 35. E. bidens McAtee. Beamer: Species of Erythroneura 305 PLATE XXI 28 E NUDATA 29 E BEAMERI 30 E VAGA 31 E ANCORA 32 E OMASKA m-^ M-C^S 33 E DEUCATA 34 E. CORNI 35 E BIDENS 20—6037 306 The University Science Bulletin PLATE XXII 36. E. rubrella McAtee. Dorsolateral view of aedeagus, foot of style and pygofer hook. Lateral view of aedeagus. 37. E. ontan Robinson. 38. E. aza Robinson. 39. E. rubra (Gill.). 40. E. festiva n. sp. 41. E. compta McAtee. 42. E. comes (Say). 43. E. gilensis Beamer. '44. E. tacita n. sp. Beamer: Species of Erythroneura 307 PLATE XXII 44 E TACIT A THE UNIVERSITY OP KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 15. Alconeura of the United States (Homoptera, Cicadellidae) MELVIN E. GRIFFITH Department of Entomology, University of Kansas CONTENTS PAGE Generic distinctions 310 Equipment and method 311 Key to species 317 Descriptions 317 1. A. planata Ball and DeLong 318 2. A. necojnnata, n. sp 318 3. A. beameri, n. sp 319 4. A. balli Beamer 319 5. A. tricolor (Van Duzee) 320 6. A. jidminea Lawson 321 7. A. dorsalis (DeLong) 321 8. A. unipuncta (Gillette) 322 9. A. quadrimaculata Lawson 324 10. A. derecta, n. sp 325 11. A. insulae, n. sp 325 12. A. dodonana Beamer 326 13. A. lappa, n. sp 327 14. A. macra, n. sp 327 15. A. rotundata Ball and DeLong 328 16. A. nudata Ball and DeLong 329 17. A. languida, n. sp 330 18. A. luculenta, n. sp 331 19. A. cornigera, n. sp 332 Alconeura quadrivittata (Gillette) 332 Literature cited -^33 Abstract: The Alconeura group, separated from the Dikraneura by Ball and DeLong (1925), retains its generic rank with the addition of nine new species. The numerous specimens studied were collected chiefly from the (309) 310 The University Science Bulletin southern and western regions of the United States. Specific distinctions are primarily drawn from characters of the dissected male genitalia, and the technique for preparation of these structures is given in detail. A key to the nineteen species described from the United States is provided, and complete descriptions with locality notes are presented for them: Alconeura planata Ball and DeLong (p. 318), A. necojiinata, n. sp. (p. 318), A. beameri, n. sp. (p. 319), A. balli Beamer (p. 319), A. tricolor (Van Duzee) (p. 320), A. jvlminea Lawson (p. 321), A. dorsalis (DeLong) (p. 321), A. unipuncta (Gillette) (p. 322), A. quadrimaculata Lawson (p. 324), A. derecta, n. sp. (p. 325), A. insidae, n. sp. (p. 325), A. dodonana Beamer (p. 326), A. lappa, n. sp. (p. 327), A. maa-a, n. sp. (p. 327), A. rotundata Ball and DeLong (genotype) (p. 328), A. nudata Ball and DeLong {^unipuncta var. nudata) (p. 329), A. languida, n. sp. (p. 330), A. luculenta, n. sp. (p. 331), A. cornigera, n. sp. (p. 332). A note on A. quadrivittata (Gillette) from the Bahamas is added, and the literature of the genus is given. Illustrations are of the aedeagus and pygofer process of each species, and two typical forewing tips. Types of new species are in the Francis Huntington Snow Entomological Collections at the University of Kansas. GENERIC DISTINCTIONS THE Alconeura group was first generically distinguished from the Dikraneura in 1925 by Ball and DeLong, who considered an angulate or pedunculate second apical cell determinative, overlook- ing, as McAtee (1926) suggests, the extreme variability of this character. However, the insufficiency of the first distinction should not cause a too hasty rejection of the generic significance of the group, since there are plain indications that the present assembly of species can be recognized most conveniently as a separate genus. It should be stated at once that the pedunculate apical cell is dis- tinctly a characteristic of the Alconeura, and while it may appear exceptionally among the Dikraneura, it is never as admirably de- veloped. In none of our species of Alconeura is the apical cell simply angulate. The most obvious individualizing mark of the genus is the dark apical dot which appears in the third or fourth cell and frequently touches or lies upon the third vein. This may also be observed in the Dikraneura, but there it is not accompanied by the pedunculate apical cell. There is evidence that the dissected male genitalia msiy be dis- criminative in the two genera, but this is problematical, since the forms of genitalia are highly specific and vary a great deal within a genus. Certainly the clear division of Alconeura and Dikraneura by external properties is dissuasion enough from the tedious under- taking of logically gathering in two major classes these less acces- sible structures. Griffith: Alconeura of the U. S. 311 Nineteen species of the Alconeura have been distinguished, hav- ing in common these characteristics: a marginal vein on the under- wing, with two closed cells at the apex; four apical cells on the forewing, the second cell pedunculate; a dark apical dot in the third or fourth cell, often touching or lying upon the third vein; the ver- texal angle but slightly obtuse or acute; a length of 2.5 mm. to 3.5 mm. ; and, usually, bright coloration. Alconeura rotundata was de- scribed as the genotype by Ball and DeLong; this species displays very well the characters enumerated above and, indeed, its orange- yellow markings on a creamy-white background are typical of the major part of the genus. The notable addition of nine new species to a genus in which but ten were properly recognized in 1934 may be ascribed to the good fortune of having available a wondrous amount of material collected by Dr. R. H. Beamer and others, and type specimens of all the de- scribed species. Sincere appreciation for their kindness and co- operation must be expressed to Dr. E. D. Ball, University of Ari- zona; Dr. D. M. DeLong, Ohio State University; Mr. P. W. Oman, United States National Museum; and particularly to Dr. R. H. Beamer, University of Kansas. EQUIPMENT AND METHOD This study of the Alconeura was undertaken with the definite pur- pose of establishing previously described species and distinguishing new species by means of the dissected male genitalia, the impetus for this taxonomic endeavor coming directly from antecedent studies of the Erythroneura. The technique for dissection and study of the genital structures is necessarily given here in some detail, and may be esteemed a culmination in method, for literally hundreds of speci- mens were dissected, with many complete sacrifices, in its perfection. It is a sound principle for a worker to know well his tools, and this acquaintance is best made before the work is begun. The list following will be found helpful in arranging the equipment in an orderly outlay, which must be regarded as essential to the surest and quickest attack on any sort of problem in anatomizing. The articles are presented approximately in order of employment: A dissecting microscope. A thick slice of art eraser, holed to admit the head of the insect pin. A small dissecting needle with a sharp, flattened tip for removing the insect abdomen. Dissection number labels for the pinned specimens, giving the generic name, number of the dissection, and the name of the dissector. 312 The University Science Bulletin A tiny wire loop provided with a convenient handle for hfting the abdo- men into and from the solutions. Two small porcelain casseroles: one to hold caustic potash (10% solution) and the other distilled water. An alcohol lamp and stand for heating the caustic potash. Glycerine : the medium within which the dissection is made. Microscope slides. A small linen towel for wiping off the slides. A soft wax pencil for noting temporarily upon the slide the dissection number. A binocular microscope provided with dissecting stage and hand rests. Two very fine needles (minuten nadeln) ; one straight, one bent into a right angle, provided with convenient handles. Microscope object labels. De Faure's mounting fluid. (Gum arable, 30 grms.; chloral hydrate, 50 grms;. chlorhydrate of cocaine, 0.5 grm.; glycerine, 20 cc; distilled water, 50 cc.) This beautifully clear mounting fluid must be very carefully mixed and filtered. Any other satisfactory mounting fluid may be substituted. Cleaning fluid for slide cover glasses. (Hydrochloric acid, 1 cc; alcohol, 95%, 100 cc.) Slide cover glasses (circles, 15 mm.). A microscope with reasonably high magnification for examining the finer details of the dissected structures. Technique, in whatever manner described, must always be indi- vidual in application ; however, it should be remembered that noth- ing is more destructive to quick attainment of worthy results than a haphazard method. While the experience of a very few dissec- tions will best enable the worker to locate the equipment most ad- vantageously, it must be borne in mind from the first that a logical sequence in the placement of tools and a plan of action will un- doubtedly secure the happiest conclusion of the process in the shortest time. Accuracy and a certain finesse are most to be de- sired; yet it is undeniably heartening to feel that more time may be given to studying the dissections than to the routine of making them. The systematic approach suggested here will be found im- perfect, but it has made possible an average of twenty dissections an hour and may prove helpful to those who wish to determine posi- tively large series of the Alconeura or other Eupterygine leaf hoppers. 1. Removing the Abdomen Fill a porcelain casserole with caustic potash and place it upon a stand over the lighted alcohol lamp, adjusting the wick so that the solution does not bubble vigorously. As the heated solution Griffith: Alconeura of the U. S. 313 evaporates during a series of dissections, replenish alternately with distilled water and caustic potash. Fill a casserole with distilled water and place near the alcohol lamp. Place the art eraser centrally on the dissecting microscope stage. A binocular microscope may be used, but the dissecting microscope is preferable since, after removal of the abdomen, the lens arm may be swung away, leaving a clear field for operations with the wire loop and for removing the specimen. Invert the pinned insect upon the art eraser; a plain view is, of course, necessary and interfering labels on the pin must be turned slightly aside. Remove the abdomen, observing through the lens, with the flat- tipped needle. This instrument is best held like a pen and the abdomen should be removed by gently thrusting the tiny chisel point between abdomen and thorax. Breaking-off is most easily accomplished at this point; although, exceptionally, the genital cap- sule alone may be removed most satisfactorily. Withdraw the lens arm, and dipping the wire loop into the caustic potash, touch it to the abdomen, lifting it into the solution. 2. Labeling the Pinned Specimen Each dissected specimen must be numbered so that the perma- nent microscope slide mount of the genital structures may be definitely referred to it. This is best done with small regular-sized three-line labels upon which are printed the generic name and the name of the dissector, the middle line being left blank for writing in the dissection number, and it is best to use India ink for this purpose. 3. Preparing for Dissection Carefully clean a microscope slide and note the dissection num- ber across one end, using the wax pencil. Place a droplet of glycerine on the slide and center it upon the binocular microscope stage with an end toward the observer. This position allows working with the guiding fingertips close to the dis- section without troublesomely nudging the slide. Remove the abdomen from the caustic potash, in which it should have been heated from two to five minutes, using the wire loop, and submerge it in the distilled water. (At this point the abdomen 314 The University Science Bulletin of another specimen may be removed and transferred to the caustic potash.) Place the abdomen within the glycerine droplet, observing with the microscope. Use transmitted light. 4. Dissecting Take up the fine needles, the right-angle in the left and the straight-point in the right hand, grasping them, not as a pen is held, but with the handles resting lightly on the tips of the first two fingers and over the backs of the second pair. Using the handrest for the wrists, establish the fingers upon the stage and bring the needles together within the glycerine. Observing with the highest power lenses which permit a clarity of vision, turn the abdomen so that the genital capsule is normally oriented and directly centered in the field with the greater part of the abdomen extending toward the observer and with the pygofer processes visible on top. This is the proper position for dissection and should be reestablished after an unsuccessful trial. Insert the right-angle within the hollow of the capsule with the point toward the plates. Place the straight-point alternately upon the two plates, gently pulling the needles apart. Under optimum conditions this single operation will remove the aedeagus and the styles, if carefully done, to a point where they may be easily detached from the pygofers and the rest of the genital apparatus. The plates are then pulled away quite simply and the dissection is complete. In more un- toward circumstances it may be necessary to abandon the tugging at the plates and literally to pry out the aedeagus and styles with the straight-point while holding the capsule as before with the right- angle. The pygofers may be left intact on the capsule until per- manently mounted. The slide may now be set aside for study and the final mounting. (Here the next abdomen will be found ready for dissection; it should be dropped immediately into the distilled water and replaced in the caustic potash with another. This overlapping of processes so that a series may be quickly done can be easily mastered, but the first two or three dissections are best attempted singly.) The dissection may be preserved indefinitely in glycerine and studied most conveniently within that medium since it permits a free turning about of the structures. Griffith: Alconeura of the U. S. 315 5. Mounting Permanently Upon a gummed microscope object label note the dissection num- ber in the upper left-hand corner and the locality data of the pinned insect at the bottom. These notations should be made in India ink, and a series referable to specimens to be dissected can best be made all at one time. The center of the label, it will be seen, is left blank for penciling in the name of the insect or the dissection num- ber of a slide upon which the scientific name is given and which may typify the species. (For example, dissections in this study were assigned the key letters "GA" representing "Griffith Alconeura," and in most cases the permanent mounts bear, in addition to their proper corner labels, the cryptic middle signs "See GA 92, See GA 103;" etc., these references being to mounted dissections considered typical of their several species. And how much simpler it is to write Alconeura languida upon the one slide "GA 92," referring all subsequent dissections of the species to that number, than to note tediously the scientific name upon each mounted example.) Place the glycerine mount upon the binocular microscope stage and center it with one end of the slide toward the observer. Clean a slide and, attaching the label at one end, place it edge-to- edge with the glycerine mount. Drop a tiny bit of De Faure's mounting fluid in the center of the slide. Holding the needles as before and observing carefully through the lenses, lift up the aedeagus and styles on the right-angle. Thrust immediately into the De Faure's fluid and without removing the needle, move the slide to the center of the stage, thus pushing the glycerine mount slightly aside. Using a high power of the binocular lenses, arrange the genital structures with the aedeagus flat upon its side as in the included figures. Withdraw the needles carefully and dip into the hot caustic potash to clean. Using the tweezers, draw a cover glass from the cleaning fluid (which is best kept in a small, tightly covered dish) and dry upon the linen towel. Supporting the glass horizontally with the tweezers, touch a droplet of water to the underside, using the wire loop in the left hand. Gently lower the cover glass upon the slide until the drop of water merges with the mounting fluid, then release the glass immediately. (This bit of technique with cover glass and water 316 The University Science Bulletin drop obviates the usual difficulty with air bubbles.) Tap the cover glass with the tweezers, watching through the binocular, to insure complete spreading of the mounting fluid to the edge of the cover glass. Return the glycerine mount to its original central position, thrust- ing the permanent mount back. Place a small drop of De Faure's fluid upon the permanent prep- aration next the aedeageal mount and farthest from the label. Intro- duce the pygofers from the glycerine into this second droplet as before. Split the side of the genital capsule opposite that to which the pygofer processes are attached so that they may be pressed flat and still be in proper bilateral relation with each other. Apply the cover glass as before and press down tightly upon the flattened pygofers. Allow the prepared slides to dry for a day and then ring the mounts with clear lacquer to prevent deleterious evaporation. The dissected male genitalia structures are now permanently fixed in the best position for careful study and more powerful microscopes may be applied to them. With reasonable care in the use of the microscope, and the keys and figures given on following pages, posi- tive determination of described species of Alconeura should be ac- complished in a pleasingly swift fashion. Since all species are established determinatively in this paper by means of the dissected male genitalia, presentation of an elaborate key employing other characters would be worth neither the time necessary for its composition, nor, certainly, the time which might be given to its use. Indeed, the inclusion of figures which show quite clearly the plain differences exhibited in the various forms of genital structure make any involved key superfluous. However, a key to species, by whatever characters, is an effective way of pointing out descriminative points which might otherwise be overlooked, and the simple device which follows may be found work- able, though the figures, which have been arranged in a systematic manner considered most helpful, should be consulted often. The question of those troublesome female specimens collected without their attendant mates has not been unforeseen, but the de- scriptions of color patterns must be relied upon to place them as properly as can be until a larger series containing males may be col- lected. The task of correlating male and female genital characters is a very possible one, but it is much too extensive a piece of work to be finished and included in the present study. Griffith: Alconeura of the U. S. 317 Key to Species 1. Green, yellow, golden-brown, or orange-brown on white 2 Bright orange to yellow on white 9 2. (1) Green, yellow or golden-brown, apical dot in base of fourth cell 3 Orange-brown on white, apical dot in third cell 8 3. (2) Largely green, aedeagus with three slender processes arising in basal third... 4 Yellow or golden-brown, aedeagus without lanciform processes 7 4. (3) Green with red markings 5 Green without red markings 6 5. (4) Vertex to scutellum red tricolor (Van Duzee), 320 Vertex to scutellum green, apex of forewing flecked with pink balli Beamer, 319 6. (4) Distance from tip of median ventral process to margin of aedeageal shaft less than width of shaft planata Ball and DeLong, 318 Distance from tip of median ventral process to margin of aedeageal shaft greater than width of shaft necopinata, n. sp., 318 7. (8) Yellow, aedeagus diamond-shaped in ventral view beameri, n. sp., 319 Golden-brown, aedeagus not diamond-shaped in ventral view, .fulminea Lawson, 321 8. (2) Color on vertex, pronotum and scutellum fused ; pygofer process with small spur on outer margin dorsalis (DeLong), 321 Color on vertex, pronotum and scutellum in spots ; pygofer process without apical spur unipuncta (Gillette), 322 9. (1) Apical dot in base of fourth cell, often touching cross vein 10 Apical dot in third cell or upon third vein, removed from cross veins by more than its diameter 11 10. (9) Clavus with large stripe; wing-like short processes at tip of aedeageal shaft; pygofer process undivided at tip rotundata Ball and DeLong, 328 Clavus with thres spots ; without wing-like processes at tip of aedeageal shaft ; pygofer process divided at tip dodonana Beamer, 326 11. (9) Aedeageal shaft without important processes; pygofer process almost as long as aedeagus 12 Aedeageal shaft with important processes ; pygofer process much shorter than aedeagus 14 12. (11) Pygofer process double insulae, n. sp., 32,5 Pygofer process single 13 13. (12) Pygofer process scimitar-shaped quadrimaculata Lawson, 324 Pygofer process almost straight derecta, n. sp., 325 14. (11) Pygofer process well -developed 15 Pygofer process almost atrophied 17 15. (14) Aedeagus with pair of lateral processes at least half as long as shaft cornigera, n. sp., 332 Aedeagus with lateral processes less than half as long as shaft 16 16. (15) Basal processes of aedeagus more than five lappa, n. sp., 327 Basal processes of aedeagus less than five macra, n. sp., 327 17. (14) Tips of lateral processes on aedeagus bending dorsad languida, n. sp., 330 Tips of lateral processes on aedeagus bending ventrad 18 18. (17) Tip of aedeageal shaft bending ventrad nudata Ball and DeLong, 329 Tip of aedeageal shaft straight hiculenta, n. sp., 331 DESCRIPTIONS The problem of systematically presenting the species of the Al- coneura according to their external resemblances and the more im- portant affinities of the dissected male genitalia is, by no means, an easy one. However, the most cursory examination allows a division of the genus in two groups: Those first separated in the preceding key. Although the orange-yellow and white phases make up the greater part of the genus, the best introduction may be given with 318 The University Science Bulletin the other series which shows most clearly relationship with the Dikraneura. In descriptions of male genitalia the basal width of the aedeagus is measured from the dorsal margin immediately distad of the mem- branous tuft to the ventral margin just distad of the articulatory piece. This line of measurement is illustrated in the figure of Al- coneura balli Beamer. Quotations given are from the original descriptions, and types recorded, unless otherwise stated, are in the Francis Huntington Snow Entomological Collections at the University of Kansas. 1. Alconeura planata Ball and DeLong Alconeura planata Ball, E. D., and DeLong, D. M., Ann. Ent. Soc. Am., XVIII, 337, 1925. Resembling Dikraneura kunzei Gillette, but with pedunculate second apical cell and black dot in base of fourth apical cell. Color. Largely green. Vertex greenish-yellow with mesal white line often extending from apex to scutellum ; pronotum mostly green, with greenish-yellow anterior margin; scutellum greenish-yellow with white markings. Forewing light green with blue-green mot- tlings particularly evident along claval suture. Apical veins lighter. Venter fuscous ; face greenish-yellow near vertexal apex. Genitalia. Aedeagus in lateral view over twice as long as width at base; shaft curving dorsad, bent in basal fourth into right angle with base, narrowing gradually, tip about one fourth width of base, abruptly curving dorsad; three lanciform processes, two lateral and one ventromedian, arising in basal third and paralleling shaft to apical third, laterals slightly longer. Pygofer process almost as long as aedeagus, base heavy, bent near middle into right angle, tapering evenly to attenuated tip. "Described from eight examples taken by the senior author at Helper and Soldiers Summit, Utah, August 13, 1906. A remarkably distinct little species from the Wasatch Mountains.'^ Types studied: Allotype, male, and paratype, female (E. D. Ball Collection), 2. Alconeura necopinata, n. sp. Resembling Alconeura planata Ball and DeLong, but with white line on vertexal margin very distinct and ventromedian lanciform process on aedeagus, forming almost 45° angle with shaft rather than nearly paralleling it; pygofer process heavier and less radically bent. Color. Largely green. Vertex greenish-yellow, with white line on anterior margin joining eyes, and with mesal white line often ex- Griffith: Alconeura of the U. S. 319 tending to scutellum; pronotum mostly green, frequently with green- ish-yellow margins; scutellum variably tinged with yellow and green. Forewing yellowish-green, usually mottled sparsely in faint blue-green; apical dot in base of fourth apical cell. Tip of forewing lighter. Venter fuscous; face often lighter near vertexal apex. Genitalia. Aedeagus in lateral view about twice as long as width at base; shaft bending dorsad, apical portion at right angles with base, narrowing gradually, tip about one third width of base; three lanciform processes, two lateral and one ventromedian, arising in basal third and extending almost to tip of shaft, two laterals paral- leling, median forming almost 45° angle with shaft. Pygofer proc- ess about as long as aedeagus, base heavy, slightly curving and evenly tapering to point. Holotype, male; allotype, female; paratypes, twelve pairs: Mint Canyon, California, July 6, 1933, and July 31, 1935. Palmdale, California, July 6, 1933. R. H. Beamer. 3. Alconeura beameri, n. sp. Resembling Alconeura necopinata, n. sp., but yellow rather than green and aedeagus without lanciform processes. Color. Mostly yellow. Vertex with white line on anterior margin joining eyes, mesal white line often extending to scutellum; pronotum frequently white or lighter yellow anteriorly and at sides ; scutellum yellow or greenish, usually figured with white. Forewing yellow with white or lighter spots particularly evident along claval suture ; apical dot in base of fourth apical cell, infrequently very faint or apparently absent. Venter yellow; face usually with greenish tinge. Genitalia. Aedeagus in ventral view diamond-shaped, lateral ex- panded portions with minute conical teeth on ventral surface; in lat- eral view bending dorsad, apical portion of shaft approaching right angle with base, narrowing slightly, curving dorsad to blunt tip. Pygofer process slightly more than half length of aedeagus, sickle- shaped; apical two thirds slender, sharp-tipped, bending at right angles to heavy base. Holotype, male; allotype, female; paratypes, twelve pairs: Ja- cumba, California, August 12, 1935. Congress Junction, Arizona, August 13, 1935. R. H. Beamer. 4. Alconeura balli Beamer Alconeura balli Beamer, R. H., Can. Ent., LXVI, 17, 1934. "Resembles superficially Alconeura planata Ball and DeLong, but may easily be separated by the pink tips of the tegmen. 320 The University Science Bulletin "General ground color yellowish-green mottled with lighter. Head conical, quite pointed with two slightly darker yellow marks or blotches near base. Pronotum with anterior border whitish-yellow, lacking the mottling, remainder of pronotum mottled. Scutellum of about same color as anterior border of pronotum. Tegmen pale greenish-yellow mottled with white to cross veins. Usually pink to tip slightly suffused with fuscous. Black spot in base of cell M4. Venter dark throughout. Longitudinal veins white spotted, cross veins and apicals entirely white." Genitalia. Aedeagus in lateral view over twice as long as width at base; apical portion at right angle with base and half as wide; three lanciform processes, two lateral and one ventromedian, aris- ing in basal third and extending into apical third, paralleling line of shaft. Pygofer process about two thirds length of aedeagus, widest portion at base nearly one fourth length, bending abruptly in basal half and evenly tapering to point. Types studied: "Holotype, male and allotype, female, Santa Rita Mts., Arizona, July 17, 1932, R. H. Beamer. Numerous paratypes, same data." Ashfork, Arizona, August 8, 1932; Prescott, Arizona, August 7, 1932, and July 7, 1933; Granite Dell, Arizona, July 30, 1933; R. H. Beamer. 5. Alconeura tricolor (Van Duzee) Dicraneura tricolor Van Duzee, E. P., Trans. San Diego Soc. Nat. Hist., II, 56, 1914. Alconeura tricolor (Van Duzee), Ball, E. D., and DeLong, D. M., Ann. Ent. Soc. Am., XVIII, 336-337, 1925. Resembling Alconeura planata Ball and DeLong, but with vertex, pronotum and scutellum red. Color. Mostly red and green. Vertex, pronotum and scutellum bright red. Vertex with milky-white line on anterior margin and usually from apex mesally to scutellum; pronotal lateral margins touched with milky-white. Forewing green, often mottled with white along claval suture. Apical dot in base of fourth cell. Venter fuscous and red, appendages red; face with inverted V in milky- white formed by stripes extending mesad from antennal bases. Genitalia. Aedeagus in lateral view about twice as long as width at base; shaft bending dorsad, apical portion at 45° angle with base, narrowing in distal half to about one half width of base; three lanci- form processes, two lateral and one ventromedian, arising in basal third and extending past tip of shaft, two laterals curving ventrad but closely paralleling, median forming almost 45° angle with shaft. Griffith: Alconeura of the U. S. 321 Pygofer process nearly as long as aedeagus, slender, bent near mid- dle, very slightly tapered from base to point. "Described from thirteen examples representing both sexes, taken at Lakeside and Foster in May" (California). Types studied: A pair of homotypes (by R. H. Beamer), San Jacinto Mountains, California, June 30, 1933. Dulzura, California, August 9, 1935; Campo, California, August 10, 1935; Jacumba, Cali- fornia, August 12, 1935. R. H. Beamer. 6. Alconeura jubninca Lawson Alcotieura futminea Law.son, P. B., Bull. Brooklyn Ent. Soc, XX\', 44, 1930. Resembling Alconeura necopinata, n. sp., but with white line on vertexal margin continued across eyes, along lateral margins of pronotum and claval suture; aedeagus without lanciform processes. "Vertex, pronotum and scutellum rich golden brown. Vertex with narrow brown submarginal line and broad white marginal band which extends backward across eyes, sides of pronotum, and along length of claval suture. Elytra olive green with distinct black spot in fourth apical cell opposite third cross vein. Underside for the most part greenish yellow. Abdomen sometimes largely fuscous." Genitalia. Aedeagus in lateral view over twice as long as width at base, cur\-ing dorsad, with lines of dorsal and ventral margins nearly inirallel to apical third, which bends sharply dorsad, tapering abruptly to ])oint; dorsal and ventral margins in ventral view- slightly ex])an(k'd. Pygofer })rocess approximately one third longer than shaft of aedeagus, from middle evenly tapering to point. Types studied: Holotype, male; allotype, female; paratypes, two males and one female: Cameron county, Texas, August 3, 1928. Nogales, Arizona, June 25, 1933. R. H. Beamer. 7. Alconeura dorsalis (DeLong) Dikraneura unlpuncta dorsalis DeLong, D. M., Journ. N. Y. Ent. Soc, XXXII, 67-6S, 1924. Alconeura imipuncta dorsalis (DeLong), Ball, E. D., and DeLong, D. M., Ann. Ent. Soc. Am., XVIII, 330, 1925. Dikraneura {Alconeura) dorsalis DeLong, McAtee, W. L., Journ. N. Y. Ent. Soc, XXXIV, 1G4, 1926. Alconeura imipuncta dorsaVs (DeLong), Lawson, P. B., Bull. Brooklyn Ent. Soc, XXV, 46, 1930. Resembling Alconeura jidminea Lawson by white line margining dorsal longitudinal brownish band, but with apical dot in third, not fourth apical cell; pygofer process not as straight, curved into sickle form with marginal spur in distal half. 21—6037 322 The University Science Bulletin Color. Creamy-white with orange-brown markings. "Disc of vertex and pronotum, scutellum and cla\iis of elytra covered with a broad longitudinal brownish band which is bordered by a white band extending along margins of vertex from apex, across margins of pronotvun on to elytra at base and along claval suture half way to apex." Corium suffused semihyaline with orange-browai vitta curv- ing in from near humeral angle to margin apical half of claval suture; with darker line from midcostal margin to cross veins. Apical veins light with fumose margins; apical dot central in third apical cell. Tip of forewing fumose. Venter stramineous; face yel- low. Genitalia. Aedeagus in lateral view about twice as long as width at base, narrowing in apical third to about one half basal width, curving dorsad and gradually narrowing to rounded tip; line of ven- tral margin nearly semicircular; dorsal margin with distinct basal tooth. Pygofer process almost as long as aedeagus, about one fifth as wide at base, constricted in middle, widening apically and curving into sickle form with tooth on outer margin. "This is a very common variety found in the everglades in com- pany with unipuncta and is extremely abundant upon Trima jiori- dana in everglade hammocks wliicli are composed partially or en- tirely of this plant." Types studied: A pair of paratypes, Miami, Florida, April 13, 1921 (D. M. DeLong collection). Other records from Florida are: Key Largo, August 9, 1930, R. H. Beamer. Cocoanut Grove, August 9, 1930, L. D. Tuthill. Biscayne Bay, P. R. Uhler col- lection. 8. Alconcura unipuncta (Gillette) Dicraneura ■unipuncta Gillette, C. P., Proc. U. S. Nat. Mus., XX, 718, 1S98. Alconcura unipuncta (Gillette), Ball, E. D., and DeLong, D. M., Ann. Ent. Soc. Am., XVIII, 335-336, 1925. Dikraneura (Alconcura) unipuncta Gillette, McAtee, W. L., Jouin. N. Y. Ent. Soc, XXXIV, 163, 1920. Alconcura unipuncta (Gillette), Lawson, P. B., Bull. Brooklyn Ent. Soc, XXV, 4'), 1930. Resembling Alconeum dor.salis (DeLong), but without dorsal longitudinal brownish band; pygofer process without marginal spur in apical half. Color. Creamy-white with orange-brown markings. Vertex usu- ally with orange inverted V from apex to posterior margin; pro- notum most fre(|ucntly with four orange vittae, ends of lateral i)airs joined or more broadly fused; scutellum variably marked with yel- low to orange-brown. Forewing with two orange to orange-brown spots on clavus, one frecjuently almost filling i)asal portion, an- Griffith: Alconeura of the U. S. 323 other just past middle, normally forming one half large mid-dorsal spot apparent when wings are closed, tip of clavus fumose; corium with orange to orange-brown vitta almost paralleling claval suture from near humeral angle to tip of clavus; fumose orange stripe ex- tending in from midcostal margin to cross veins. Apical veins light with fumose borders; apical dot large, in third apical cell. Tip of forewing fumose. Venter stramineous, smokily suffused. (Jcnitalia. Aedeagus in lateral view about twice as long as width at base, narrowing in apical third to about one half basal width, curving dorsad to rounded tip; line of ventral margin nearly semi- circular; dorsal margin with stout basal tooth. Pygofer process almost as long as aedeagus, slender, constricted just proximad to apical third which curves abruptly to sharp point. "Described from three females and one male from the United States National Museum, one of which is labeled 'Cociuillet, Col- lector, Calif., through C. V. Riley,' and three are labeled 'Co- quillet, Los Angeles, Calif.' " Types studied: Holotype, male (Type 3414, U. S. N. M.), and a paratype, female, found to be Alconeura quadrimaculata Lawson, a much lighter species. This apparent confusion of the two species evidently accounts for the emphasis on orange coloration in Gil- lette's description of Alconeura vnipuncta, which is typically marked, in even darker brown than Alconeura dorsalis (DeLong). This species is very common and widely distributed, though typi- cally Californian. Records from that state are: San Diego county. July 4, 1929; Alpine, July 9. 1929; Mint Canyon, July 6, 1929: San Jacinto Mountains, July 21, 1929; Marin county, August 3, 1929; Winters, August 6, 1929 ; Strawberry, August 8, 1929 ; Boulder Creek, July 18, 1933; SaHnas, July 18, 1933; Gaviota, July 19, 1933; San Ardo, July 18, 1933; Nicolaus, June 27, 1935; Siskiyou, N. F., July 14, 1935; Nipomo, July 24. 1935; Claremont, July 29. 1935; Anza, August 6, 1935; R. H. Beamer. Giant Forest, July 28, 1929, L. D. Anderson. Van Duzee (1914) reports the species from Foster, Mus- sey's, and Alpine (San Diego county). , Ball and DeLong (1925) note that "specimens have been ex- amined from Grant's Pass, Oregon; San Francisco, Sacramento, and Pasadena, California (Ball); Ash Creek, Arizona; Utica, Missis- sippi; Alabama and Biscayne Bay, Florida, (U. S. N. M.) ; Miami, Florida, (DeLong)." ]\IcAtee (1926) records the species from Washington, D. C, July 2, 1913, W. L. IMcAtee; and Soledad, Cuba, IVLarch 14, 1925, J. G. Myers. 324 The University Science Bulletin A specimen was taken in Glendale, Nevada, January 3, 1930, by David E. Fox on Pluchea sericea, and a number of other specimens in the U. S. N. M. bear the label "Ariz., Collection C. F. Baker." Among the eleven species of Alcojieura which are distinctively yel- low and white, general resemblances are at once apparent, and in many instances coloration markedly intergrades. It is submitted, however, that closely related species not uncommonly exhibit such external similarity, and certainly in these cases there is no compa- rable intergradation of the male genital characters, although there are indubitable resemblances. Fully colored specimens of any two given species, however close, show plainly differences in the mark- ings, one being invariably more definitely and extensively striped or spotted. Unfortunately, the optimum in coloration cannot always be obtained, and, as for many species in widely separate genera, the male genitalia must be regarded here as the positive means of classi- fication. 9. Alconeura quadrimaculata Lawson Alcoiieura qiiadrimaciilata Lawson, P. B., Bull. Brooklyn Ent. Soc, XXV, 45, 1930. Resembling Alconeura unipuncta (Gillette) , but with much lighter coloration, marked with orange, not orange-brown; aedeagus with- out stout basal tooth on dorsal margin ; pygofer process recurved into heavy, sharp-pointed hook. "Vertex, pronotum and scutelliun milky white, with traces of four yellow or orange-red lines on pronotum and hints of color in basal angles of scutellum. Elytra with two bright, well-separated, orange- red spots on each clavus, a yellow line between first and second sec- tors and sometimes a suggestion of yellow latcrad of this near the costal margin; some of apical veins margined with fuscous, par- ticularly along costal margin; a small but distinct black spot near middle of third apical vein. Underside pale except for black tarsal claws and tip of ovipositor." Genitalia. Aedeagus in lateral view nearly twice as long as width of base; narrowing in apical third to less than one third basal width, curving dorsad to rounded tip; line of ventral margin nearly semi- circular. Pygofer process nearly as long as aedeagus, base as wide as apical portion of shaft; constricted in middle, apex recurving, forming sharp-pointed hook. Types studied: Holotype, female; allotype, male; paratypes, ten females and eight males: Pima county, Arizona, July 27, 1927. Also, one paratype, male, from Gila county, Arizona, August 6, 1927. R. H. Beamer. San Diego county, California, July 7, 1929; Alpine, Griffith: Alconeura of the U. S. 325 California, July 9, 1929; Sabino Canyon, Arizona, July 14, 1932; Baboquivari Mountains, Arizona, July 19, 1932; Patagonia, Arizona, August 22, 1935; R. H. Beamer. Verde Valley, Arizona, September 20, 1922; H. R. Brisley. Santa Rita iMountains, Arizona, June 16, 1933; Patagonia, Arizona, June 24, 1933; P. W. Oman. Los Angeles county, California, Coquillet. Three specimens taken in Pasadena, California, April 4, 1928, bear the label "sycamore, Arroyo Seco." 10. Alconeura derecta, n. sp. Resembling Alconeura quadnmaculata Lawson, but smaller, with py gofer process straight, not hook-shaped. Color. Creamy-white with orange markings. Vertex with orange inverted V extending from apex to posterior margin, widening and touching eyes ; pronotum with irregular orange \'ittae forming an ]M ; scutellum orange and creamy-white. Forewing with two large yel- low spots on clavus, one basal and ovate, other slightly beyond mid- dle, usually forming one half of dorsal spot evident when wings are at rest; corium with bright yellow vitta curving in from costal margin and i)aralleling distal two thirds of claval suture almost to cross veins, joined by shorter smoky-yellow stripe from mid-costal mar- gin. Apical veins yellowish, margined with fumose; apical dot in third apical cell. Tip of forewing darkened. Venter stramineous. Genitalia. Aedeagus in lateral view about twice as long as width at base, narrowing abruptly in apical third to approximately one third basal width and curving dorsad to truncate tip; line of ven- tral margin roughly semicircular. Pygofer process as long as aedeagus, slender and nearly straight, so twisted as to appear con- stricted just before obliciuely truncated apex. Holotype, male; allotype, female; paratype, female: Key Largo, Florida, August 9, 1930. Batesburg, South Carolina, August 24, 1930. R. H. Beamer. 11. Alconeura insulae, n. sp. Resembling Alconeura derecta n. sp.. but more delicately marked with yellow; obliciue lines near margin of vertex forming inverted V extending to eyes; two processes on pygofer. Color. Creamy-white with golden-yellow markings. Vertex with yellow inverted V near anterior margin, extending to eyes, often indistinct apically, continuing medianly to posterior margin in two parallel lighter lines; pronotum with yellow lines forming irregular trapezoid in each half, adjacent median sides of these lateral figures almost parallel; angles of scutellum yellow. Forewing with yellow 326 The University Science Bulletin spots on clavus, one basal and ovate, another just beyond middle and forming one half of dorsal spot completed when wings are closed; corium with yellow stripe from near humeral angle to middle of claval suture, and with similar strii)e extending obliquely inward from midcostal margin, enlarging at tip of clavus, joining small fumose mark. Apical veins pale yellow, fumosely bordered near costal margin; apical dot in midapical cell, touching or upon third vein. Tip of forewing suffused with yellow. Venter stramineous. Genitalia. Aedeagus laterally compressed, in lateral view major portion about twice as long as wide; thicker dorsoventrally near laterally expanded apex, and with slender process briefly paralleling ventral margin in l)asal tliird of aedeageal shaft. Pygofer with two processes, semiequal in length, obliquely joined at base forming U- shaped projection occupying space approaching two thirds size of aedeagus; inner spur slightly sinuate, gradually tapering to point; outer spur tapering abruptly in apical tliird and incurved. Holotype, male; allotype, female; paratypes, twelve pairs: Key Largo, Florida, August 9, 1930, R. H. Beamer. 12. Alconexira dodonana Beamer Alconeura dodonana Beamer. R. H,. Can. EnT., I^X\'I. 17. 1!I34. "Resembling Alconeura qiiadrimariilata Lawson, somewhat. l)ut with many more orange markings, and a flattened face instead of arched. "General ground color yellowish white. Vertex with inverted V- shaped orange mark with arm of ^' bi'oadened to touch eyes. Pro- notum \yh\\ inverted undulating ^^'-shaped orange mark reaching both anterior and jiosterior margins. Scutellum all orange except small sciuare anteiiomesal yellowish wliite spot. Clavi with large, almost circular, basal orange spot and an elongated bilobed mesal one. Coria with diagonal orange dash at anterior end of costal plaque ending in enlarged romided area at claval suture; another slightly narrower and shorter diagonal dash at posterior end of costal plaque and an oval spot near its inner end. Cross veins light, mar- gined with fuscous. Small round black sjiot in base of cell M^. Venter stramineous suffused with orange. T\p of ovipositor often black." Genitalia. Aedeagus in lateral view more than three times as long as width of base; shaft curving dorsad, evenly tapering to slender dorsad-bent tip; line of ventral margin semicircular; pair of lanciform processes arising at base and paralleling aedeagus, though not quite as sharply curved. Pygofer jirocess about two thirds as Griffith: Alconelra of the U. S. 327 long as aedeagiis, base heavy, slightly narrower in distal portion, apex bifurcate. Types studied: "Holotype, male and allotype, female, Sabino Canyon, Arizona, July 14, 1932, R. H. Beamer: 19 female and one male paratypes same data. One female paratype, Baboquivari j\Its., Ariz., July 19, 1932, R. H. Beamer." "All specimens were collected on Dodonaca viscosa var. angusti- folia." 13. Alconcura lappa, n. sp. Resembling Alco?ieura quadrimaculata Lawson. l)ut smaller and wdth pronotal markings more definitely ^•ittate, not fused; aedeagus with several l)asal spur-like jjrocesses; pygofer }irocess reduced to stout tooth. Color. Creamy-white with orange-yellow markings. Vertex w'ith orange inverted V, apically obscure, extending to posterior margin, frequently touching eyes; pronotum with four orange vittae, each lateral pair often fused posteriorly and mure faintly joined ante- riorly; angles of scutellum orange. Forewing with two yellow spots on clavus, larger one near base and ovate, other in middle, becoming half of dorsal spot formed when wings are at rest; corium with yel- low stri])e bordering distal two thirds of cla\'al suture, joined ob- liquely at base with similar stripe coloring proximal half of costal margin. Apic;d veins yellow, funiosely ourliiied; apical dot in third apical cell, touching or upon third \-ein. Tip of forewing yellow, delicately suffused, ^'enter stramineous to brighter yellow. Genitalia. Aedeagus with many spur-like jirocesscs on basal half; in lateral xlvw. shaft ajiically slentler (about one third width of base), divided into ventral and dorsal arms; basal width of aedeagus less than one fourth length. Pygofer process about one half length of aedeagus, semitriangular with broad base and sharp tip. Holotype, male; allotype, female; paratypes, fifteen males and twelve females: Hauchuca Mountains, Arizona, July 8, 1932, R. H. Beamer. Additional paratypes: two males and one female, Sabino Canyon, Arizona, October 8, 1932 (E. D. Ball collection) ; one male, Santa Rita IMountains, Arizona, June 27, 1933, P. W. Oman (U. S. N. M.). Chiricahua INIountains, Arizona, July 8, 1932; Sabino Can- yon. Arizona, July 12 and July 14, 1933; R. H. Beamer. 14. Alconcura macra, n. sp. Resembling Alconcura lappa, n. sp., but tips of forewings more deeply suffused with fumose-yellow, apical dot distinctly larger; 328 The University Science Bulletin aedeagus with less than five basal spurlike processes; pygofer proc- ess with attenuated tip. Color. Creamy-white wdth orange-yellow markings. Vertex with orange-yellow inverted V, apically indistinct, lateral stripes extend- ing with little widening to posterior margin, often touching eyes; pronotum marked with four orange-yellow vittae, lateral pairs lightly joined at anterior end, often apparently connected posteri- orly; scutellum orange-yellow^ excepting small anteromedian white square. Forewing with two orange-yellow claval spots, one basal, elongate-oval, one in middle part completing oval dorsal spot when wings are closed; corium with orange-yellow stripe cur^'ing inward from costal margin and paralleling distal two thirds of claval suture to cross veins, meeting fumose mark interruptedly proceeding in from yellowish spot on midcostal margin. Apical veins yellow with dark borderings; apical dot on third vein. Tip of forewing yellow, smokily suffused. Venter stramineous, occasionally brighter yellow. Genitalia. Aedeagus with less than five spurlike processes on basal half; in lateral view, shaft apically slender (about one third width of base), divided into ventral and dorsal arms; basal width of aedeagus nearly one fifth of lengtli. Pygofer process about one half length of aedeagus, semitriangular, broad at base with sharp, attenuated tip. Holotypc, male; allotype, female; paratypes, ten pairs: Pensacola, Florida, July 12, 1934, R. H. Beamer. Additional paratypes: six males and four females, Alabama, collection C. F. Baker; two males, Utica, Mississippi, August; two males and one female, Tallulah, Louisiana (on cotton), September 6, 1929, R. L. McGarr; two fe- males, Mullins, South Carolina (on Hypericiuvi) , May 5, 1932, P. W. Oman (U. S. N. M.). Sanford, Florida, July 25, 1934; Hilliard, Florida, July 28, 1934; Vinton, Louisiana, December 25, 1931; R. H. Beamer. Sanford, Florida, June 6, 1933, C. 0. Bare. Hamilton, Mississippi, July 15, 1930, L. D. Tuthill. Douglas county, Kan- sas, August 17, 1928, and July 25, 1929, P. B. Lawson. 15. Alconeiira rotunclata Ball and DeLong Alconeura rotuiidata Ball. E. D., and DeLong, D. M.. Ann. Ent. Soc. Am., XVIII, 335, 1925, Alconeura rottindata Ball and DeLong, Lawson, P. B., Bull. Brooklyn Enf. Soc., XXV, 40, 1930. (Evidently confused witli Alcoiiewa nridnta Ball and Delong. and Alconeura lanyuida, n. sp.) Resembling Alconeura vnipuncta (Gillette) but stouter, marked with yellow not darkened with fumose, apical dot in base of fourth, not in third cell; tip of aedeagus in ventral view with winglike processes. Griffith: Alconeura of the U. S. 329 Color. Creamy-white with orange-yellow markings. Vertex with two orange-yellow vittae, almost forming inverted V; orange spot next anterior margin of eye; pronotmii witli four irregular orange- yellow vittae; angles of scutellum marked with orange-yellow. Forewing white, semihyaline with two yellow vittae paralleling claval suture, one on clavus from basal portion into distal third, one on corium from near humeral angle almost to cross veins; large yellow mark on midcostal margin. Apical veins yellow with fulvous margins; apical dot in base of fourth cell. Tip of forewing fulvous. Venter stramineous. Genitalia. Aedeagus in lateral view about tlirec times as long as basal width, narrowed in apical third to nearly one third middle width, curving slightly dorsad; witli pair of winglike jirocesses at tip, widely divided to point of attachment into two sharp-pointed spurs on either side of apex of shaft. Pygofer process little wider than apical portion of aedeagus, about six times as long, sharply bent in middle, tapering evenly to i)oint. "Described from three females and one male from Ames, Iowa. Taken by the senior author August 5, 1895, on Aviorpha fruticosa, and one male from Onaga, Kansas, July 9, 1898 (Crevecoeur) , and five specimens from Ames, Iowa, in collection of Prof. Herbert Os- born. This is the stoutest species in the genus. It is much broader than uni'pnncta, which it otherwise resembles." Type studied: Paratype, male, Ames, Iowa, August 5, 1895, Herbert Osborn collection (U. S. N. M.). Garnett, Kansas, July 7, 1933, P. W. Oman. Scott county, Arkansas, August 24, 1928 (on willow) ; Arkansas City, Kansas, June 12, 1935; R. H. Beamer. This handsome species was chosen by Ball and DeLong as the genotype and displays admirably the genotypical features, though the curious structure of the aedeagus shows it to be somewhat re- moved phylogenetically from the rest of the species described. 16. Alconeura midata Ball and DeLong Atconcvra vnipuncta nudata Ball, E. D., and DeLong, D. M., Ann. Ent. Soc. Am., XVIII, 336, 1925. Resembling Alconeura rotundata Ball and DeLong, but with apical dot on third vein, not in base of fourth cell; aedeagus with tonguelike ventromedian extension, without winglike processes at tip; pygofer process reduced to small rounded tooth. Color. Creamy-white with orange-yellow markings. Vertex with inverted V in orange-yellow from apex to posterior margin, eyes narrowly bordered witJi yellow; pronotum with four orange-yellow 330 The University Science Bulletin vittae, lateral pairs in males fused posteriorly; angles of scutellum orange and yellow. Forewing chiefly yellow, claviis almost entirely so; corium with two yellow vittae, diverging from humeral angle, one paralleling distal two thirds of claval suture, one along basal half of costal margin. Apical veins light with fulvous margins; apical dot on third vein. Tip of forewing darkened. Venter stra- mineous, often relieved with brighter yellow. Genitalia. Aedeagus in lateral view more than six times as long as basal width; with jiair of slender ventrolateral processes arising in basal third, paralleling shaft to tip, bending slightly ventrad at tips; median tonguelike extension lying between processes and shaft, reaching into apical fourtli; ti]) of shaft projecting ventrad into short tooth. Py gofer process almost wanting, represented by very slight elevation. "Described from five exami^les from Pasadena and San Francisco. California. Collected by the senior author." The very short description given by Ball and DeLong was drawn apparently from exceptionally light specimens, and cannot be valued as truly representative of this brightly colored species, so very dis- tinct from Alconeura iinipuncta (Gillette). The description above was prepared from a jiair of i)lesiotyiU's selected from a large series. Types studied: Allotype, male, and paratype, female, San Fran- cisco, California, September 9. 1007 lE. D. Ball collection i. Plesio- types, male and female, San Jacinto ^Mountains, California, July 21, 1929, R. H. Beamer. Haugan, Montana, August 9, 1931; Yellow- stone Park, Frying Pan Lake, August 15, 1931; Grand Teton Xa- tional Park, August 18, 1931; Mt. Shasta, California, June 29, 1935; Siskiyou N. F., California, July 14, 1935; Eureka, California, July 15,1935; R. H. Beamer. 17. Alconeura languida, n. sp. Resembling Alconeura nudata Ball and DeLong, but with lighter markings and less general yellowness; tips of paired ventrolateral processes on aedeagus and apical third of shaft bending abruptly dorsad. Color. Creamy-white with orange-yellow markings. Vertex with orange inverted V, diverging lines almost parallel from aj^ex to posterior margin; pronotum with four orange vittae; median pair continued across scutellum, apex of latter orange. Forewing with yellow vitta on clavus from near base almost to tip; corium witli two yellow stripes, one paralleling claval suture from near humeral Griffith: Alconelra of the U. S. 331 angle almost to cross veins, one from base on costal margin halfway, briefly continuing obliquely inward along veins to transverse fumose mark. Apical veins yellow, fumosely outlined; apical dot in third cell, touching or upon third vein. Tip of forewing slightly darker. Venter stramineous. Genitalia. Aedeagus in lateral view more than seven times as long as basal width; with pair of slender ventrolateral processes arising in basal third and attaining length of aedeagus, slightly sin- uate, bending dorsad at tips; median tonguelike extension lying be- tween processes and shaft, reaching past middle of and bending dorsad with apical third of shaft; tip of shaft recurving ventrad. Pygofer almost wanting, represented by toothlike projection, often appearing obliciucly truncate. Holotype, male; allotype, female; paratypes, twelve pairs; Chiri- cahua ]\Iountains. Arizona, July 8, 1932. Huachuca Mountains, Arizona, July 8, 1932; Ashfork, Arizona, August 8, 1932; Yellow- stone Park, August 15, 1932; Big Bear Lake, California, July 26, 1932; Flagstaff, Arizona, August 1, 1933; Santa Rita Mountains, Arizona, August 18, 193."); R. H. Beamer. 18. AlcuNcura luculoita, n. sj). Resembling Alconcura nudata Ball and DeT.ong, but even more extensively yellow-colored and smaller; shaft of aedeagus with tip straight, not curving ventrad. Color. Creamy-white, but largely obscured by broadening and fusing of orange-yellow markings. Vertex with orange-yellow in- verted V joining orange-yellow lines bordering eyes at apical and posterior margins; pronotum with four orange-yellow vittae, these often quite broadly fused; scutellum with orange-yellow markings more or less fused. Forewing largely yellow, claval area completely so; corium with two very broad yellow stripes, one bordering claval suture almost to cross \'eins, other slightly separate, extending from humeral angle along costal margin to cross veins. Apical veins light, margined with fulvous; apical dot in third cell, touching or upon third vein. Tip of forewing slightly darkened. Venter yellow and brown; face bright yellow. Genitalia. Aedeagus in lateral view more than seven times as long as basal width; with pair of slender ventrolateral processes arising in basal third, attaining length of aedeagus, tips bending slightly dorsad and abruptly recurving ventrad; median tongue-like extension between processes and shaft reaching past middle of apical third of shaft; tip of shaft straight. Pygofer process almost want- ing, represented by rounded toothlike projection. 332 The University Science Bulletin Holotype, male; allotype, female; paratype, male: Chiricahua Mountains, Arizona, July 8, 1932, R. H. Beamer. Additional para- types: two males and twenty-two females, Ashfork, Arizona (on Fallugia paradoxa) , June 30, 1933, P. W. Oman (U. S. N. M.). Wickenburg, Arizona, July 8, 1932; Las Cruces, New Mexico, June 8, 1933; R. H. Beamer. Organ, New Mexico, June 8, 1933, P. W. Oman. 19. Alconcura cornigera, n. sp. Resembling Alconeura luculenta, n. sp., but with orange markings more definite; aedeagus divided into ventral and dorsal arms; pair of slender lateromedian processes reaching only into apical third of shaft; pygofer process hornlike, sharp-pointed, not simply a rounded tooth. Color. Creamy-white with orange-yellow markings. Vertex with orange inverted V, lateral stripes parallel to posterior margin; pro- notum with four irregular orange vittae, broadening posteriorly; scutellum orange, except anteromedian white rectangle. Forewing almost all yellow on claval area; corium with yellow stripe border- ing distal two thirds of claval suture and one from Immeral angle along costal margin almost to cross veins. Apical veins yellow, edged with fumose lines; a])ical dot in third cell, touching third vein. Tip of forewing slightly darker. Venter light yellow. Genitalia. Aedeagus in lateral view more than three times as long as middle width; divided into ventral and dorsal arms, former curving to approach latter closely at tip; pair of slender pointed processes extending along sides from near base to distal third of shaft. Pygofer process about as broad at base as middle width of aedeagus and curving out almost into right angle, sides evenly con- verging to pointed tip. Holotype, male: Huachuca Mountains, Arizona, August 1, 1927, R. H. Beamer. Alconcura (juadrivittata (Oillette) Although specimens have not been taken in the United States, it should be noted here that Ball and DeLong (1925) place a species, described as Dicranciira quadrivittata by Gillette (1898), among the Alconeura. McAtee (1926) makes the following note on the six'cies: ''Ball and DeLong have shown that the type locality for this species is Long Island, Bahamas. It needs only a glance at the figure of teg- men given bv C.illotte, or that of Ball and DeT>ong, to show how Griffith: Alconeura of the U. S. 333 slight a variation would serve to place the angulate but sessile third apical cell in the stalked categoiy and thus give this species two stalked apical cells, a character which seems to be the only important one of the segregate Kahaono Kirkaldy. The type specimen has been available for the present study." The third apical cell in the forewing of Alconeura dodonana Beamer is similarly angulate; this condition is quite variable, but simple angularity seems to be the extreme point in variation from the normal. There is at present no reason to conclude that it may be otherwise in Alconeura quadrivittata (Gillette), which is known only from the single male type now in the National Museum. While possibly confined to its island home, this troublesome species should be kept in mind by collectors in the more southern regions of the United States. LITERATURE CITED 1898. Gillette. Clarence P. American leaf hoppers of the subfamily Typhlo- cybinae. Proc. U. S. Nat. Mus., XX, pp. 718, 723-724, Figs. 19-21, 40-42. 1914. Van Duzee, E. P. A preliminarj^ list of the Hemiptera of San Diego Coimty, California. Trans. San Diego Sec. Nat. Hist., II, p. 56. 1924. DeLono, Dwight M. Some new Cicadellidae (Homoptera) from the southern Unifed States. Journ. N. Y. Ent. Soc, XXXII, pp. 67-68. 1925. Ball, E. D., and DeLong, D. M. The genus Dikraneura and its allies in North America. Ann. Ent. Soc. Am., XVIII, pp. 334-337, Plates XXI, XXIII. 1926. McAtee, W. L. Notes on Neotropical Eupteryginae, with a key to the varieties of Alebra albostriclla (Homoptera, Jassidae). Journ. X. Y. Ent. Soc, XXXIV, pp. 158-159, 163-164. 1930. Lawsox, Paul B. Two new .Alconeura (Homoptera, Cicadellidae) with notes on other species. Bull. Brooklyn Ent. Soc, XXV, pp. 44-46, Figs. 1-8. 1934. Beamek, R. H. Notes on leaf hoppers (Homoptera, Cicadellidae). Can. Ent., LXVI, p. 17. 1934. McAtee, W. L. Genera and Subgenera of Eupteryginae (Homoptera, Jassidae). Proc. Zoolog. Soc. London, Part 1, pp. 105, 111. ILLUSTRATIONS Figures of the dissected male genitalia are presented in the specific order of the descriptions. It will be noted that the aedeagus has been pictured consistently with the ventral side toward the top of the page ; this is believed the best position, since attention should be centered most on the ventral margin and its processes. The pygofer process in the position shown would lie below the other member of the pair if the illustration included both for each species. The last two figures show the apeces of typical forewings, the apical cells and alternate positions of the apical dot. 334 The University Science Bulletin PLATE XXIII A: aedeagus; ventral margin upper. B: pygofer; inner margin upper. Fig. 1. Alconeura planala Ball and DeLong. Fig. 2. Alcoiieura necopinata, n. sp. Fig. 3. Alconeura beameri. n. sp. A: Ventral view of tip of aedeagus. Fig. 4. Alconeura balli Beamer. Line of measurement for basal width shown on aedeagus. Fig. 5. Alconeura tricolor (Van Duzee). Griffith: Alconeura of the U. S. 335 PLATE XXIII 336 The University Science Bulletin PLATE XXIV A: aedeagus; ventral margin upper. B: pygofer; inner margin upper. Fig. 6. Alconcura julminea Lawson. Fig. 7. Alconeura dorsalis (DeLong). Fig. 8. Alconeura unipuncta (Gillette). Fig. 9. Alconeura quadrimaculata Lawson Fig. 10. Alconeura derecta, n. sp. Griffith: Alconel'ra of the U. S. PLATE XXIV 337 22—6037 538 The University Science Bulletin PLATE XXV A: aedeagus; ventral margin upper. B: pygofer; inner margin upper. Fig. 11. Alconeura insidae, n. sp. Fig. 12. Alconeura dodonana Beamer. Fig. 13. Alconeura lappa, n. sp. Fig. 14, Alconeura macra, n. sp. Fig. 15. Alconeura rotundata Ball and Delong. A: ventral view of tip of aedeagus. Griffith: Alconeura of the U. S. 339 PLATE XXV 340 The University Science Bulletin PLATE XXVI A: aedeagus; ventral inarsiin III )pcr. B: pygofer ; inner margin upper. Fig. 16. Alconeura naidala Ball and DeLong. Fig. 17. Alconeura languida, n. sp. Fig. 18. Alconeura luculenta, n. sp. Fig. 19. Alconeura cornigera, n. sp. Fig. 20. Alconeura quadrimaculata La\v.sun. Apex of left forewing; from left to right: first, second (or i)edunculate), third (or midapical), and fourth apical cells. The apical dot is in the third cell, partly lying upon the third vein. Fig. 21. Alconeura tricolor (Van Duzee). Apex of left forewing; apical cells as in Fig. 20. The apical dot is in the base of the fourth cell. Griffith: Alconeura of the U. S. 341 PLATE XXVI I6a THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 16. A Generic Revision of American Bythoscopinae and South American Jassinae* By P. W. OMANf CONTENTS PAGE Introduction 345 Acknowledgments 346 Morphological Notes and Terminology 346 Characters of the Subfamilies 347 Key to the American Genera of Bythoscopinae 349 Descriptions of Bythoscopine Genera 350 Key to the South American Genera of Jassinae 355 Description of Jassine Gent ra 358 Literature Cited 400 Illustrations 403 Index 420 Ab.stuact: The Aniciican genera of Bythoscopinae and the South American genera of Jassinae are reviewed. A key to the American subfamilies of Cicadel- lidae, and keys to the genera of the subfamilies indicated are included, as well as drawings illustrating the head structures and wing venation of the genotypes of most of the genera treated. Taxonornic terminology for leaf hoppers is discussed and some new terms introduced. The new categories described and nomenclatorial changes made are as follows. New genera: Euragallia (type, AgaUia furculola Osborn), AgalUota (type, Agallla punctata Oman) , Brasa (type, Macropsis rugicoUh Dozier), Ncopsls (type Pcdiopsis elegans Van Duzee), By- thonia (type Nionia (?) ricgosa Osborn), Cariancha (type Cariancha canboba, n. s]).), Brincadonis (type Brincadorus laticeps, n. sp.), Cerrillus (type Hccalus notatus Osborn). Arrngada (type Huleria rugosa Osborn), Egenus (type Egcnus acuminatus, n. si).), Bunamus (type Bonamus linealus, n. sp.), Bolarga (tj'pe * Submitted to the Department of Entomology and the Faculty of the Graduate School of the University of Kansas in partial fulfillment of the requirements for the degree of Master of Arts, 1935. t Bureau of Entomology and Plant Quarantine, United States Department of Agriculture. (343) 344 The University Science Bulletin Fambolocmtus buUvUntus Osbcrn), GarapUa (type Garapita garbosa, n. sp.), Agitdus (type Agudiis iypicus, n. sp.), Ciimom (type Cumora angulnta, n. sp.), Kanorba (type Kanorba reflcra, n. sp.), Caphodus (type Caphodus macidatus, n. sp.), Baroma (type Baroma reticulata, n. sp.), Bahita (type Eutettix in- jiiscata OHborn), Alaini.s (type Eutettrix dentatus Osborn), Hcgira (type Hegira bnniuea, n. sp.), Fnltain (type Faltnln brachyptera, n. .sp.), Brazosa (type Thamnotettix picturcllus Bilker). .4/n(V( (type A/ora longicauda, n. sp.), Onura (type Onura vbiirncola, n. s)).), Bcuala (type DcUorcplKdu^ liimidns Osborn), Cortona (type Cortona inimita, u. sp.), Agclina (tyiie Agelina punctata, n. sp.), Neobala (type Thainuotcttix pallidus Osborn), Connia (tyjie Spangbcrgiella fasci- ata Osborn) , and Coclidiaua (type Ncococlidia rubrdliiicta Baker) . New subgen- era: Aceratagallia (BergaUia) (type Bythoscopus .•i Stal), Dcltoci phalus (Haldorus) (type TJiuiniiotcttix vcnlaus Osborn). New synonymy: Bythoscopus Gcrniar {=^Eurinoscopu!< Kirkaldy) ; Por/z/yoyw/.s Uhler (^Strngauiopsis Baker) ; Pachyopsis laetus Uhler {^^ Macropsis idioceroidcs Baker); Spangbergiella Signoret (=: BergifUa Baker); Bolarga bolivmnun (Osborn) (^ Bolarga boli- vianu.'< \',ir. pj)ius l^all. Types of the new species are in the c<)ll(>c(ion of the United States Xalional IVIuseiun. Oman: Bythoscopinae and Jassinae 345 INTRODUCTION FOR over a century and a half workers in taxonomic entomology- have been contributing to our knowledge of the Cicadellidae or leaf hoppers of South America. Most of these contributions have been concerned primarily with the description of new species, these being assigned to the then established European or North American genera known to the describer. Although a great deal of work has been done in recent years on the proper differentiation of the North American genera, the proper assignment of the South American representatives of these groups has not kept pace, and what few genera have been erected for South American forms ha\'e in most cases been based on unusual and phylogenetically isolated species, so that their erection adds little to a natural arrangement of the more complex portions of the higher groups. Conseciuently, the accepted limits of the established genera have been so enlarged liy the in- clusion of many species representing diverse evolutionary trends that these heterogeneous groups cannot be defined and become use- less as a basis for classification. It is similarly impossible to use the mass of specific descriptions as a basis for the identification of species in the group, and it becomes evident that before any ap- preciable order can be l)rouglit from the confusion that exists, genera must be defined, keys presented for the separation of those genera, and the pre\-iously described s])ecies referred to their correct ge- neric positions. The present paper is an attempt to do this for the subfamilies indicated, but in many cases the assignment of pre- viously named species to tlieir respective genera has been impos- sible, owing to the lack of adequate descriptions, and must await the examination of types or authentically determined specimens. At the time this study was undertaken it was the intention of the writer to limit it to the South American fauna, l)ut the North American Bythoscopinae were treated also when it became ap- parent that they could be included without greatly enlarging the paper. The writer realizes the inadequacy of a restricted fauna as a basis for a generic revision, and has endeavored to eliminate er- rors as far as possible by constant reference to literature and col- lections dealing with other faunal regions. In this he has been par- ticularly fortunate in having available for study genotypes of most of the F.uropean, Philippine, and Australian genera and a consider- 346 The University Science Bulletin able number of African and Indian species. Generic concepts here presented have been based primarily on the study of genotypes, plus as many other species as were available. If authentically de- termined material of the type species of a genus has not been avail- able for study it is so stated in the discussion of the genus. ACKNOWLEDGMENTS The ])reparation of this paper has been expedited by the ready assistance given by various entomological workers. Expression of gratitude is made to Dr. R. H. Beamer, of the University of Kansas, under whose direction the major portion of the work was undertaken and who has assisted much with notes and suggestions; to ]Mr. H. G. Barber, of the Bureau of Entomology and Plant Quarantine, U. S. Department of Agriculture, who has given freely of his time for the discussion of problems of common interest to hemipterists; to Mr. R. E. Snodgrass, of the same bureau, for the generous assistance in problems of a mori)hoIogical nature; to Dr. E. D. Ball, of the Uni- versity of Arizona, for valued suggestions concerning taxonomic terminology ; to ]\Ir. E. P. Van Duzee, of the California Academy of Sciences, for kind suggestions and encouragement; to Dr. Hugo Kahl, curator of insects at the Carnegie Museum, for many kind- nesses while the writer was working at that institution, and for the loan of valuable material; to jMr. C. H. Ballou, of Costa Rica, for specimens of Chinaia bella Bruner and Metcalf ; to Mr. J. N. Knull, curator of insects at Ohio State University, for the loan of valuable material and for notes and drawings of other material not available to the writer. MORPHOLOGICAL NOTES AND TERMINOLOGY The WT'iter makes no attempt in the present paper to enter into a detailed discussion of the morphology of the Cicadellidac, but pro- poses merely to define the terms used and to illustrate the struc- tures mentioned sufficiently to permit the user of the paper to in- terpret correctly and easily the verbal descriptions of the writer. A necessity for so doing has arisen through the introduction of a few terms new to tlic literature on Homoptera, and the adoption of others that have been little used by taxonomic workers. The choice of terms has been made only after careful consideration of the needs of the taxonomist, recognizing that for the ]~)urpose of writing in- telligible and at the same time reasonably brief descriptions, termi- nology borrowed from morphological workers is freciucntly unsatis- factory. Oman: Bythoscopinae and Jassinae 347 According to recent work on the insect head by Snodgrass, 1935 (pp. 111. 329-331), the frons in the Cicadellidae is greatly reduced and not evident as a distinct area. The area previously referred to by most workers as frons, is, according to Snodgrass, the postclypeal jilate, while the clypeus of certain authors is the anteclypeal plate of Snodgrass. The writer has chosen, for taxonomic work, to restrict the term clypeus to the postclypeus of Snodgrass and use the term clypellus for his anteclypeus (PI. XXVII, fig. IB). The clypeus and clypellus together constitute the front. The entire cephalic aspect of the head is referred to as the face (PI. XXVII, fig. IB), and the area between the eye and the adjacent ocellus, or between the eye and the lateral frontal suture, if it is present, is referred to as the ocellocular area. The entire dorsal surface of the head, ex- clusive of the eyes, is called the crown (PI. XXVII, fig. IC). In referring to the veins of the wings, the Comstock-Needham sys- tem has been followed with the now generally accepted emendations, as indicated (PI. XXVII, figs. 1 and lA). and certain other modi- fications in naming which are now well established in literature. The forewing of leaf hoppers normally has a basal cross vein between media and cubitus, called the first cross vein, and fre- cjuently has a second cross vein in the region at the base of the central anteapical cell. The claval suture ajipears to be cubitus 2. but the name claval suture is here retained. For the cells of the wings the writer has retained the well-established terms in use with but few additions. The anteapical cells, typically three in number, are designated inner, central, and outer, respectively, beginning nearest the claval suture, and the inner is considered as being pres- ent even though the second cross vein between M and Cu is absent, in which case the cell remains open basally. Similarly these cells are considered as distinct entities even though divided by extra cross veins. The apical cells, for purposes of stability in naming, are numbered from the inner margin of the wing outward, the third apical cell thus always having its base at the distal end of the central anteapical cell. CHARACTERS OF THE SUBFAMILIES The following key is presented for the separation of the subfamilies of Cicadellidae known to occur in America. 1. Ocelli ?itiuitf(l iirar di-k of crown, remote from eyes ami from anterior margin of ^ crown Ocelli on or near distal margin of crown, or on face, or absent .■;■.■" ^ 9 (1) Lateral frontal sutures extending over obtuse anterior margin of crown to vicinity " of ocelli Hind wing usuallv with submarginal vein evanescent from hamulus to apex leaving onlv three apical cells. Form usually somewhat cylnidrical, not flattened ....". Cicadellmae 348 The University Science Bulletin Lateral frontal sutures terminating below anterior margin of crown, which is usually distinct. Hind wing always with four apical cells. Form flattened dorsoventrally Gyponinae 3. (1) Ocelli on face, which is separated from crown by a distinct margin. . . .Koebelinae Ocelli either on face or not, or absent, but if on face, then face antl crown not .separated by a distinct margin 4 4. (3) Lateral frontal sutures terminating at or slightly above antennal pits. Inter- mediate tibiae without numerous large, spinelike setae Bythoscopinae Lateral frontal sutures extending above antennal pits to or near ocelli or oceliar vestiges. Intermediate tiliiae usually with numerous large, spinelike setae. ... 5 5. (4) Distance between ocelli less than distance between antennal pits. Face broad and relatively flat, crown always short and broad. Ocelli always on face. Eurymelinae Distance lietween ocelli or ocellar vestiges eqvial to or greater than distance be- tween antennal pits. Face usually not especially flat, crown of various shapes. Ocelli on or near margin between crown and face or absent 6 6. (5) Forewings usually with cross veins anterior to the apical series. Veins distinct basally ; ocelli us\ially present. Facial sclerites not highly fused, lori not par- ticularly elongated Jassinae Forewings usually without cross veins anterior to the apical series. Veins in- distinct basally; ocelli often absent. Faciae sclerites showing a high degree of fusion, lori unusually long and narrow. Small, fragile forms, usually brightly colored Eupteryginae In tlie consideration of the leaf hopper fauna of the entire world, or even that of one of the major faunal regions, it soon becomes evi- dent that the differentiation of subfamilies is an extremely difficult task. Consequently, the writer freely admits that the key pre- sented above is unsatisfactory in some respects, but feels that it is better to attempt to key out intermediate forms with the use of somewhat indefinite character^^, or a combination of characters, than to ignore such forms and use brief but positive alternatives. Idiocerus and allies lun'e been removed from the Bythoscopinae, with which they have little in common, and referred to the Euryme- linae. This will at first seem to be an erroneous disposition of the group, since Eurymela shows so many cercopid-like characters, but a study of species of Eurymeloides. Churna, and other Australian and Oriental genera convinces the writer that the arrangement is a natural one. Kirkaldy, 1906 (p. 345), McAfee, 1934 (p. 94), and perhaps others have pointed out that the much-used character of the position of the ocelli or ocellar vestiges is not a sufficient basis for the sep- aration of the Jassinae and Bythoscopinae. McAfee proposes to base the separation on the development of the bristles or spines on the lower surface of the tore tibiae, but the writer believes this to be even less reliable than the position of the ocelli. The separation here proposed is indicated in the preceding key. In addition to the characters indicated, tlie transverse suture on the scutellum is usually curved forward laterally in tlie l)ythoscopines and backward in the iassines, although this is bv no means a reliable character. Oman: Bythoscopinae and Jassinae 349 According to the above interpretation it has been found necessary t/O place Ionia Ball in the Bythoscopinae as a relative of Acerata- gallia and to transfer Chinaia Bruner and Metcalf from the By- thoscopinae to the Jassinae, where it is closely related to Jassus and allies. The writer has been unable satisfactorily to place Dicy- phonia picturata Osborn, 1924, but feels certain that the assign- ment of the species to the genus Dicyphonia Ball is erroneous. Key to the American Genera of Bythoscopinae 1. Lateral margins of pronotum carinate and moderately long. Form distinctly depressed. First apical cell of forewing very narrow. Ledges above antennal sockets transverse or nearly so 11 Lateral margins of pronotum short and not carinate. Form often broad but not depressed. First apical cell of forewing normal. Ledges above an- tennal sockets, when present, always oblique 2 2. (1) Hind wing with three apical cells. Distance between ocelli equal to, or usually greater than, twice the distance from ocellus to eye 9 Hind wing with four apical cells. Distance between ocelli not over twice dis- tance from ocellus to eye 3 3. (2) Distance between ocelli equal to or less than distance from ocelhis to eye. Lateral margins of genae not distinctly sinuated below eyes 8 Distance between ocelli always greater than distance from ocellus to eye. Lateral margins of genae usually distinctly sinuated below eyes 4 4. (3) Surface of pronotum finely granulated, neither coarsely pitted nor trans- versely rugulose 6 Surface of pronotum either coarsely pitted or transversely rugulose 5 5. (4) Surface of pronotum tran.sversely rugulose Aceratagallia , p. 3.")0 Surface of pronotum coar.sely pitted Agalliana p. 3.50 6. (4) Posterior margin of crown sinuately curved laterally and crown distinctly ex- tended lateralU' l>Hhind eye. I^ength iif crown medially usually less than next the eye. Eyes never bulbous Agalliopsis, p. 3.51 Posterior margin of crown smuothly rounded with only a narrow lateral ex- tension behind eye. Length of crown uniform, or, if shorter medially, eyes somewhat bulbous 7 7. (6) Crown shorter medially than eyes; eyes somewhat bulbous. Face rather flat, pronotum short. Seventh ventral segment of female sliort and expos- ing luiderlying membranes, styles of male genitalia not forked or with forks obsolete Euragallia, p. 3.51 Crown of uniform length. Face distinctly convex, pronotum longer in propor- tion to width. Genital characters not as above Agallia, p. 351 8. (3) Forewing at least partly membranous, tapering apically. Pronotum and scutellum not distinctly elevated or "humped" in profile Agalliota, p. 351 Forewing entirely opaque and subcoriaceous, broad apically. Pronotum and scutellum distinctly elevated or "humped" m profile Brasa, p. 352 9. (2) Hindwing with inner fork of radius and outer fork of media united apically (PI. XXVn, fig. 2.\); submarginal vein present between hamulus and apex of wing Neopsis, p. 353 Hind wing with forks of veins not united apically ; submarginal vein evanescent from hamulus to apex of apical cells (PI. XXVII, fig. 4) 10 10. (9) Pronotal striae transverse; anterior margin of pronotum broadly rounded, Oncopsis, p. 353 Pronotal striae oblituie, converging anteriorly; anterior margin of pronotum angulate medially Macropsis, p. 352 n. (1) Hind wing with three apical cells (PI. XX\'II, fig. 2.\). Inner apical mar- gin of forewing and first apical cell membranous and appendix-like, fre- quently bent down at a distinct angle to remainder of wing, Bythoscopus, p. 353 Hnul wing with four apical cells. Inner apical cell of forewing not as above. Pachyopsis, p. 354 350 The University Science Bulletin DESCRIPTIONS OF BYTHOSCOPINE GENERA AceratagaUia Kirkaldy* AceratagalUa Kirkaldy, Hawaiian Sugai- Planters' Atisoc. Expt. Sta., Div. Ent. Bull. 3, p. 30. 1907. Type of the genus, Bythoscojms sanguinolentus Provancher, 1872 (p. 376) , a Nearctic species. As defined in the key, this genus occurs in the Nearctic, Neotropical, Palearctic, and Oriental regions. The typical forms, referred to the subgenus AceratagaUia, are known only from North America. The typical forms of the genus, and those belonging to the other two subgenera, may be differentiated by the following key: 1. Styles of internal male genitalia forked. South America Bergallia Styles of internal male genitalia not forked. North America 2 2. (1) Very .small, extremely robust species, ocelli visible from above. Color, reddish-brown Ionia Larger and less robust species, the ocelli not visible from above. Color usually some shade of fuscous, brown, or cinereous, never reddish -brown, AceratagaUia AceratagaUia subgenus Ionia Ball loiiUi Ball, Bull. Brooklyn Ent. Soc, vol. 28, p. 22(1, 1933. Type of the subgenus, Ionia triunata Ball, 1933. This is the only known representative of the subgenus. The writer does not consider the difference in size and position of the ocelli sufficient basis for treating Ionia as distinct from AceratagaUia, with which it agrees in having the frontal sutures terminating at the antennal sockets, a transversely striated pmnotuni, and ty|iical acoratagallian genitalia in the males. AceratagaUia subgenus BergaUia, new subgenus Type of the subgenus, Bythoscopus signatus Stal, 1859, a Neotropi- cal species known only from Argentina. This subgenus is erected to accommodate the South American representatives of AceratagaUia. AgaUiana Oman Analliaiia Oman, U. 8. Dept. Agr. Tech. Bull. 372, p. 70, 1933. Type of the genus Bythoscopus sticticoUis Stal, 1859 (p. 291), a Neotroi)ical species. The genus is apparently limited in distribution to the Neotropical region, and is known to include, in addition to the genotype, AgaUiana grossa Oman, mnigera Oman, jjisca Oman, and ancora Oman, 1934. * The writer is treating the American genera of the AgaJlia gi-oup in detail in a forthcom- ing i>ublication and considers it unnecessary to include complete descriptions here. Oman: Bythoscopinae and Jassinae 351 AgaUia Curtis Agallia Curtis, Ent. Mag., vol. 1, p. 193, 1833. Type of the genus, Agallia consobrina Curtis, 1833, which is at present considered to be a synonym of Jassiis puncticeps Germar, 1832 (fasc. 17, ]). 2), a Palearctic species. The genus {sensus stric- tus) is known to occur commonly in the Palearctic, Nearctic, and Neotropical regions, and possibly also in the Oriental region, al- though a careful study of the Oriental species may reveal characters which will justify generic segregation. Euragallia, new genus Type of the genus, Agallia furculata Osborn, 1923, a Neotropical species. So far as known the genus is limited in distribution to the Neotropical region. Euragallia bears a superficial resemblance to Moonia Distant, type, Moonia sancita Distant, 1908, from British India, but differs in the ])osition of the ocelli, the shape of the prono- tum, and other pertinent characters, and is more closely related to Agallia Curtis. ^4 ga lliops is Ki rka Idy Ac/alUopsis Kiikaldv, Hawaiiiin Sugai- Planters' Assoc. Exiit. Sta., Div. Ent. Bull. 3, p. 31, 1907. Type of the genus, Jas.sus norcllii^s Say, 1831, an American species. The genus is apparently limited in distribution to the Nearctic and Neotropical regions. Agalliopsis bears a distinct resemblance to Nehela Buchanan White, tyi>e Nehela vulturiana Buchanan ^^'hite, 1878, from St, Helena (synonym, Igerna Kirkaldy, 1903 [Pachynus Stal, 1866, not Pachynus Bonaparte, 1854 j, type Bythoscopus (Oncopsis) hima- culaticollis Stal, 1855), but that genus has the ocelli close together, and although the crown extends laterally behind the eyes, it is not shortened medially. Xchcla appears to be more closely related to Agallia Curtis. Agalliota, new genus Type of the genus, Agallia punctata Oman, 1934, a Neotropical species. So far as known the genus is limited in distribution to the Neotropical region. 352 The University Science Bulletin Brasa, new genus (PI. XXVIII, fig. 1) Resembling Macropsis Lewis in general form, but more closely related to Agallia Curtis by the venation of the hind wing. Differing from Macropsis in having four apical cells in the hind wing and in having no pronotal striations, and from Agallia and related genera by the position of the ocelli, which are close together, the broad fore- wings, and the "humped" appearance of the thorax. Crown very short and of uniform length. Face slightly convex, frontal sutures distinct, genae broad. Pronotum several times as long as vertex, lateral margins obsolete, disk and posterior portion strongly elevated, anterior margin with depressed areas behind eyes. Scutellum distinctly elevated above forewings, especially anteriorly. Forewing opacjue and subcoriaceous, nearly flat and held in a nearly vertical position as in Macropsis, apex broad and rounded, ajipendix very small or absent, closed anteapical cells two in number. Type of the genus, Macropsis nigicollis Dozier, 1926 (p. 264), known only from the island of Haiti. No other .species known to the writer belong in the genus. Macropsis Lewis Macropsis Lewis, Trans. Ent. Soc. Lond., vol. 1, p. 49, 1830. Pediopsis Burmeister, Gen. Ins., pi. 10, 1838. (Type, Jassus tiUae Gerniar, 1831, a Palearctic species now known to occur in the northeastern part of the United States.) Galboa Distant, Trans. Linn. Soc. Lond., vol. 13, pp. 45-46, 1909. (Type CJalhoa typica Distant, 1909, from Seychelles.) Crown very short, angled medially. Face convex, lori and genae small and indistinctly separated from each other and from front. Pronotum several times as long as vertex, pronotal striae distinct and oblique, converging anteriorly. Scutellum large. Forewing sometimes subcoriaceous, appendix very small or absent. Last ven- tral segment of female with lateral points of attachment very short, posterior margin with only faint notches and few modifications use- ful in specific separation. Male plates long, slender, and lingerlike. Type of the genus. Cicada virescens Gmelin, 1788 (p. 2111), a Palearctic species now known to occur in the Nearctic region also. As here considered the genus is known to be distributed throughout the world with the exception of South America. Typical species are particularly abundant in the Nearctic and Palearctic regions. Gal- boa Distant is included as a synonym on the authority of China, 1925. Oman: Bythoscopinae and Jassinae 353 Oncopsis Burmeister (PI. XXVII, fig. 4) Oncopsis Burmeister, Gen. Ins., pi. 10, 1838. Crown short and broad, not angled as in INIacropsis. Clypeus more convex than in Macropsis and face shorter and broader. Pro- notum with striae transverse, but usually not so coarse as those of Macropsis. Forewings broad apically, appendix absent. Genitalia as in Macropsis. Type of the genus, CicaAa fiavicollis Linnaeus, 1761 (p. 891), a Palearctic species. So far as known the genus is limited in distribu- tion to the Nearctic and Palearctic regions. Neopsis, new genus Intermediate in character between Macropsis Lewis and Bythos- copus Germar; related to the former by the shape of the pronotum and head, and resembling the latter in the venation of the hind wing. Differing from Macropsis in having the inner fork of the radius and the outer fork of the media of the hind wing anastomosing distally, and from Bythoscopus in having the lateral pronotal margins obso- lete and without carinae. Crown short and rounded to the front, subangular medially as in IVIacropsis. Clypellus small, genae comparatively narrow. Prono- tum several times as long medially as median length of vertex, sur- face with numerous fine, nearly transverse striae. Forewing tapered distally, not broad as in Macropsis, venation irregular and difficult to see, the normally central anteapical cell apparently open basally, appendix absent. Genitalia of both sexes differing from the con- ventional Macropsis type. Type of the genus, Pediopsis elegans Van Duzee, 1907 (p. 58), described from Jamaica and known to occur in South America. No other species are referred to the genus. Bythoscopus Germar (PI. XXVII, fig. 2, 2A) Bythoscopus Germar, Silb. Rev. Ent., vol. 1, p. 180, 1833. Batracomorphus Lewis, Trans. Ent. Soc. Lond., vol. 1, p. 31, 1836. (Type, Batracomor- phus irroratus Lewis, 1836, a Palearctic species.) Stragania Stal, Rio Jan. Hemip., vol. 2, p. 49, 1862. (Type, Stragania ornatula Stal, 1862, a Brazilian species.) Gargaropsis Fowler, Biol. Cent. Amer. Homop., vol. 2, p. 167, 1896. (Type, Gargaropsis innervis Fowler, 1896, a Mexican species.) Eurinoscopus Kirkaldy, Hawaiian Sugar Planters' Assoc. Expt. Sta., Div. Ent. Bull. 1, p. 346 1906. (Type, Eurinoscopus lentiginosus Kirkaldy, 1906, an Australian species.) (New synonymy.) 23—6037 354 The University Science Bulletin Crown short and broad, but distinctly longer in proportion to its length than that of most genera of the subfamily. Crown, pronotum, scutellum, and portions of face usually with fine, transverse striae. Face very short and broad, clypellus but slightly longer than broad, lateral margins of genae strongly sinuated. Forewings with surface usually thickly set with fine setae, sometimes subcoriaceous. Male plates usually covered by the unusually large valve. Type of the genus. Cicada lanio Linnaeus, 1761 (p. 892), a Pale- arctic species. The genus appears to be practically cosmopolitan in distribution and is represented in North America by a number of species. South American species referred to the genus, include Stra- gania ornatula Stal, S. divisa Stal, and S. pallescens, Stal, 1862; Macropsis smithii Baker, M. sordidus Baker, and M. stramineus Baker, 1900; Bythoscopus pallidus Osborn and B. nigrifrons Osborn, 1924; and will probably contain some other species described by Stal. The writer has examined the genotype of Eurinoscopus Kirkaldy and is of the opinion that the additional elytral cross veins which he mentions in his description are not sufficient basis for retaining that genus as distinct from Bythoscopus. Pachyopsis Uhler (PI. XXVII, fig. 3) Pachyopsis Uhler, Bull. U. S. Geol. Geog. Surv., vol. 3. p. 466, 1877. Straganiopsis Baker, Invert. Pacif., vol. 1, p. 10, 1903. (Type, Macropsis idioceroides Baker, 1900, which is at present considered to be a s>iionym of Pachyopsis laetus Uhler, 1877.) (New synonymy.) The genus is here removed from synonymy, having previously been considered a synonym of Bythoscopus Germar. Characters of diagnostic value are given below. Closely related to Bythoscopus Germar, with which it has been placed, but differing from that genus in having four apical cells in the hind wing, and in having the face less strongly receding. Characters of the head and pronotum as in Bythoscopus except that the clypellus is broader and the lateral margins of the pronotum are shorter and less diverging posteriorly. Forewing with a small appendix or none, inner apical cell not membraneous as in Bythos- copus. Type of the genus, Pachyopsis laetus Uhler, 1877, a Nearctic species, known to occur only in the southwestern part of the United States. The genus will also contain, in addition to Macropsis laetus var. pactum Ball, 1900, an apparently undescribed species from Panama and two South American species. Oman: Bythoscopinae and Jassinae 355 Key to the South American Genera op Jassinae 1. Second cross vein of forewing joining media anterior to base of central an- teapical cell (PI. XXVIII, fig. 2); apex of forewing rounded, appendix small. Second apical cell of hind wing very narrow. Head extremely short and broad, ocelli distant from the eyes (PI. XXXI, fig. 1), Bythonia, p. 358 Second cross vein of forewing, if present, joining central anteapical cell (except in Idiotettix, which has the forewing subtruncate apically and the appendix very large). Second apical cell of hind wing rarely narrow. Width of head and position of ocelli variable 2 2. (1) Head, thorax, and at least basal portions of forewings with numerous circular pits. Hind wing with apical margin concavely excavated opposite third and fourth apical cells (PI. XXVIII, fig. 3). Color black Niorua, p. 358 Head, thorax, and forewings without distinct circular pits. Apex of hind wing convexly rounded. Color variable, seldom black 3 3. (2) Submarginal vein of hind wing evanescent or joining wing margin opposite apical cells (PI. XIXVIII, fig. 4A). Forewing without appendix, Xestocephaltis, p. 359 Submarginal vein of hind wing present and not joining wing margin apically. Forewing either with or without appendix 4 4. (3) Head and anterior portion of pronotum coarsely and irregularly rugose (PI. XXXI, fig. 7) Arrugada, p. 362 Head and pronotum not as above 5 5. (4) Costal area of hind wing much expanded for a short distance near base (PI. XXX, fig. 9A). Forewing always broad 56 Costal area of hind wing not expanded basally. Forewing either broad of narrow 6 6. (5) Ocellocular area produced into a distinct ledge above the antennal socket, 51 Ocellocular area not produced into a ledge above antennal socket 7 7. (6) Cubitus of forewing appearing forked* near first cross vein (PI. XXVIII, fig. 7). Forewing never with more than one closed anteapical cell and usually only one claval vein Spangbergiella, p. 364 Forewing not as above 8 8. (7) Forewing acuminate distally and without appendix (PI. XXVIII, fig. 6). Crown flat and anterior margin carinate Egenus, p. 363 Forewing not acuminate, or if so, then crown neither flat nor carinate an- teriorly 9 9. (8) Forewing with two closed anteapical cells, the outer short and subtriangular, the central strongly constricted; appendix lacking (PI. XXVIII, fig. 9). Crown broad and flat, with anterior margin carinate Luheria, p. 363 Forewing not as above, or if so, then crown not flat 10 10. (9) Ocelli exactly on crown margin over one third the distance from eyes to median line. Crown flat, anterior margin carinate; lateral margins of pronotum long and carinate Cerrillus, p. 362 Ocelli either near eyes, or crown and pronotum with margins not carinate, or both 11 11. (10) Forewing with only one closed anteapical cell or with none. Appendix al- ways present 47 Forewing with two or more closed anteapical cells. Appendix present or absent 12 12. (11) Clypellus with lateral margins straight and parallel except basally; apex of clypellus extended beyond normal curve of lateral margins of genae (PI. XXXIV, fig. 9A). Apex of head subcorneal 46 Clypellus usually constricted, with margins not parallel, or apex of clypellus not extended beyond normal curve of lateral margins of genae. Apex of head seldom subcorneal 13 13. (12) Lateral margins of genae not sinuated below eyes (PI. XXXII, figs. 3A, 3B); genae visible behind eyes in dorsal view (PI. XXXII, fig. 3). Crown pro- duced and angled ; reflexed veinlets to costal margm of forewing always present (PI. XXIX, fig. 1) Scaphytopius, p 367 Lateral margins of genae usually sinuated below eyes; genae never visible behind eyes in dorsal view. Crown variable; reflexed veinlets to costal margin of forewing either present or absent 14 14. (13) Forewing with numerous ramose lines and false veins 28 Forewing without ramose lines and false veins 15 15. (14) Forewing with two or more reflexed veinlets to the costal margin in the vicinity of the outer anteapical cell 16 Forewing without reflexed veinlets to costal margin 21 • Thia is probably media separating from cubitus. See remarks, page 365. 356 The University Science Bulletin 16. (15) Crown of nearly uniform length, broad, and with a distinct transverse furrow just behind anterior margin. Broad, somewhat flattened species, Bahita, p. 379 Crown much longer medially than next eyes, or, if short, lacking the trans- verse furrow 17 17. (16) Crown flat, anterior margin carinate Hecaloidia, p. 378 Crown not flat, or if so, anterior margin not carinate 18 18. (17) Crown only slightly longer medially than next the eyes Menosoma, p. 379 Crown about twice as long medially as next the eyes 19 19. (18) Forewing with reflexed veinlets to costal margin much broadened; central anteapical cell very narrow distally (PI. XXIX, fig. 3) . . . .Scaphoidula, p. 368 Reflex veinlets to costal margin of forewing not broadened ; central ante- apical cell not extremely narrow distally 20 20. (19) Outer anteapical cell of forewing less than half as long as central anteapical cell (PI. XXIX, fig. 7). Crown not distmctly angled (PI. XXXII. fig. 7) Garapita, p. 369 Outer anteapical cell of forewing over half as long as central anteapical cell and not so broad. Crown angled (PI. XXXII, fig. 6) Osbornellus, p. 369 21. (15) Crown produced, ocelli very large and situated as near to apex of head as to eyes (PI. XXXII, fig. 8). Antennae as long as body Portanus, p. 370 Crown either produced or not, but if produced, ocelli never as above. Antennae of various lengths 22 22. (21) Forewings not reaching beyond third abdominal segment.... Faltala, p. 385 Forewings reaching at least to ninth abdominal segment, usually extending beyond abdomen 23 23. (22) Forewing with only two closed anteapical cells 30 Forewing with three or more closed anteapical cells 24 24. (23) Crown extremely long and angled (PI. XXXII, fig. 9), with a flat disk. Fore- wing narrowed apically from inner margin (PI. XXIX, fig. 9). . .Agudus, p. 371 Crown not as above. Forewing with appendix longer and not narrowed from inner margin 25 25. (24) Outer anteapical cell of forewing divided into 3-5 smaller cells. Apex of forewing subtruncate (PI. XXIX, fig. 6) Kanorba, p. 374 Outer anteapical cell of forewing not divided. Apex of forewing rounded.. 26 26. (25) Head narrower than thorax (PI. XXXIII, fig. 1); apex turned slightly up- ward Cumora, p. 374 Head as wide or wider than thorax ; apex seldom turned upward 27 27. (26) Broad, comparatively flat species. Pygofcr with numerous, stout, closely set setae Sanctanus, p. 371 Not distinctly broad or flat species. Pygofer with setae usually rather sparse Deltocephalus, p. 373 28. (14) Ramose lines and false veins of forewing occurring only in area between radius and cubitus (PI. XXIX, fig. 8) Caphodus, p. 375 Ramose lines and false veins of forewing not limited to area between radius and cubitus 29 29. (28) Head very short and broad ; ocelli large, distant from eyes about three times their own diameter (PI. XXXIII, figs. 5, 5A) Baroma, p. 377 Head not unusually broad ; ocelli near eyes Phlepsius, p. 376 30. (23) Forewing without appendix, inner margin extended in a nearly straight line from tip of clavus to apex 31 Forewing with a distinct appendix, inner margin rounded from tip of clavus to apex 32 31. (30) Crown longer medially than next to eyes; head narrower than pronotum (PI. XXXIV, fig. 5) Acinopterus, p. 388 Crown very short and of uniform length ; head as wide as pronotum (PI. XXXI, fig. 4) Cariancha, p. 360 32. (30) General form broad and flat; crown flat and short with anterior margin thin. Crown and pronotum transversely striated Brincadorus, p. 361 General form occasionally broad but not flat ; crown with anterior margin not thin. Crown and pronotum not striated 33 33. (32) Forewing with only one claval vein, or with second claval vein very close to inner margin. Face very short, in profile swollen between antennae, Onura, p. 387 Forewing with two distinct claval veins. Face neither unusually short nor swollen between antennae 34 34. (33) Crown flat or nearly so, distinctly separated from the face by a subcarinate margin. Species slender 35 Crown not flat, or if so, very short and species robust. Crown not separated from face by a distinct margin 36 Oman: Bythoscopinae and Jassinae 357 35. (34) Outer anteapical cell of forewing petiolate at least basally (PI. XXVIII, fig. 8). Crown with two shallow, transverse furrows Bolarga, p. 366 Outer anteapical cell of forewing truncate basally (PI. XXIX, fig. 4). Crown flat Bonamus, p. 365 36. (34) Crown over twice as long medially as next eyes (PI. XXXTV, fig. 10). Ocelli small and distant from eyes Benala, p. 390 Crown not twice as long medially as next eyes. Ocelli either near eyes or unusually large 37 37. (36) Appendix extending around apex of forewing. Clypellus tapering distally (PI. XXXIII, fig. lOA) Exitmnus, p. 3S2 Appendix not extending around apex of forewing. Clypellus not as above, 38 38. (37) Crown sloping abruptly downward anteriorly; clypeus much swollen in pro- file (PI. XXXIV, fig. lA). First and second apical cells of forewing un- usually large (PI. XXX, fig. 1) Eusceloidia, p. 384 Crown not sloping downward abruptly ; clypeus not swollen as above. Apical cells of forewing of more uniform size 39 39. (38) Head distinctly wider than pronotum. Wedge-shaped leaf hoppers 40 Head about as wide as pronotum or slightly narrower. Not wedge-shaped leaf hoppers 41 40. (39) Outer anteapical cell of forewing narrowed at both ends. Clypellus short and nearly parallel margined Opsins, p. 381 Outer anteapical cell of forewing not narrowed distally. Clypellus extending beyond genae but with apex rounded and basal portion constricted, Alaca, p. 386 41. (39) Frontal sutures extending past ocelli onto crown Hegira, p. 383 Frontal sutures terminating at ocelli 42 42. (41) Pronotum unusually large and humped in profile (PI. XXXIV, fig. 3). Clypellus extended beyond genae Brazosa, p. 386 Pronotum not unusually large and not humped in profile. Clypellus not distinctly extended beyond genae 43 43. (42) Crown comparatively flat and of nearly uniform length, never angular or subangular but always broadly rounded 44 Crown rounded downward, disk not flat; length of crown variable, fre- quently longer or shorter medially than next the eyes 45 44. (43) Outer anteapical cell of forewing broader distally than basally. .. .iSandora, p. 380 Outer anteapical cell of forewing of nearly uniform width Atanus, p. 381 45. (43) Clypellus broad, scarcely or not at all constricted. Usually broad species, color green to brown Chlorotettix, p. 389 Clypellus narrow, usually distinctly constricted. Usually comparatively slender species, often with an angled vertex Thamnotettix, p. 388 46. (12) Forewing with three closed anteapical cells and no appendix. . .Penestirellus, p. 391 Forewing with two closed anteapical cells and usually a distinct appendix, Stirellus, p. 391 47. (11) Basal segment of hind tarsus with a distinct and deep groove in basal third ventrally. Slender, somewhat flattened species 50 Basal segment of hind tarsus without a distinct and deep groove basally. Species not flattened 48 48. (47) Forewing without closed anteapical cells Cortona, p. 390 Forewing with one closed anteapical cell 49 49. (48) Hind wing with four apical cells. Crown produced and angular or sub- angular Baldulus, p. 392 Hind wing with three apical cells. Crown not angular Agelina, p. 392 50. (47) Head wider than pronotum Nesosteles. p. 394 Head narrower than pronotum Baldutha, p. 393 51. (6) Sternal sclerites of mesothorax greatly enlarged, so that anterior coxa and trochanter reach only slightly beyond middle. Pronotum broadest near posterior margin. Appendix of forewing small or absent 55 Sternal sclerites of mesothorax not enlarged. Pronotum broadest just be- hind eyes. Appendix extending around apex of forewing 52 52. (51) Crown conical; median length over twice that next the eye Conala, p. 396 Crown not conical, but broad and short; median length not over one and one half times that next eye 53 53. (52) Forewing with less than three closed anteapical cells 54 Forewing with three closed anteapical cells (PI. XXX, fig. 7). . . .Idiotettix. p. 395 54. (53) First cross vein of forewing absent (PI. XXX, fig. 5). Basal portion of forewing coriaceous Exohdza, p. 395 First cross vein of forewing present. Wings hyaline Neobala. p. 396 358 The University Science Bulletin 55. (51) Hind wing with three apical cells. Crown distinctly produced anteriorly be- yond eyes and distinctly longer medially than next eyes Coelidiana, p. 397 Hind wing with four apical cells. Crown produced anteriorly only slightly beyond eyes Chinaia, p. 397 56. (5) Anterior femora and tibiae broad and foliaceous Gabrita, p. 399 Anterior femora and tibiae neither broad nor foliaceous 57 57. (56) Tibiae of first two pairs of legs flattened Terulia, p. 399 Tibiae of first two pairs of legs not flattened Jassus, p. 398 DESCRIPTIONS OF JASSINE GENERA Bythonia, new genus (PI. XXVIII, fig. 2; PI. XXXI, fig. 1) Intermediate in character between Pachyopsis Uhler and Nionia Ball, but differing from both in having the crown more distinctly separated from the face and in having the second apical cell of the hind wing very narrow. Easily separated from Nionia by the ab- sence of circular punctures such as occur on the head, thorax, and forewings of the species of that genus. Crown very short and broad, slightly shorter medially than next eyes. Face broad and short, ocellocular space with a distinct ledge above antennal socket. Fronotum broad, lateral margins distinctly carinate, surface transversely rugose. Forewing subcoriaceous, ex- cept extreme tip and appendix, which are membranous. Anteapical cells three in number; apical cells four; veins forming outer three apical cells practically parallel. Second cross vein joining media before base of the middle anteapical cell. Hind wing with second apical cell unusually narrow; costal cell distad of hamulus distinct and unusually large. Type of the genus, Nionia (?) rugosa Osborn, 1923 (p. 32), a Bolivian species. No other species which could be referred to this genus are known to the writer. Nionia Ball (PI. XXVni, fig. 3; PI. XXXI. fig. 2) Nionia Ball, Proc. Biol. Soc. Wash., vol. 28, p. 165, 1915. Correctly placed by Ball as a relative of Tartessus Stal, but even more closely related to Thymbris Kirkaldy and Epipsychidion Kir- kaldy, with which it agrees in having an extremely short, broad, and curved crown, a strongly receding face, and no appendix on fore- wing. It differs from the above genera in being shorter and more robust and in having circular pits on the head, thorax, and at least the basal portion of the forewings. Not closely related to any American genus known to the writer. Oman: Bythoscopinae and Jassinae 359 Robust species, the known American forms all black in color, sometimes with apices of forewings reddish-brown. Head wider than thorax, eyes small, crown very short in its entire width, but usually slightly longer medially than at any other point. Ko dis- tinct margin between crown and face. Ocelli distant from the eyes about four times their own diameter. Face rounded, clypellus un- usually broad and tapering apically. Genae broad, lateral margins strongly sinuated. Ocellocular area produced into a distinct ledge over antennal socket. Pronotum long, but lateral margins very short and only weakly carinate. Scutellum small. Forewing cori- aceous, circular pits arranged in rows along veins, anteapical cells three in number, the outer one often open distally. Hind wing with apical margin having a (J -shaped excavation opposite the third and fourth apical cells, this excavation falling on the median line of the dorsum when the wing is folded in its normal position. Type of the genus, Goniagnathus palmeri Van Duzee, 1891 (p. 171), a Nearctic species. The genus is known to be represented in South America by Bytkoscopus postica Stal, 1860, and Nionia major Osborn, A'^. gagatina Osborn, with its variety brunnea Osborn, and N. minor Osborn, 1924. Nionia (?) rugosa Osborn, 1923, has been transferred to the genus Bythonia. As pointed out by Kirkaldy, 1906 (p. 305 and 345), Tartessus Stal and its relatives are in many respects intermediate in character between typical Jassinae and typical Bythoscopinae. Nionia and Bythonia are the American representatives of this group, although Bythonia has less apparent affinities with Tartessus than has Nionia. Xestocephalus Van Duzee (PI. XXVIII, figs. 4, 4A; PI. XXXI, fig. 3) Xestocephalus Van Duzee, Bull. Buffalo Soc. Nat. Sci., vol. 5, p. 214-215, 1894. Related to Acucephalus Germar, with which it agrees in the posi- tion of the ocelli, the absence of an appendix on the forewing, and in having no submarginal vein at the apex of the hind wing. Dif- fering from Acucephalus in that the head is narrower than the tho- rax and the outer anteapical cell of the forewing is small and much basad of its normal position. Kirkaldy placed Xestocephalus as a relative of Phrynomorphus Curtis, but the writer believes that it is properly placed near Acucephalus, as indicated in the original de- scription by Van Duzee. Crown produced and rounded to the front, median length dis- tinctly greater than length next eyes. Eyes small. Ocelli distant 360 The University Science Bulletin from eyes about one third or one half the distance from eyes to apex of head. Face strongly receding, sometimes clypellus and lori somewhat flattened. Lateral margins of genae sinuated. Lateral margins of pronotum short, slightly diverging posteriorly, and car- inate. Forewing hyaline; outer anteapical cell slender, usually lying along basal one half of central anteapical cell, but sometimes ex- tending slightly farther distad ; first apical cell small. Female pygo- fer armed with numerous large setae, ovipositor curved downward posteriorly. Male plates and pygofer armed with either spinelike or feathery setae or both. Type of the genus, Xestocephalus puUcarius Van Duzee, 1894, a Nearctic species. The genus appears to be almost cosmopolitan in distribution, typical representatives being present on all continents of the world except Europe. The entire genus must be carefully worked before the specific status of some of the South American forms can be determined. Members of the genus recorded from South America are: Deltocephalus superbus Provancher, 1890, Xestocephalus puUcarius Van Duzee and X. tessellatus Van Duzee, 1894, X. brunneus Van Duzee, 1907, and X. irroratus Osborn, 1924. Cariancha, new genus (PI. XXXI, fig. 4) Short, broad leaf hoppers resembling Goniagnathus Fieber in gen- eral form but not closely related to any genus known to the writer. Distinguished by the extremely short, broad head with the crown rounded to the front, and the forewings without appendices. Crown very short, about five times as broad as long. Ocelli dis- tant from the eyes about three times their own diameter. Face short and broad; clypellus unusually narrow; lateral margins of genae sinuated below eyes. Pronotum as broad as head, about four times as long as crown, surface faintly and irregularly transversely striated, lateral margins very short and feebly carinate. Forewing tapered to bluntly pointed apex, closed anteapical cells two in num- ber and of approximately equal size. Pygofer with numerous short, stout setae. Type of the genus, Cariancha cariboba, n. sp., from Brazil. Cariancha cariboba, new species General color, shining reddish-brown. Crown pale, with an in- definite transverse band of brown to fuscous above ocelli; base of pronotum and scutellum usually with similarly colored areas. Fore- wing subhyaline, with small whitish spots in cells near apex. Oman: Bythoscopinae and Jassinae 361 Length of female, 6 mm. ; width of pronotum, 2.25 mm. Crown of nearly uniform length throughout its width. Forewing with two or three extra veins to costal margin opposite outer anteapical cell; apex of outer anteapical cell close to costal margin. Last ventral segment of female with posterior lateral angles rounded, posterior margin with a median V-shaped notch. Holotype female and two female paratypes labeled ''Rio," P. R. Uhler collection. No other data. Types in the collection of the United States Museum, Cat. No. 51662. Brincadorus, new genus (PI. XXXI, fig. 5) Related to Stroggylocephalus Flor, which it resembles in general form and shape of the broad, short crown, but differing from that genus and from Acucephalus Germar in that the forewing has a dis- tinct and membranous appendix and the hind wing has a distinct submarginal vein apically. Crown broadly rounded anteriorly, anterior portion and thin an- terior margin with numerous fine, transverse striae. Ocelli distant from eyes about two and one half times their own diameter. Face short and relatively flat, clypellus unusually short and broad, lat- eral margins of genae sinuated below eyes. Pronotum broad, sur- face of posterior portion with fine transverse striae. Forewing short and broad, closed anteapical cells two in number. Type of the genus, Brincadorus laticeps, n. sp., from Brazil and Bolivia. So far as known, this is the only species belonging to this genus. Brincadorus laticeps, new species General ground color yellowish-white. Crown with an indefinite transverse band of reddish-brown ; thorax, forewing, except tip, and dorsum of abdomen washed with the same color. Length of male, 5.75-6 mm. A broad, fiat species. Clypellus swollen, distinctly curved in profile. Posterior margin of pronotum shallowly and angularly excavated; scutellum large. Male valve large, apex bluntly rounded or subtruncate. Plates extending upward and backward around the posterior ventral angle of the pygofer and tapering to sharp, spinelike apices which con- verge behind the pygofer. Aedeagus visible, extending downward and backward from near the apex of the valve. Holotype male and 1 male paratype, Chapada, Brazil, January, C. F. Baker collection. Also 1 male paratype labeled Rurrenabaque, 362 The University Science Bulletin Beni, Bolivia, W. M. Mann, December, 1921, Mulford Biological Exploration, 1921-'22. Types in the collection of the United States National Museum, Cat. No. 51663. Cerrillus, new genus (PI. XXXI, fig. 6) Related to Hecalus Stal by the flat crown with a thin margin, but differing from that genus in having the head, pronotum, and scu- tellum finely rugose rather than granulose and in having the ocelli about half way between the eyes and the apex of the head rather than next the eyes. No obvious sexual dimorphism. Crown over twice as long medially as next the eyes, apex sub- angular. Lateral margins of genae sinuated. Pronotum short and broad, lateral margins carinate and slightly diverging posteriorly, posterior margin broadly, shallowly, and angularly excavated. Fore- wing short, but exceeding abdomen, appendix distinct, closed an- teapical cells two in number. Type of the genus, Hecalus notatus Osborn, 1923 (p. 27), known to occur in Brazil and Bolivia. This is the only species at present included in the genus. Arrugada, new genus (PI. XXVIII, fig. 5; PI. XXXI, fig. 7) Apparently related to Huleria Ball by the venation and angulate head, but differing from that genus in the character of the head and anterior half of the pronotum, which are coarsely and irregularly rugose, and in having the numerous setae on the ventral surface of the basal segment of the hind tarsus not arranged in the normal two rows but distributed over most of the ventral surface. Crown about twice as long medially as next the eyes, apex slightly turned upward; margin between crown and front distinctly carinate or subfoliaceous. Ocelli distant from the eyes about three times their own diameter. Lateral margins of genae incised below the eyes, below incision broadly rounded to apex of clypellus; genae comparatively broad next clypellus. Antennae mounted on a short tubercle which resembles a basal antennal segment. Pronotum large, lateral margins unusually long and distinctly carinate, disk and posterior half faintly, transversely rugose. Forewing subcori- aceous, long and comparatively slender, with a distinct appendix and normally with two closed anteapical and four apical cells. Venation of hind wing normal. Oman: Bythoscopinae and Jassinae 363 Type of the genus, Huleria rugosa Osborn, 1924 (p. 404) , a Bo- livian species. The genus will also include Huleria affinis Osborn, 1924, from the same region. Luheria Osborn (PI. XXVIII, fig. 9; PI. XXXI, fig. 8) Luheria Osborn, Ann. Cam. Mus., vol. 15, no. 1, p. 31, 1923. Resembling Selenocephalus Germar in the shape of the short, broad crown, but differing from that genus in having the outer anteapical cell of forewing roughly triangular and the central an- teapical cell constricted near distal end. In the opinion of the writer this genus is not closely related to Huleria Ball, near which it was placed by Osborn, since Huleria has an elongate and subfoliaceous crown, the ocelli are distant from the eyes, and the anteapical cells of the forewing are slender and parallel-sided. Crown but little longer medially than next the eyes, anterior mar- gin broadly and evenly rounded, with a few faint transverse striae. Ocelli distant from eyes slightly more than twice their own diameter. Face broad, clypellus constricted near base, lateral margins of genae evenly curved except for a faint sinuation below each eye. Pro- notum broad and nearly flat, lateral margins faintly carinate. Fore- wing broad, scarcely tapered distally, lacking a distinct appendix, claval veins connected by one or two cross veins, apical cells very short. . Type of the genus, Luheria constricta Osborn, 1923, described from Brazilian specimens and now known to occur also in northern Argentina. No other species belonging to the genus are known to the writer. Egenus, new genus (PI. XXVIII, fig. 6; PI. XXXI, fig. 10) Most closely related to Parabolocratus Fieber, with which it agrees in having two elongate anteapical cells in the forewing, but differing from that genus in that the forewing is acuminate and the front is swollen. Differing from Dorycephalus Kuschakewitch, with which it agrees in the form of the forewing, in having the crown much shorter and the margin not foliaceous, and in the structure of the pronotum. Crown produced and broadly rounded anteriorly, the anterior margin thin and turned upward, leaving a semicircular concave area anterior to the ocelli. Ocelli about one and one half times their 364 The University Science Bulletin diameter from the eyes. Clypeus strongly convexly rounded between antennae when seen in profile. Pronotum short, lateral margins short and carinate, carina curved slightly upward posteriorly in lateral view. Forewing long, appendix absent, apical and anteapical cells slender. Ovipositor sheath not extending beyond the short pygofer. Type of the genus, Egenus acuminatus, n. sp., from Argentina. This is the only species included in the genus at present. Egenus acuminatus, new species A large, pale-yellow species without definite markings but prob- ably green in life, as are most species of Parabolocratus. Length of female, 8.5 mm. Crown slightly wider than long, me- dian length over twice that next the eye and equal to median length of pronotum. Pronotum broad, but slightly narrower than head, posterior portion with shallow transverse rugae. Costal margin of forewing but slightly curved ; forewing tapered beyond clavus about equally from each margin. Last ventral segment of female distinctly longer than preceding segment, posterior margin with a faintly incised median lobe, lateral angles rounded. Pygofer short, but extending slightly beyond ovi- positor sheath. Holotype female from Loreto, Prov. Missiones, Argentina, Dec. 14, 1931, A. A. Ogloblin. Type in collection of the Utiited States National INIuseum, Cat. No. 51664. Spangbergiella Signoret (PI. X:SVIII, fig. 7; PI. XXXII, figs. 1, lA) Spangbergiella Signoret, Ann. Ent. Soc. France, ser. 5, vol. 9, pp. 273-274, 1879. BergieUa Baker, Can. Ent., vol., 29, pp. 157-158, 1897. (New synonymy.) Elongate, somewhat flattened leaf hoppers; the crown distinctly produced medially and the margin always foliaceous in the females and frequently so in the males; the pronotum broad and short, with lateral margins long and distinctly carinate. Crown extremely variable in length, always twice, usually several times as long medially as next eyes ; margin of crown varying from foliaceous to bluntly rounded ; disk of crown usually flat or slightly convex. Ocelli on narrow margin of crown, next to or near eyes. Pronotum with lateral margins nearly straight, either nearly parallel or slightly diverging posteriorly from eyes. Forewing long and rather slender, appendix very narrow, usually not distinctly mem- Oman: Bythoscopinae and Jassinae 365 branous; claval veins usually one, but occasionally two, in number; first cross vein rarely present and then very short; usually entirely absent and the media joining cubitus for a short distance; one ante- apical cell present, the base of this pointed. Ovipositor sheath ex- tending beyond pygofer. Type of the genus, Glossocratus vulneratus Uhler, 1877 (p. 464) , a North American species. The genus occurs only in America so far as known. South American species referred to the genus include Spangbergiella lacerdae Signoret, 1879, Hecalus lynchii Berg, 1879, Spangbergiella punctato-guttata Berg and S. felix Berg, 1884, and Parabolocratus uruguayensis Berg, 1884. The writer has been un- able to examine either types or authentically determined specimens of P. uruguayensis Berg, upon which Baker based his genus Berg- iella, but finds nothing in either Berg's or Baker's characterizations which would seem to justify separating Bergiella from Spangberg- iella. The status of lynchii is in doubt, having been placed as a synonym of vulnerata Uhler by Signoret, 1880, Berg, 1884, and Ball, 1899 ; however, since Signoret's figures of S. lacerdae, August, 1879, from Bahia, Brazil, scarcely agree with vulnerata Uhler, it seems more probable that lacerdae Signoret is a distinct species and has as a synonym lynchii Berg, December, 1879. Lawson, 1932 (p. 116), has characterized Spangbergiella as hav- ing ". . . inner branch of first sector (of forewing) forking well cephalad on disc of corium, radius three-branched resulting in five apical cells . . .," an interpretation of the venation which the writer believes unwarranted, since specimens are occasionally found in which the venation at the base is of the normal type except that the cross vein is extremely short. The writer's interpretation of the venation is given in the description of the genus. Bonamus, new genus (PI. XXIX, fig. 4; PI. XXXI, fig. 9) Related to Spangbergiella Signoret, but with crowm margin never foliaceous and forewing with a distinct appendix and two anteapical cells. Crown nearly flat, distinctly longer medially than next the eyes, not angled anteriorly, anterior margin not thin or foliaceous. Face rather flat, lateral margins of genae strongly sinuated below eyes. Pronotum short, lateral margins short and only weakly carinate. Forewing long and slender, anteapical cells lying about parallel to claval suture, the outer cell shorter than the central one. Ovipositor 366 The University Science Bulletin sheath long, extending beyond slender pygofer for about one third its total length. Genital segments of both sexes with numerous large setae. Type of the genus, Bonamus lineatus, n. sp., from Argentina. A second species, from the same locality and apparently undescribed, will also be included in the genus. Bonamus lineatus, new species A pale blue-green to yellowish-white species with narrow dor- sal, longitudinal lines of orange-red. Crown usually whitish, some- times suffused with blue-green, with three parallel stripes, one next each eye and one in the middle, these continued across pronotum and scutellum. Pronotum with an additional pair of stripes, one behind each eye, these continued across middle of clavus of each forewing and meeting on commissural line. Forewing subhyaline, suffused with blue-green to yellowish-white, marked with orange-red along commissural line, across middle of clavus, along outer margin of claval suture, and in cells of corium basad of apical cross veins, these markings less distinct than the dorsal ones. Length, 5-6 mm. Crown as long as basal width, median length over twice that next eye and equal to median length of pronotum. Pronotum slightly narrower than head, posterior margin broadly and shallowly excavated. General form, slender. Male valve elongate and triangular, plates together somewhat spoon-shaped, ventral surfaces covered with long setae. Last ven- tral segment of female long, median portion broadly notched about half way to base of segment, with a blunt toothlike projection at base of notch. Pygofer and distal third of ovipositor sheath with numerous long setae. Holotype male, allotype female, and 3 male and 10 female para- types from Loreto, Prov. Missiones, Argentine, collected by A. A. Ogloblin in 1931, the holotype on December 6, the allotype on De- cember 1, and the paratypes on various dates from November 27 to December 14. Types in collection of the United States National Museum, Cat. No. 51665. Bolarga, new genus (PI. XXVIII, fig. 8 ; PI. XXXII, fig. 2) Resembling Laevicephalus De Long in general appearance, but apparently not closely related to the Deltocephalus group. Similar to OshornellUsS Ball in the shape of the anteapical cells of the fore- wing, but lacking the reflexed veinlets to the costal margin. Oman: Bythoscopinae and Jassinae 367 Crown produced and siibangular, disk with two broad, shallow, transverse furrows, one between the eyes and the other just in front of the ocelli. Margin between crown and face subcarinate. Ocelli near eyes. Face comparatively fiat; genae broad and lateral mar- gins broadly notched below the eyes. Pronotum short, lateral mar- gins not carinate. Forewing with a distinct appendix, second cross vein absent, outer anteapical cell broader distally, central anteapical cell strongly constricted near apex, fourth apical cell unusually small, costal area with a cross vein to costal margin at each end of outer anteapical cell. Pygofer with numerous stout setae which are arranged in rather definite groups. Type of the genus, Parabolocratus holivianus Osborn, 1923 (p. 32), from Bolivia. An apparently undescribed species from nothern Argentina also belongs here. The writer considers holivianus variety 'pallida Osborn, 1923, unworthy of varietal rank and places it as a synonym of holivianus. (New synonymy.) Bolarga is not closely related to Parabolocratus, where its type species was placed by Osborn, and is easily separated from that genus by the shape of the cells of the forewing. Scaphytopius Ball (PI. XXIX, fig. 1 ; PI. XXXII, figs. 3, 3A, 3B) Scaphytopius Ball, Can. Ent., vol. 63, p. 218, 1931. Easily separated from other genera of the Jassinae by the flat crown, broad genae which extend up behind the eyes and are visible from above, and the reflexed veinlets in the costal area. Crown produced and angled, disk flat ; margin between crown and face distinct, often carinate. Ocelli near eyes. Front very narrow and elongate, distinctly constricted just beyond base of clypellus. Genae broad, lateral margins not sinuated. Pronotum short, slightly wider than head; lateral margins short, diverging posteriorly and subcarinate. Forewing with a distinct appendix, numerous reflexed veinlets to the costal margin opposite outer anteapical and outer discal cells, and usually three, but sometimes only two, closed ante- apical cells. Type of the genus, Platymetopius elegans Van Duzee, 1890 (p. 94), a Nearctic species. The genus is known to occur only in the Nearctic and Neotropical regions. In addition to the species re- ferred to Scaphytopius by Ball, 1932 (p. 252), the genus as above defined will include the species placed in Deltopinus Ball (type, Platymetopius nigriviridis Ball, 1909), Cloanthus Ball (type, Platy- 368 The University Science Bulletin metopius angustatus Osborn, 1905) , Platymoideus Ball (type, Platy- metopius trilineatus Ball, 1916), Nasutoideus Ball (type, Playtme- topius nasutus Van Duzee, 1907), and Convelinus Ball (type, Platy- metopius nigricollis Ball, 1916), those groups being here considered as subgenera of Scaphytopius. In making a phylogenetic arrange- ment of the species the writer has found these segregates very useful, but is of the opinion that the differences in markings and the sexual dimorphism exhibited by certain groups are an insufficient basis for the establishment of genera. These groups are separated in a key by Ball, 1931 (p. 217). Species of Scaphytopius recorded from South America are: Jas- sus (Deltocephalus) marginelineatus Stal, 1858; Jassus {Delto- cephalus) anticus Stal, 1860; Platy metopius loricatus Van Duzee, 1894; Platymetopius fuliginosus Osborn and P. sulphureus Osborn, 1923; and Scaphytopius bolivianus (new name for Platymetopius lineolatus Osborn, 1923, not Platymetopius lineolatus Motschulsky, 1859). Scaphoidula Osborn (PI. XXIX, fig. 3; PI. XXXII, fig. 4) Scaphoidula Osborn, Ann. Cam. Mus., vol. 15, no. 1, p. 41, 1923. Related to Scaphoideus Uhler by the shape of the outer anteapical cell of the forewing, but with the crown margin more rounded, the refiexed veinlets to costal margin of forewing much broadened and not situated anterior to base of outer anteapical cell, and with the large setae near the tip of the pygofer not arranged in groups as is typical of Scaphoideus. Crown about twice as long medially as next the eye, apex bluntly angled, disk convex, margin between crown and face rounded. Head about as wide as pronotum. Ocelli comparatively large and situated next the eyes. Lateral margins of genae sinuately curved, genae very narrow next clypeus. Pronotum longer than vertex, lateral margins short and faintly carinate, posterior margin broadly and shallowly excavated between basal angles of scutellum. Forewing slender, with a distinct appendix and two closed anteapical cells, the outer about half as long as the second and not lying parallel to costal margin, the second narrowed apically. Antenna about half as long as forewing. Type of the genus, Scaphoidula cingulata Osborn, 1923, a Neo- tropical species. Three closely related species from Central America and northern South America are also referred to this genus. Oman: Bythoscopinae and Jassinae 369 Osbornellus Ball (PI. XXXII, fig. 6) ' Osbornellus Ball, Jl. Wash. Acad. Sci., vol. 22, no. 1, p. 17, 1932. Closely related to Scaphoidula Osborn by the reflexed veinlets to the costal margin and the shorter outer anteapical cell of the fore- wing, but differing from that genus in that the outer anteapical cell of the forewing lies nearly parallel to the costal margin and the re- flexed veinlets to the costal margin are not broadened. Head and thorax as in Scaphoidula ; head often slightly narrower than pronotum. Second anteapical cell of forewing not narrowed apically, costal area occasionally with a reflexed veinlet anterior to base of outer anteapical cell. Type of the genus, Scaphoideus auronitens Provancher, 1889 (p. 277), a Nearctic species. South American species belonging to the genus are Scaphoideus hyalinus Osborn, S. affinis Osborn, S. julvom- aculatus Osborn, and S. mexicanus Osborn. Ball, 1932, placed mexi- canus Osborn as a doubtful synonym of Portanus stigmosus (Uhler), but the cotype of mexicanus in the United States National Museum collection is an Osbornellus closely related to consors Uhler. A specimen identical specifically with this cotype was sent to Mr. J. K. Knull, curator of insects at Ohio State University, who kindly com- pared it with the three cotj'pes in the collection there and reported it to be the same. It appears, therefore, that mexicanus is properly placed in the genus Osbornellus. Garapita, new genus (PI. XXIX, fig. 7; PI. XXXII, fig. 7) Related to Scaphoidula Osborn and Osbornellus Ball in having a shorter outer anteapical cell and reflexed veinlets in the costal area of the forewing, but differing from those genera in that the outer anteapical cell is even smaller, the crown is shorter and less angled, and the head is distinctly wider than the pronotum. Crown relatively flat, margin between crown and face rounded, median length about twice that next the eyes. Ocelli near eyes. Clypellus unusualh' broad and convex; genae narrow, lateral mar- gins sinuated below the eyes. Lateral margins of pronotum short, not distinctly carinate. Forewing long and slender, appendix dis- tinct, closed anteapical cells two in number, the outer petiolate and distinctly less than half as long as the central cell, the latter not so 24—6037 370 The University Science Bulletin narrow distally as in Scaphoidula, reflexed veinlets to costal margin usually three in number and situated in vicinity of outer anteapical cell. Pygofer with numerous stout setae. Type of the genus, Garapita garbosa, n. sp., from northern Argen- tina. A closely related species from the same locality also belongs here. Garapita garbosa, new species Body below yellow except for fuscous marks on abdomen; above black with longitudinal stripes of yellow. Crown marked with yel- low as follows: Narrow anterior margin, three small spots on disk marking corners of a triangle with the apex anterior, and a dash near each ocellus, all these spots frequently minute or absent in the male, and a small, triangular basal area which is the beginning of a broad median stripe across pronotum and scutellum. Forewing black to fuscous except for narrow yellow marks along commissural line, claval veins, claval suture, and media, and the hyaline costal area and inner apical cell, the hyaline costal area being opposite the anteapical cells and interrupted by the black veinlets to costal mar- gin. Length, 4.25-5 mm. Crown bluntly angled in female, less pro- duced in male. Pronotum longer than crown, posterior margin broadly and shallowly concave. Forewing broad apically, apical cells unusually large. Male valve small and triangular, plates slender and elongate, tapering rather gradually to the somewhat acuminate tips. Last ventral segment of female much longer than preceding segment, posterior margin truncate or slightly produced medially, subcarinate medially on posterior third. Holotype male, allotype female, and 4 male and 6 female para- types from Loreto, Prov. Missiones, Argentina, collected in 1931 by A. A. Ogloblin. The holotype was taken on December 10, the allo- type on December 14, and the paratypes on various dates from No- vember 29 to December 18. Types in the collection of the United States National Museum, Cat. No. 51666. Portanus Ball (PI. XXXII, fig. 8) Portanus Ball, Jour. Wash. Acad. Sci., vol. 22, no. 1, p. 18, 1932. Resembling Osbornellus Ball in general form, but differing in that the ocelli are larger and distant from the eyes, the antennae are longer, and the two anteapical cells of the forewing are of nearly equal size. Oman: Bythoscopinae and Jassinae 371 Crown about twice as long medially as next the eye, apex rounded, not angled as in Osbornellus. Margin between crown and face rounded. Ocelli unusually large, situated on anterior margin of crown at about midpoint between eyes and apex of head. Antenna as long as or longer than wings, basal segments large; antennal socket unusually deep. Forewing with two closed anteapical cells which are nearly parallel-sided and of about equal length, veinlets to costal margin not reflexed; a distinct appendix absent, and with the veins usually intermittently marked with white. Type of the genus, Scaphoideus stigmosus Uhler, 1895 (p. 77), a West Indian species. South American species referable to the genus are Scaphoideus boliviensis Baker (S. bicolor Osborn, 1923, nee S. bicolor Ball, 1909), S. hasemani Baker [S. punctulatus Osborn, 1923, nee S. punctulatus Melichar, 1903), and *S. longicornis Osborn. Sanctanus Ball (PI. XXIX, fig. 2; PI. XXXir, fig. 5) Sanctanus Ball, Jour. Wash. Acad. Sci., vol. 22, no. 1, p. 10, 1932. Related to Amplicephalus DeLong by the structure of the head, but differing from that genus in that the second cross vein of the forewing is alwaj^s present, the second anteapical cell is always con- stricted and divided, and there is usually a slightly reflexed vein to the costal margin from near the base of the outer anteapical cell. Crown relatively flat, anterior margin usually produced and angled medially, sometimes only slightly longer medially than next eyes, median length nearly equal to or less than median length of pronotum. Ocelli near eyes. Front relatively slender; lateral mar- gins of genae sinuated below eyes. Lateral margins of pronotum very short. Forewing relatively large, frequently with extra short veins in clavus and anteapical cells; appendix distinct, reaching to apex of forewing. Pygofer with numerous large setae, sometimes ar- ranged in rather indefinite groups, never in compact groups as in Scaphoideus Uhler. Type of the genus, Jassus sanctus Say, 1831 (p. 307), a Nearctic species. Representatives of the genus known to occur in South America are Scaphoideus fasciatus Osborn, 1900, Jassus {Delto- cephalus) ornatipennis Stal, 1858, and Jassus (Deltocephalus) lepi- dellus Stal, 1860. ^j Agudus, new genus (PI. XXIX, fig. 9; PI. XXXII, fig. 9) ^gf^^. Resembling Platymetopius Burmeister (sensus strictus) in that the crown is long and angled, but more closely related to certain 372 The University Science Bulletin species now referred to Sanctanus Ball in the characters of the ve- nation. Differing from Platymetopius and Sanctanus in having the crown distinctly longer than pronotum and flat on the disk. Crown produced and angled, anterior margin not carinate except near apex. Face convex, clypellus unusually broad, lateral margins very short. Forewing long and slender, tip curved away from abdo- men, appendix small; inner margin of forewing tapering to apex more sharply than outer margin, closed anteapical cells normally three in number, the central one much larger than either the inner or the outer. Pygofer of both sexes armed with numerous long setae. Type of the genus, Agudus typicus, n. sp., from Argentina. Four other species, represented by specimens from northern Argentina and Paraguay, also belong to this genus. One of these species is un- doubtedly Platynietopius longiceps Berg, 1879, as the specimens fit his description exactly except that they are not "densissime punctu- latus" and do not have the face "distincte punctato," but are finely granulose, Agudus typicus, new species General ground color pale yellowish-white. Crown washed with smoky-brown except for a narrow median line from apex to median suture, and irregular lines on crown margin anterior to ocelli; the pale lines margined with fuscous, especially at apex of crown and outer margins of parallel basal stripes. Face washed with brownish- fuscous except for a narrow and frequently interrupted median line, six or seven narrow transverse lines which do not reach median line, and a broad stripe just below crown margin. Pronotum with seven faint brown longitudinal stripes, deflexed lateral margins marked with contrasting fuscous-brown and yellow. Scutellum unevenly washed with yellowish-brown. Forewing with costal margin on basal half bright yellowish-white, remainder subhyaline; veins pale, faintly margined with fuscous; cells faintly embrowned, with four fuscous spots, one on clavus near claval suture, one at base of inner discal cell, one at base of central anteapical, and one along inner margin of inner anteapical cell. Length of female, 5.25 mm. Ratio of median length of crown to median length of pronotum, 2^ to II/2 ; ratio of length of crown to shortest interocular space, 2^/2 to ly^. Lateral margins of crown nearly straight, curved slightly at apex. Last ventral segment of female with a longitudinal median carina, posterior margin broadly, concavely excavated. Oman: Bythoscopinae and Jassinae 373 Holotype female and 2 female paratypes from Loreto, Prov. Mis- siones, Argentina, collected in 1931 by A. A. Ogloblin, the holotype on December 1, the paratypes on December 6. Types in the collec- tion of the United States National Museum, Cat. No. 51667. Deltocephalus Burmeister Deltocephalus Burmeister, Genera Ins., vol. 1, pi. 14, 1838. Small, frequently short leaf hoppers, grass-feeding so far as known. Forewing usually with three anteapical cells. Crown produced and frequently angled, frequently flat and with a distinct margin anteriorly, but never separated from the face by a carina. Ocelli next eyes. Lateral margins of genae sinuated. Fore- wing either long or short. Type of the genus, Cicada pulicaria Fallen, 1826 (p. 34) , a Pale- arctic species now known to occur in the Nearctic region also. The genus as here defined is world wide in distribution and contains a great diversity of forms, many groups of which have been segre- gated as subgenera. The South American species are too numerous to review here. The typical forms of the genus may be separated from the other two South American subgenera by the following key: 1. Crown distinctly angled. Species unusually long and slender Haldorus, Crown produced but not distinctly angled. Species not unusually slender. ... 2 2. (1) Width of crown greatly exceeding median length. Large species. . .Awplicephalus, Width of crown about equal to median length. Smaller species. . .Deltocephalus, Deltocephalus subgenus Amplicephalus De Long Deltocephalus subgenus Amplicephalus De Long, Ohio State University Studies, vol. 2, p. 83, 1926. The characters for the separation of this subgenus are indicated in the key. Type of the subgenus, Deltocephalus osborni Van Duzee, 1892c (p. 304), a North American species. Deltocephalus subgenus Haldorus, new subgenus Differing from the typical Deltocephalus in being more elongate, slender, with a flat crown and pointed head. Forewing with central anteapical cell nearly always constricted and divided. Type of the subgenus, Thamnotettix venatus Osborn, 1924, a South American species. The subgenus will also include about twenty-five closely related South American species, most of them apparently undescribed. 374 The University Science Bulletin Cumora, new genus (PI. XXXIII, fig. 1) Related to the Deltocephaliis group, with which it agrees in ve- nation, but differs in having the crown more angular and turned up- ward apically. Crown about twice as long medially as next the eyes, anterior margin not carinate, but crown distinctly separated from face. Ocelli next eyes. Face relatively flat, lateral margins of genae broadly sinuated. Head slightly narrower than pronotum. Pronotum short, lateral margins not carinate. Forewing long and slender, appendix distinct, central anteapical cell constricted and divided. Type of the genus, Cumora angulata, n. sp., from northern Argen- tina. A second species from the same locality is also referred to this genus. Cumora angulata, new species Face fuscous with numerous pale areas. Ground color of dorsal regions tawny. Anterior margin of crown with four small fuscous spots, pronotum with four longitudinal reddish stripes, and veins of forewing milky-white. Length, 5 mm. Clavus with veins usually connected. Male valve comparatively large; plates small and tapered to blunt tips. Last ventral segment of female short, with a single median tooth on posterior margin. Holotype male, allotype female, and 4 female paratypes from Loreto, Prov. Missiones, Argentina, collected in 1931 by A. A. Oglo- blin, the holotype on December 10, the allotype on November 27, and the paratypes on various dates from December 1 to December 18. Types in the collection of the United States National Museum, Cat. No. 51668. Kanorba, new genus (PI. XXIX, fig. 6; PI. XXXIII, fig. 2) Resembling Scaphoideus Uhler in general appearance, but with disk of crown concavely excavated, outer anteapical cell of forewing divided into several small cells, and with reflexed veinlets to costal margin opposite outer discal cell of forewing. Crown produced and angled, margin between crown and face dis- tinct but not carinate. Ocelli near the eyes. Face broad and com- paratively flat; clypellus broad distally, constricted just beyond base; genae broad, lateral margins slightly sinuated. Pronotum short, lateral margins not carinate. Forewing broad, subtruncate Oman: Bythoscopinae and Jassinae 375 apically, appendix distinct, claval veins connected by a cross vein, second cross vein present, costal area with two reflexed veinlets to costal margin opposite outer anteapical cell and three or four oppo- site the outer discal cell. Pygofer with numerous stout setae. Type of the genus, Kanorba reflexa, n. sp. No other species are referred to the genus. Kanorba reflexa, new species General ground color creamy-white. Face washed with pale brown. Crown with an irregular brown area on the disk with ex- tensions to the anterior margin next the ocelli and median line. Pronotum with four broad, indefinite, longitudinal brown stripes; scutellum with brown areas basally. Forewing subhyaline brown with numerous hyaline areas ; tip and reflexed veinlets to costal mar- gin fuscous. Length, 5-5.25 mm. Crown with apex slightly turned upward, other structures as given for the genus. Male valve short and triangular. Plates together broad basally and tapered to pointed tips, the two plates not touching at any point along median line. Last ventral segment of female with a faint median notch on truncate posterior margin. Holotype male, allotype female, and 1 female paratype from Co- rumba, Brazil, C. F. Baker collection, the holotype and allotype labeled "March" and the paratype "May." Types in the collection of the United States National Museum, Cat. No. 51669. Caphodus, new genus (PI. XXIX, fig. 8; PI. XXXm, fig. 4) Resembling Osbornellus Ball in general habitus, and related to that genus in the structure of the head, but allied to Phlepsitis Fieber by the presence of ramose lines and false veins in the forewings. Differing from Osbornellus by the presence of the false veins and from Phlepsius by the shape of the anteapical cells. Crown produced and angled as in Scaphoidula Osborn, the margin between the crown and face distinct but not angled. Ocelli near eyes. Face and pronotum as in Scaphoidula. Forewing with a dis- tinct appendix, outer anteapical cell broader distally and with a reflexed veinlet to costal margin from each end, central anteapical cell narrowed medially. Second cross vein usually absent, ramose lines and false veins restricted to area between radius and cubitus. Pygofer with numerous large setae. 376 The University Science Bulletin Type of the genus, Caphodus maculatus, n. sp. The genus will also contain Scaphoideus obliquus Osborn, 1923, from Brazil. Caphodus maculatus, new species Closely related to obliquus Osborn, differing in being slightly smaller, having a less flattened crown, shorter and more tapered male plates, and with the posterior margin of the last ventral seg- ment of the female excavated instead of produced. General ground color yellowish-white. Crown with four small fuscous spots on anterior margin and a pair of large orange-red spots on the disk. Face fuscous, interrupted on clypeus with numerous short, transverse, pale marks. Pronotum with a pair of orange-red spots basally and four faint orange spots on disk, the latter some- what elongate and resembling traces of longitudinal vittae. Scutel- lum with indefinite orange marks basally. Forewing with a distinct fuscous spot on clavus next claval suture, smoky brown coloration on most of clavus except three opalescent dashes next commissural line, a fuscous spot on corium between media and cubitus anterior to base of central anteapical cell, and another fuscous area at apex of central anteapical cell following outer apical vein to wing margin. Veins and false veins mostly brown to fuscous, the two reflexed vein- lets to costal margin black. Length, 5.25-5.75 mm. Structurally this species is identical with obliquus except for the proportionally smaller size. Male valve broad basally, triangular in shape. Plates broad bas- ally, but tapering gradually to pointed tips, margins with a row of large setae and above these a row of fine hairlike setae. Last ven- tral segment of female long, with a median carina which ends in a blunt median tooth on posterior margin; posterior margin broadly excavated and with a caudal projection at each side which curves slightly toward the median line. Holotype male, allotype female, and 10 male and 2 female para- types from Loreto, Prov. Missiones, Argentina, collected in 1931 by A. A. Ogloblin, the type on December 18, the allotype on December 8, and the paratypes on various dates from November 25 to Decem- ber 18. Types in the collection of the United States National Mu- seum, Cat. No. 51670. Phlepshis Fieber Phlepsius Fieber, Verh. Zool.-Bot. Ges. Wien, vol. 16, p. 503, 1866. Relatively large, frequently robust species, with numerous ramose lines and false veins in the forewings. Color usually some shade of brown or near brown. Oman: Bythoscopinae and Jassinae 377 Crown of varj'ing lengths and shapes, either margined or rounded to the front. Face usually broad, lateral margins of genae sinuated. Lateral margins of pronotum not at all or only feebly carinate. Forewing broad, appendix distinct, closed anteapical cells either two or three in number. Costal area frequently with reflexed veinlets to costal margin in South American species. Type of the genus, Jassus intricatus H,errich-Schaeffer, 1838, a Palearctic species. South American species referred to the genus include Phlepsius lacerdae Signoret, 1879; Phlepsius gracilis Osborn, hasemani Osborn, saranensis Osborn, sparsus Osborn, loricatus Os- born, annulatus Osborn, clypeatus Osborn, and signatus Osborn, 1923; and Eutettix punctatus Osborn, 1923. The genus as here defined contains a great diversity of forms, and careful study of all of the species of the group is needed to de- cide the status of the folloTvang segregates which have been estab- lished in the North American fauna and are at present considered as subgenera: Paraphlepsius Baker, type Paraphlepsius ramosus Baker, 1897; lowanus Ball, type Phlepsius (lowanus) handlirschi Ball, 1918; Texananus Ball, type Phlepsius (Texananus) mexicanus Ball, 1918; Dixianus Ball, type Phlepsius utahnus Ball, 1909; Zion- inus Ball, type Phlepsius extremus Ball, 1901; and Pendaru^ Ball, type Phlepsius slossoni Ball, 1905. B aroma, new genus (PI. xxxni, figs. 5, 5A) Resembling Idiotettix Osborn in the shape of the short, broad head and elongate form, but differing from that genus in having no ledge above the antennal socket, and in the small size of the appendix of the forewing. More closely related to Phlepsius Fieber, with which it agrees in having numerous ramose lines and incomplete cross veins in the forewing, but differing from Phlepsius by the extremely short, broad head. Differing from Remadosus Osborn and Ball in the shape of the facial sclerites and in the structure of the forewing. Head broader than pronotum ; crown about seven times as broad as long, broadly curved anteriorly, of uniform length and rounded to the face. Ocelli large, distant from the eyes about three times their own diameter. Face nearly flat, front strongly constricted at base of clypellus ; genae broad, lateral margins broadly rounded from be- low eyes. Pronotum broad and rather short, disk faintly, trans- versely striated, lateral margins short and faintly carinate. Scutel- lum unusually large. Forewing long, venation irregular, but with three closed anteapical cells and several extra cross veins to costal 378 The University Science Bulletin margin; appendix distinct, but not extending around apex of wing as in Idiotettix. Pygofer short and broad, without large setae. Type of the genus, Baroma reticulata, n. sp., from northern Argen- tina. This is the only species belonging to the genus known to the writer. Baroma reticulata, new species General ground color pale yellowish-white, often with a bluish- green tint. Anterior and middle pairs of legs, front, and entire dor- sal surface mottled with brown, deepened to fuscous on apex of fore- wing; head less heavily marked with brown. Length, 7.75 mm. ; width of head, 2.3 mm. Pronotum evenly con- vex, without depressed areas. Inner apical cell rather large, remain- ing cells slender. Male valve broad and triangular, plates elongate, tapering to pointed tips, pygofer very broad and flattened dorsoventrally, easily visible beyond plates laterally. Last ventral segment of female with posterior margin concavely excavated from lateral angles and with a narrow median V-shaped notch. Holotype male, allotype female, and 1 male paratype from Loreto, Prov. Missiones, Argentina, collected by A. A. Ogloblin in 1931, the holotype on December 18, the allotype and paratype on December 12. Types in collection of the United States Museum, Cat. No. 51671. Hecaloidia Osborn (PI. XXIX, fig. 10; PI. XXXIII, fig. 3) Hecaloidia Osborn, Ann. Cam. Mus., vol. 15, no. 1, p. 28, 1923. Resembling certain species of the genus Phlepsiu^ Fieber, but dif- fering from that genus in lacking ramose lines in the forewings. Related to Mesamia Ball by the reflexed costal veinlets. Appar- ently not closely related to either Hecalus Ball or Huleria Ball, near which it was placed by Osborn, since reflexed veinlets in the costal area of the forewings do not occur in those genera. Crown flat or slightly concave; anterior margin thin, but not foliaceous, with faint transverse striae. Length of crown on me- dian line, about twice that next eye; apex subangular. Ocelli situ- ated next the eyes. Face broad and flat, nearly straight in profile; lateral margins of genae strongly sinuated below eyes, genae very narrow next clypellus. Pronotum slightly longer than crown, lat- eral margins short and faintly carinate. Forewing elongate, with Oman: Bythoscopinae and Jassinae 379 distinct appendix, two closed anteapical cells and a few reflexed veinlets in costal area, some of these anterior to the base of the short outer anteapical cell. Type of the genus, Hecaloidia nervosa Osborn, a Bolivian spe- cies. No other species belonging to this genus are known to the writer. Menosoma Ball (PI. xxxm, fig. 6) Menosoma Ball, Florida Ent., vol. 15, p. 4-5, 1931. Resembling Eidettix Van Duzee in general habitus. Differing from that genus in having the crown more produced medially and rounded to the face, the transverse furrow in the crown absent or very faint, and usually a few reflexed veinlets to the costal margin of the forewing near the outer apical cell. Menosoma also lacks the distinct striae which occur on the crown and pronotum of typi- cal species of Eutettix. Crown short, but usually distinctly longer medially than next eyes. Ocelli large, situated near eyes. Front narrow, strongly con- stricted at base of clypelius. Genae broad, lateral margins sinuated below eyes. Forewing with a distinct appendix; closed anteapical cells usually two in number, the outer sometimes divided. Type of the genus, Menosoma stonei Ball, 1931, known only from the southern part of the United States. The following members of the genus occur in South America: Eutettix cinctus Osborn and Ball, 1898; Mesamia jasdata Osborn, 1923; and Eutettix neocinctus Osborn, and E. elegans Osborn, 1923. Bahita, new genus (PI. XXIX, fig. 5; PI. XXXIII, fig. 8) Intermediate in character between Eutettix Van Duzee and Menosoma Ball; differing from the former in lacking the distinct striae on the crown and pronotum and in having reflexed vein- lets to the costal margin of the forewing; distinct from Menosoma by having a much shorter head and distinct transverse furrow on the crown, which is distinctly margined anteriorly. Crown short and of nearly uniform length, sometimes slightly longer medially than next the eye ; anterior margin carinate or sub- carinate. Crown with a distinct transverse furrow across entire width anterior to disk. Ocelli near eyes. Frontal sutures usually 380 The University Science Bulletin extending past ocelli onto vertex and terminating at transverse fur- row. Face convex; front broader than in Eutettix or Menosoma; genae broad, with lateral margins sinuated. Pronotum short, lat- eral margins not carinate. Forewing with a distinct appendix; closed anteapical cells two in nmnber, the outer short and lying obliquely with respect to costal margin; the central narrowed me- dially; costal area with two or three reflexed veinlets to costal margin near base of outer anteapical cell. Pygofer with a few stout setae. Type of the genus, Eutettix infuscatiis Osborn, 1923 (p. 52), from Brazil and Bolivia. The genus, which appears to be limited in dis- tribution to Central and South America, is well represented in that region. The following described species are referred here: Jassiis (Athysanus) palliditarsus Stal, 1860; Eutettix irroratus Osborn, E. femoratus Osborn, and E. laticeps Osborn, 1923; and Aligia plena Van Duzee, 1933, from Costa Rica. Bandar a Ball Bandara Ball, Bull. Brooklyn Ent. Soc, vol. 26, p. 93, 1931. Closely related to Eutettix Van Duzee, but without the distinct striae on the crown and pronotum which are characteristic of that genus. Differing also in having the outer anteapical cell of the fore- wing distinctly shorter than in Eutettix and broader distally than basally. Crown about as in Eutettix, but transverse furrow usually not distinct. Ocelli distant from the eyes about one and one half times their own diameter. Face short and broad as in Eutettix. Lateral margins of pronotum short and faintly carinate. Forewing with a distinct appendix; closed anteapical cells two in number, the second more narrowed medially than is true of Eutettix; outer apical cell unusually large and usually somewhat semicircular in shape. Setae of pygofer short. Color tawny or yellow, usually with opalescent spots on forewing. Type of the genus, Eutettix johnsoni Van Duzee, 1894 (p. 137), a Nearctic species. The only South American representative of the genus known to the writer is Mesamia hyalina Osborn, 1923, de- scribed from Colombia. Specimens at hand from Trinidad, Gre- nada, and the Dominican Republic are considered to be this species. Osborn's description of hyalina as having "Abdominal segments of Oman: Bythoscopinae and Jassinae 381 female above punctate at spiracles''' apparently refers to the dark depressions near the lateral margins of each segment on the dorsum, as all Cicadellidae examined by the writer have had the spiracles ventral in position. Atanus, new genus (PI. XXXIII, fig. 7) Closely related to Opsins Fieber, with which it agrees in general habitus, but differing in having the head of nearly the same width as the pronotum, the crown more nearly flat, the front more elon- gate and narrow, and the margins of the outer anteapical cell of the forewing nearly parallel. Crown rather flat, not rounded to the face anteriorly, but lacking a distinct margin, median length distinctly greater than the length next the eyes. Face triangular, front narrow, genae joining clypellus only as a very narrow sclerite. Lateral margins of pronotum not carinate. Forewing with a distinct appendix, closed anteapical cells two in number, the outer shorter and more slender than the inner and with sides nearly parallel. Pygofer with numerous stout setae. Type of the genus, Eutettix dentatus Osborn, 1923 (p. 57), de- scribed from Bolivia and also occurring in northern Argentina. The genus will also include Eutettix tesselatus Osborn, 1923; Thamnotet- tix lohatus Osborn, 1923; and twenty-five or more species closely related to dentatus. These are for the most part apparently unde- scribed, although several of the species described by Berg, 1881, 1884, as Athysanus probably belong here. Opsins Fieber Opsins Fieber, Verb. Zool.-Bot. Ges. Wien, vol. 16, p. 505, 1866. Comparatively small, rather wedge-shaped leaf hoppers with a short, broad head and two' anteapical cells in the forewing. Genae joining the clypellus as comparatively broad sclerites. Crown slightly longer medially than next the eye, rounded down- ward to the front. Ocelli large, distant from the eyes about two and one half times their own diameter. Face broad and short; lateral margins of genae strongly sinuated below eyes; clypellus with sides approximately parallel. Lateral margins of pronotum very short. Forewing opaque except apically and along costal margin ; a distinct appendix present ; outer anteapical cell narrowed at both ends. Setae on pygofer short and stout. 382 The University Science Bulletin Type of the genus, Opsins stactogalus Fieber, 1866, a species now cosmopolitan in distribution but thought to have come originally from the Mediterranean region. Exitianus Ball (PI. XXXIII, figs. 10, lOA) Exitianus Ball, Trans. Amer. Ent. Soc, vol. 55, p. 5, 1929. Mimodrylix Zachvatkin, Moscow Univ., Sci. Proc, No. 4, p. 108, 1935. (Type, Tham- notettix capicola Stal, 1855, an African species). (New synonymy.) Most closely related to Nephotettix Matsumura, with which it agrees in the shape of the clypellus and in having the appendix ex- tended around the apex of the forewing. Differing from Nephotettix in lacking the transverse furrow on the crown, in having the crown rounded to the face rather than margined anteriorly, and in having the outer anteapical cell comparatively large, normally truncate at both ends, and extending to the fourth apical cell, rather than very small, acuminate posteriorly, and not extending to the base of the fourth apical cell. Exitianus differs from Athysanus Burmeister and Euscelis Brulle, where the species have frequently been placed, by the shape of the clypellus and the appendix of the forewing. Crown longer medially than next the eyes, disk convex and an- terior margin rounded to the face. Ocelli large, distant from the eyes about their own diameter. Clypellus unusually long, narrowed gradually from base to apex. Lateral margins of genae sinuated be- low eyes. Forewing hyaline, appendix unusually large, closed an- teapical cells two in number. Pygofer and distal portion of oviposi- tor sheath of female with a few stout setae. Plates of male with a row of large setae along lateral margins, posterior margins of pygo- fer with two or three unusually stout setae. Ovipositor sheath ex- tending beyond pygofer. Type of the genus, Cicadula exitiosa Uhler, 1880 (p. 72), which is at present considered to be a synonym of Jassus [Thamnotettix) obscurinervis Stal, 1859 (p. 293), described from Brazil. Other American species belonging to the genus are the following: Athy- sanus picatus Gibson, 1919, and its synonym A. miniaturatus Gibson, 1919; Euscelis quxidratula Osborn, 1923; Euscelis pallida Osborn, 1926; and Athysanus digressus Van Duzee, 1933. Exitianus armus Ball, 1923, does not belong to the genus as here restricted, and may be referred to the already crowded Thamnotettix Zetterstedt until it can be disposed of more satisfactorily. African, European, and Asiatic species belonging to Exitianus are as follows: Thamnotettix Oman: Bythoscopinae and Jassinae 383 capicola Stal, 1855; Athysanus fusconervosus Motschiilsky, 1863; Jassus {Thamnotettix) taeniaticeps Kirschbaum, 1868, Athysanus indicus Distant, A. nanus Distant, and A. atkinsoni Distant, 1908; Athysanus transversalis Matsumura, 1908; Athysanus simillimus Matsumura, 1914; and probably several others described as Athy- sanus, Euscelis, or Phrynomorphus. Phrynornorphus Curtis, 1833, type nitidus Curtis, 1833, is a synonym of Euscelis Brulle. The spe- cific synonymy of all the species of Exitianus listed above is much involved and a study of much more material than is now available is needed to decide some of the points in question. It would be difficult to find a more homogeneous group of species than those listed above, and there can be no doubt about the affin- ities of the American, European, Asiatic, and African species. Hegira, new genus (PI. XXXIII, fig. 9) Related to Ophiola Edwards and Euscelis Brulle, but differing from both in having the face longer and the frontal sutures extend- ing beyond the ocelli onto the anterior margin of the crown. Dis- tinct from Ophiola in having the head narrower than the thorax and from Euscelis in having a larger appendix on the forewing. Crown short, but longer medially than next the eyes, rounded to the face. Ocelli near eyes. Front long, strongly constricted near base of clypellus. Genae broad, lateral margins sinuated. Lateral margins of pronotum faintly carinate. Forewing broad, appendix large, closed anteapical cells two in number, the outer shorter than the central one. Costal area with a vein to costal margin from each end of the outer anteapical cell. Pygofer with numerous large setae. Male plates with many long, slender, silky-appearing setae on outer margins. Type of the genus, Hegira brunnea, n. sp., from northern Argen- tina. A specimen of the same or a closely related species is at hand from Brazil. Hegira brunnea, new species Face brown with numerous pale-yellowish areas, the most con- stant being transverse marks on the clypeus. Crown yellowish- white with a pair of transverse brown dashes between the eyes, a pair of small spots at the apex, and a pair of larger spots on the posterior margin. Pronotum washed with brown, with numerous irregular pale areas on the anterior submargin. Scutellum brown 384 The University Science Bulletin with two yellowish-white spots on each lateral margin. Forewing subhyaline, marked with brown on veins and in cells, the intra- cellular marks restricted to the distal area. Length, 6-6.5 mm. Structure as indicated for the genus. Male valve broad and short, triangular in shape. Plates together bluntly spoon-shaped, basal width about equal to length. Last ventral segment of female large, median portion slightly longer than lateral portion, with a small median notch on posterior margin. Holotype male, allotype female, and 9 male and 7 female para- types from Loreto, Prov. Missiones, Argentina, collected in 1931 by A. A. Ogloblin, the holotype on December 14, the allotype on December 6, and the paratypes on various dates from November 27 to December 18. Types in the collection of the United States Na- tional Museum, Cat. No. 51672. Eusceloidia Osborn (PI. XXX, fig. 1 ; PI. XXXIV, figs. 1, lA) Eusceloidia Osbom, Ann. Cam. Mus., vol. 15, no. 1, p. 46, 1923. Superficially resembling macropterous specimens of the fulgorid genus Bruchomorpha Newman; apparently related to Exitianus Ball by the wing venation, but easily distinguished from that genus by the very large apical cells of the forewing and the greatly swollen clypeus. Crown short and sloping downward anteriorly, broadly rounded to the face. Ocelli distant from eyes about one and one half times their own diameter. Clypeus greatly swollen, especially next clypel- lus. Lateral margins of genae sinuated, genae very narrow next cly- pellus. Pronotum short, lateral margins very short and not cari- nate. Forewing rather broad, appendix and first and second apical cells unusually large. Closed anteapical cells two in number, the outer about half as long as the inner and lying along distal portion of that cell. The hind wings of the single specimen upon which the original description of the genus was based are badly torn, but the venation appears to be normal for the group, with four apical cells. Type of the genus, Eusceloidia nitida Osborn, a Bolivian species, and, so far as known, the only representative. Oman: Bythoscopinae and Jassinae 385 Faltala, new genus (PI. XXXIV, fig. 2) Broad, flat, and relatively short leaf hoppers, brachypterous so far as known. Resembling Athysanella Baker in general habitus, but probably not closely related to that genus. Crown flat and broad, separated from the face by a distinct mar- gin which is subcarinate at least apically. Median length of crown about twice as long as length next the eyes. Ocelli small, distant from the eyes two or three times their own diameter. Face broad and short, front unusually broad, genae with lateral margins incised below eyes, Pronotum shorter than crown, lateral margins com- paratively long and subcarinate. Macropterous forms, if they oc- cur, may be expected to have the lateral margins of the pronotum shorter. Forewing quadrangular in shape, truncate apically, not reaching to posterior margin of third abdominal segment, venation reticulated. Abdomen broad and much flattened, much broader than thorax. Pygofer with few or no setae. Type of the genus, Faltala brachyptera, n, sp., from northern Argentina. No other species are included in the genus. Faltala brachyptera, new species General ground color pale yellowish-brown, with irregular ivory- colored markings on crown, pronotum, forewing, and abdomen, these markings, except those of forewing, margined with fuscous. Mark- ings of crown illustrated in drawing (PI. XXXIV, fig. 2), those of abdomen consisting of two oblique dashes near each lateral margin on segments 3 to 7, inclusive, and 3 broader dashes on each segment between the lateral dashes, all these arranged so as to form longi- tudinal stripes. Eighth abdominal segment with a fuscous spot near each lateral margin dorsally. Median portion of posterior margin of female genital segment black. Length of male, 3 mm.; of female, 3.75 mm.; greatest width about 1.5 mm. Crown subangular apically. Posterior margin of pronotum broadly and shallowly incised. Scutellum broad and short, Male valve broad and short. Plates short and broad, extending only a short distance beyond apex of valve. Pygofer terminating in short points posteriorly. Last ventral segment of female with median portion of posterior margin slightly produced and tridentate. 25—6037 386 The University Science Bulletin Holotype male, allotype female, and 3 female paratypes from Loreto, Prov. Missiones, Argentina, collected in 1931 by A. A. Oglo- blin, the holotype on November 29, the allotype on November 25, 1 paratype on November 27, and the other 2 on November 29. Types in the collection of the United States National Museum, Cat. No. 51673. Brazosa, new genus (PI. XXXIV, fig. 3) Related to the Thamnotettix group, which it resembles in general habitus, but differing in having the pronotum much larger and humped in profile, and in the shape of the crown. Crown short and broad, median portion sometimes slightly pro- duced. Crown not separated from face by a distinct margin. Face broad, clypellus extended beyond genae, lateral margins of genae sinuated. Pronotum large, lateral margins carinate. Forewing hyaline, closed anteapical cells two in number. Type of the genus, Thamnotettix picturellus Baker, 1923 (p. 532), {Thamnotettix pictus Osborn 1923, from Brazil, not Thamnotettix pictus Lethierry, 1875). The genus will also contain Thamnotettix amazonensis Osborn, 1923, from Brazil. Alaca, new genus (PI. XXXIV, fig. 4) Related to the Thamnotettix group, with which it agrees in general habitus and wing venation, but differing in having the head broader than the pronotum and the crown short and of uniform length. Crown short and rounded downward to the face. Face convex, clypellus strongly constricted. Pronotum short, lateral margins weakly carinate. Forewing comparatively slender, closed anteapical cells two in number, appendix distinct. Type of the genus, Alaca longicauda, n. sp., from northern Argen- tina. The genus will also include Agallia multipunctata Osborn, 1923, and an apparently undescribed species from Bolivia. Alaca longicauda, new species General color sordid brown with a round black spot above each ocellus, a smaller fuscous spot on the pronotum behind each eye, and fuscous marks on the base of the scutellum. Length of male, 5.5-6 mm. Forewing subhy aline. Male valve very small, plates rather delicate and thin, strongly diverging distally and triangular in outline. Pygofer very long, about three times as long as plates, rounded apically in profile. Oman: Bythoscopinae and Jassinae 387 Holotype male and 28 male paratypes from Loreto, Pro v. Mis- siones, Argentina, collected in 1931 by A. A. Ogloblin, the holotype on December 18 and the paratypes on various dates from November 25 to December 18. Types in the collection of the United States National Museum, Cat. No. 52091. Onura, new genus (PI. XXXIV, fig. 7) Related to Thamnotettix Zetterstedt in general structure and agreeing with that genus in the venation of the corium of the fore- wing. Distinct from Thamnotettix and related genera in that the face is strongly receding and short, the apex of the clypellus does not extend caudad of the posterior margin of the eyes, and the forewing has only one distinct claval vein. Disk of crown shallowly concave, margin of crown broadly rounded to the face; face in profile most swollen between the anten- nae, constricted at the base of the clypellus. Ocelli large and situ- ated near eyes. Clypellus short and nearly parallel sided; lateral margins of genae sinuated. Pronotum short, lateral margins not carinate. Forewing long, appendix distinct, costal margin coriaceous on basal two thirds, anteapical cells lying parallel to costal margin, the outer shorter and more slender than the central one. There is occasionally a trace of the second claval vein at the inner basal angle of the clavus. Claval vein usually with a cross vein to claval suture. Pygofer sparsely clothed with rather short setae. Type of the genus, Onura eburneola, n. sp., from northern Argen- tina. Two other species, one from northern Argentina and the other from Brazil, will fall in this genus. Onura eburneola, new species Crown, anterior half of pronotum, and costal margin of forewing ivory colored. Ocelli orange. Upper portion of face, defiexed mar- gins of pronotum, and sides of abdomen also ivory, below dirty yel- low to fuscous. Crown and pronotum with faint, irregular yellow markings; posterior half of pronotum and all of scutellum dirty white. Forewing grayish-subhyaline, appearing slate-colored owing to the darker wings and dorsum of abdomen, with a fuscous stripe along radius parallel to costal margin and extending to apex of outer anteapical cell. Length of female, 4.75 mm. Crown produced and bluntly angled, median length equal to length of pronotum and nearly twice the length of the crown next the eyes. 388 The University Science Bulletin Last ventral segment of female subtruncate posteriorly, with a narrow, median, V-shaped notch which extends nearly to the base of the segment. Holotype female and 1 female paratype from Loreto, Prov. Mis- siones, Argentina, collected in 1931 by A. A. Ogloblin, the holotype on December 18 and the paratype on December 14. Type in the col- lection of the United States National Museum, Cat. No. 51674. Thamnotettix Zetterstedt Thamnotettix Zetterstedt, Ins. Lapp., column 292, 1838. Cicadula Zetterstedt, Ins. Lapp., column 296, 1838 (type. Cicada quadrinotata Fabricius, 1794, a palearctic species). Thamnus Fieber, Verb. Zool.-Bot. Ges. Wien, vol. 16, p. 505, 1866 (type, Thamnotettix confinis Zetterstedt, 1838, a palearctic species). Relatively elongate, slender species, without a distinct margin between the crown and the face of the head and with only two an- teapical cells in the forewing. Crown usually short, median length slightly longer than that next eyes. Head narrower or as wide as pronotum. Face relatively fiat, lateral margins of genae sinuated. Ocelli situated near eyes. Lat- eral margin of pronotum short, not or only feebly carinate. Fore- wing relatively broad, second cross vein absent, appendix distinct. Type of the genus, Cicada prasina Fallen, 1826, a Palearctic species. The genus as here defined continues to be the catch-all that it has been in the past. While many of the South American species described under this genus have been transferred to other genera, the number remaining is still too large to list here. As here defined the genus is cosmopolitan in distribution. The genus Cicadula, as fixed by its type quadrinotata, is here con- sidered as a synonym of Thamnotettix, while for the species of the genus Cicadula of authors the name Macrosteles Fieber (type, Cicada sexnotata Fallen, 1806) is available. Macrosteles apparently does not occur in South America. Acinopterus Van Duzee (PL XXX, fig. 2; PI. XXXIV, fig. 5) Acinopterus Van Duzee, Psyche, vol. 6, p. 308, 1892. Related to Thamnotettix Zetterstedt by the structure of the short, bluntly angled crown, but differing from that genus in that the fore- wing lacks a distinct appendix and the inner margin is continued in a nearly straight line from the apex of the clavus to the bluntly pointed apex of the forewing. Oman: Bythoscopinae and Jassinae 389 Head narrower than pronotum; crown short and rounded to the front, longer medially than next the eyes. Ocelli situated near eyes. Face relatively flat; lateral margins of genae sinuated below eyes. Pronotum short and broad, lateral margins carinate and diverging posteriorly, pronotum broadest near posterior margin. Forewing with two, or occasionally three, closed anteapical cells, often with extra cross veins, particularly in clavus and along costal area op- posite outer anteapical cell. Second apical cell of hind wing un- usually narrow. Type of the genus, Acinopterus acuminatus Van Duzee, 1892b (p. 308), an American species. So far as known, this is the only species of the genus which occurs in South America, and it appears to be limited to the extreme northern portion of the continent. Speci- mens from Colombia have been examined by the writer. Chlorotettix Van Duzee (PI. XXVII, figs. 1, lA, IB, IC, ID; PI. XXXIV, fig. 8) Chlanrotettix Van Duzee, Psyche, vol. 6, p. 306, 1892. Closely related to Thamnotettix Zetterstedt by the venation and general form; differing from that genus in having the crown more rounded and of nearly uniform length throughout its width, and in that the forewings are never opaque. Crown short and rounded to the front, median length equal to or slightly greater than that next to the eyes. Ocelli situated near eyes. Front comparatively slender; lateral margins of genae not strongly sinuated below eyes. Lateral margins of pronotum short and weakly carinate. Forewing long, without extra cross veins; closed anteapical cells two in number; appendix distinct. Pygofer with numerous stout setae. Type of the genus, Bythoscopus unicolor Fitch, 1851 (p. 58), a Nearctic species. The genus appears to be restricted in distribution to the Americas and is represented in South America by Chloro- tettix breviceps Baker and Ch. minimus Baker, 1898; Ch. bakeri Sanders and De Long, 1922 ; Ch. neotropicus Jensen-Haarup, 1922 ; Ch. aberrans Osborn, Ch. delicatus Osbom, Ch. dilutus Osborn, and Ch. truncatus Osborn, 1923; and Thamnotettix luteosus Baker. In addition to the above, there are at hand representatives of nineteen apparently undescribed species belonging to the genus. 390 The University Science Bulletin Benala, new genus (PI. XXXIV, fig. 10) Superficially resembling Deltocephalus Burmeister in size and gen- eral habitus, but different from that genus in having only two anteapical cells in the forewing and in that the ocelli are distant from the eyes. Crown over twice as long medially as next the eyes, not distinctly separated from the face, but with the apex pointed. Ocelli over one third distance from eyes to apex of head. Face strongly reced- ing, lateral margin of genae strongly sinuate. Pronotum shorter than head. Forewing with appendix distinct and inner apical cells unusually large. Type of the genus, Deltocephalus tumidus Osborn, 1923, from Bolivia. No other species are referred to the genus. Cortona, new genus (PI. XXX, fig. 6; PI. XXXIV, fig. 6) Related to Lonatura Osborn and Ball, which it resembles in size and general habitus. Differing from that genus in having a longer, more angled head and no closed anteapical cells in the forewing. Head large in proportion to body, distinctly wider than prono- tum. Crown produced and angled, rounded to face, length greater: than length of pronotum. Ocelli situated near eyes. Clypellus un- usually broad, tapering apically. Lateral margins of genae sinuated below eyes. Pronotum short, lateral margins very short. Forewing hyaline, appendix distinct, closed anteapical cells absent. Pygofer with numerous large setae. Type of the genus, Cortona minuta, n. sp., from northern Argen- tina. This is the only species referred to the genus. Cortona minuta, new species Color, uniform golden yellow, wings hyaline. Length of male, 2.3 mm.; length of head, pronotum, and scutellum, 0.9 mm.; width of head, 0.75 mm. Crown longer than pronotum, disk rather flat, median length twice that next the eyes. Forewing extending beyond tip of abdomen. Male valve broad and short. Plates together subtriangular, taper- ing to pointed tips. Holotype male from Loreto, Prov. Missiones, Argentina, Decem- ber 3, 1931, A. A. Ogloblin. Type in the collection of the United States National Museum, Cat. No. 51675. Oman: Bythoscopinae axd Jassinae 391 Stirellus Osborn and Ball / (PI. XXXIV, figs. 9, 9A) ;, : Athysanus subgenus Stirellus Osborn and Ball, Ohio Nat., vol. 2, p. 250, 1902. Related to Aconura Lethierry,* with which it agrees in the shape of the head, in having a long clypellus, a small appendix and two closed anteapical cells in the forewing, and a long ovipositor sheath. Differing from Aconura in being shorter and more robust, with a shorter and less pointed head and broader, shorter anteapical cells in the forewing. Anemochrea Kirkaldy, Anemolus Kirkaldy, Phryn- ophyes Kirkaldy, Nandidrug Distant, and Paivanana Distant are all related to Stirellus and Aconura. Small leaf hoppers. Crown produced and bluntly angled, rounded to the front, disk somewhat flattened. Head often subconical. Clypellus nearly parallel sided, extending beyond normal curve of genae, apex truncate. Lateral margins of genae sinuated. Prono- tum shorter than crown, lateral margins very short and not carinate. Forewing comparatively broad, not reaching to tip of ovipositor sheath, appendix small. Ovipositor sheath extending well beyond apex of pygofer; pygofer of female usually with a few short, stout setae. Type of the genus, Athysanus bicolor Van Duzee, 1892a (p. 114), an American species. In addition to bicolor the genus is known to be represented in South America by Athysanus (Stirellus) mexi- canus Osborn and Ball, 1902, and two other species from northern Argentina. Penestirellu^ Beamer and Tuthill Penestirellus Beamer and Tuthill, Jour. Kans. Ent. Soc., vol. 7, pp. 21-22, 1934. Related to Stirellus Osborn and Ball by the subconical head and long clypellus, but differing in having three closed anteapical cells in the forewing instead of two and in lacking an appendix. Differ- ing from Deltocephalus Burmeister and related genera by the shape of the crown and facial sclerites. Crown and face together subconical, disk of crown relatively flat, margin rounded to the face. Ocelli small and situated next the eyes. Front long, clypellus with sides nearly parallel and apex truncate or excavated and extended well beyond normal curvature of mar- gins of genae. Lateral margins of genae strongly sinuated below * The writer has not examined Acc/nura jakolevi Lethiern.-, 1876, the type of the genus. The generic concept is based on a study of several Japanese species. A) 392 The University Science Bulletin eyes. Pronotum and scutellum unusually short. Hind wing slender, with anal area much reduced in size. Ovipositor sheath extended beyond pygofer. Type of the genus, Penestirellus catalinus Beamer and Tuthill, 1934, known only from southern Arizona. The genus is represented in South America by an apparently undescribed species from north- ern Argentina. Baldulus Oman Baldulus Oman, Proc. Ent. Soc. Wash., vol 36, p. 79, 1934. Related to Macrosteles Fieber {Cicadula of authors), with which it agrees in wing venation, but differing from that genus in being elongate and slender with a produced and angled crown. Resem- bling Balclutha Kirkaldy and Nesosteles Kirkaldy in the shape and venation of the forewing, but with four apical cells in the hind wing. Crown produced and bluntly angled, rounded to the face without a distinct margin anteriorly. Ocelli small, distant from the eyes about one and one half times their own diameter. Face elongate, lateral margins of genae distinctly sinuated. Lateral margins of pronotum faintly carinate. Forewing with a distinct appendix, ante- apical venation, as here interpreted, with outer cell absent and inner cell open basally, resulting in only one closed anteapical cell. The present interpretation of the venation of the hind wing results in four apical cells instead of three. Type of the genus, Baldulus montanus Oman, 1934, known to occur only in southern Arizona. In addition to the two species originally referred to the genus, it will also include Cicadula maidis De Long and Wolcott, 1923, the only representative of the genus that is known to occur in South America. The writer has examined specimens of maidis from Puerto Rico, the type locality; Cuba; Coahuila, Mexico; Temple, California; and Tucuman, Argentina. The Argentine specimens were recorded ''on sugar beet" and the California material was reported to be damaging young sweet corn, Agelina, new genus Related to Macrosteles Fieber {Cicadula of authors), with which it agrees in the venation of forewing and in general habitus, but differing from that genus in having only three apical cells in the hind wing. In the character of the venation of the hind wings the genus shows affinities to Balclutha Kirkaldy and Nesosteles Kir- kaldy, but differs from these genera in several important structural details. Oman: Bythoscopinae and Jassinae 393 Crown produced and rounded, distinctly longer medially than next the eye. Ocelli distant from the eyes about one and one half times their own diameter. Face relatively broad. Lateral margins of pronotum short and not carinate. Forewing relatively broad and with a distinct appendix and one closed anteapical cell. Hind wing with three apical cells. Type of the genus, Agelina punctata, n. sp., from Brazil and Argentina. No other species is referred to the genus. Agelina punctata, new species General ground color sordid yellowish-green; forewings smoky. Head marked with fuscous to black as follows: Face with short, broken, transverse arcs and a curved transverse band between and below the ocelli; crown with a transverse bar behind ocelli, this broken on median line and with lateral extremities curved downward between ocelli and eyes, and a round spot on each side of median line behind transverse bar. Base of pronotum often with irregular dark marks. Veins of forewings pale. Coloration extremely vari- able, transverse bar on crown sometimes obsolete and other markings indistinct. Length of female, 3.25-3.75 mm. Structural details as indicated for the genus. Posterior margin of last ventral segment of female subtruncate, very slightly sinuated and with a faint median notch. Holotype female and 2 female paratypes from Loreto, Prov. Missiones, Argentina, December 18, 1931, collected by A. A. Oglo- blin. Type in the collection of the United States National Museum, Cat. No. 51676. Balclutha Kirkaldy (PI. XXX, figs. 4, 4A; PI. XXXV, fig. 1) Gnathodus Fieber (nee Pander, 1856), Verb. Zool.-Bot. Ges. Wien, vol. 16, p. 505, 1866 (type, Cicada punctata Tbunberg, 1782). Balclutha Kirkaldy, Entomologist, vol. 33, p. 243, 1900 (new name for Gnathodus Fieber). Eugnathodus Baker, Invert. Pacifica, vol. 1, p. 1, 1903 (type, Gnathodus abdominalis Van Duzee, 1892, which is at present considered to be congeneric with -punctata Thunberg. Slender, relatively small leaf hoppers, with a short head which is narrower than the thorax. Forewing long and slender, with a large appendix and only one closed anteapical cell. Crown slightly longer medially than next eyes, median length about one third median length of pronotum. Lateral margins of pronotum short but diverging posteriorly, pronotum widest some dis- tance behind eyes. Wing with three apical cells, the reduction 394 The University Science Bulletin caused by fusion of inner fork of radius and outer fork of media in apical region. Basal segment of hind tarsus with a distinct groove- like excavation on inner ventral surface at base. Type of the genus, Cicada punctata Thunberg, 1782 (pi. 21), a Holarctic species. An apparently undescribed species from Bolivia is the only South American form referred to this genus. Nesosteles Kirkaldy (PL XXXV, fig. 2) Nesosteles Klirkalcly, Hawaiian Sugar Planter's Assoc. Expt. Sta. Div. Ent. Bull. 1, p. 343, 1906. Anomiana Distant, Fauna British India, Rhynchota, vol. 8, p. 109, 1918 (type, Anomiana longula Distant, 1918). (New Synonymy.) Agellus De Long and Davidson, Ohio Jour. Sci., vol. 33, p. 210, 1933 (type, [Eugnathodus] Agellus neglecta De Long and Davidson, 1933). Closely related to Balciutha Kirkaldy, differing in having the head broader, the crown shorter and usually not distinctly produced medially, the pronotum not distinctly wider than the head and with its lateral margins not diverging posteriorly, and the forewing usually not so long. Other characters as in Balciutha. Type of the genus, Nesosteles hebe Kirkaldy, 1906, described from the Viti Isles. The genus, and many of the species included therein, is practically cosmopolitan in distribution, at least in the warmer regions of the world. This group is very closely related structurally to Balciutha, but is here retained as a distinct genus on the basis of the doubtfully distinct structural characters mentioned above and the apparent difference in distribution, Balciutha occurring primarily in Holarctic or Alpine regions while Nesosteles is the common form in warmer regions. The following species of the genus are recorded from South America: Eugnathodus lineatus Osborn, 1924, E. fiaves- cens Baker, 1903, and E. lacteus Baker, 1903; the last two originally described from Central America and recorded from South America by Osborn in 1924. However, E. lacteus Baker is a synonym of Gnathodus incisus Matsumura, 1902, which will also have as syno- nyms Eugnathodus bisinuatus De Long, 1923, Eugnathodus pallidus Osborn, 1926, Eugnathodus bifurcatus De Long and Davidson, 1933, Nesosteles areolata Obsorn, 1934, and Nesosteles tutuilana Osborn, 1934. (New synonymy.) The last three names are included on the authority of Davidson and De Long, 1935 (p. 106-107), who sup- pressed them as synonyms of bisinuataus. Balciutha hyalina Osborn, 1926, which was also listed by Davidson and De Long as a synonym of bisinuatus, is here removed from synonymy. Nesosteles incisa Oman: Bythoscopinae and Jassinae 395 Matsumura was described from Japan and is known to occur in the Philippines and Hawaiian Islands in addition to the localities rep- resented in the above synonymy. Identification of A^. incisa is from Japanese specimens determined by Matsumura. A'', guajanae De Long, 1923, is also known to occur in South America and the Philip- pine Islands. Idiotettix Osborn (PI. XXX, fig, 7; PI. XXXV, figs. 3, 3A) Idiotettix Osborn, Ann. Ent. Soc. Amer., vol. 22, No. 3, p. 465, 1929. Closely related to Exolidia Osborn by the shape of the head, the presence of ledges over the antennal sockets, and the shape of the appendix of the forewing, but differing from that genus in having three closed anteapical cells and a basal cross vein in the forewing. Crown short and broad and of nearly uniform length, sloping downward anteriorly and rounded to the face, median suture at base absent or very short. Ocelli distant from eyes about twice their own diameter. Pronotum broadest just behind eyes, posterior margin roundly excavated medially. Scutellum large. Forewing large, ap- pendix extended around the apex. Outer anteapical cell somewhat triangular in shape, base angled, central and inner anteapical cells truncate basally. Type of the genus, Thamnotettix magnijicus Osborn, 1924 (p. 424), from Bolivia. The genus also includes Idiotettix bolivianus Osborn, 7. fasciatus Osborn, and 7. brunnen^ Osborn, 1929, and a few apparently undescribed species as well. Exolidia Osborn (PI. XXX, fig. 5; PI. XXXV, fig. 4) Exolidia Osborn, Ann. Cam. Mus., vol. 15, No. 1, p. 75, 1923. General form robust, ledges above antennal pits distinct; fore- wing without a cross vein between R + M and Cu, and with apex obliquely subtruncate. Head short and broad; crown broad, sloping downward to the much swollen clypellus, median basal suture absent. Ocelli distant from the eyes about twice their own diameter. Pronotum broadest next posterior margins of eyes, posterior margin concavely excavated. Forewing broad apically, appendix unusually broad opposite apex of inner apical cell, cubitus not forked, R + M forked but once, outer apical veins curved, first apical cell very small. Apical cells three, or possibly four, in number. 396 The University Science Bulletin Type of the genus, Exolidia picta Osborn, a Brazilian species. No other species known to the writer at present can be referred to the genus as above defined. Neobala, new genus (PI. XXX, fig. 3) Related to Exolidia Osborn by the short, rounded crown, but separated from that genus by the venation of the forewing, which has a basal cross vein between R + M and Cu, and an anteapical cell which is truncate basally. Crown of nearly uniform length, sloping downward anteriorly and rounded to face, disk with numerous fine, longitudinal striae, basal median suture absent or very short. Ocelli distant from eyes about twice their own diameter. Face convex, clypeus somewhat swollen, genae rather broad next clypellus. Pronotum as in Exo- lidia. Forewing slender and tapered apically, apex rounded, ap- pendix large, apical veins not strongly curved as in Exolidia. Type of the genus, Thamnotettix pallidus Osborn, 1923 (p. 67), from South America. The genus, which appears to be limited in distribution to tropical South America, will also include Euscelis quadrimaculata Osborn, 1923, and Thamnotettix guaporensis (new name for Thamnotettix sordidus Osborn, 1923, nee Thamnotettix sordidus Zetterstedt, 1838). Conala, new genus (PI. XXXV, fig. 5) Most closely related to Neobala, n. g., with which it agrees in wing venation; related to Idiotettix Osborn and Exolidia Osborn as well as to Neobala by the shape of the pronotum, which is broadest just behind the eyes. The genus differs from the above groups in the much produced and nearly conical head and the fine transverse pronotal striae, and bears a superficial resemblance to Spangber- giella Signoret because of the red dorsal markings of the head and thorax, but has no close structural affinities with that genus. Crown greatly produced, apex narrowly rounded, disk slightly convex and with numerous fine, longitudinal striae, margins rounded to the face, and median basal suture absent. Ocelli small, distant from eyes about four times their own diameter. Face convex, ledge above antennal socket sometimes small. Pronotum with numerous fine, transverse striae, posterior margin angularly incised between Oman: Bythoscopinae and Jassinae 397 basal angles of scutellum. Forewing with a large appendix, one closed anteapical cell, the base of which is truncate, and two claval veins. Type of the genus, Spangbergiella fasciata Osborn, 1923 (p. 29), from Brazil. This is the only species known at present which be- longs to the genus. Coelidiana, new genus (PI. XXX, fig. 8; PI. XXXV, fig. 6) Related to, and previously included in Neocoelidia Gillette and Baker, but differing from that genus, as determined by its genotype tumidijrons Gillette and Baker, 1895, in being much more elongate with wings extending well beyond the tip of the abdomen, and in having a carina separating the crown and face. Differing from Chinaia Brunner and Metcalf by the presence of the carina between the crown and face and in having only three apical cells in the hind wing. Crown fiat or nearly so, usually somewhat pentagonal in shape, distinctly raised above the eyes and separated from them by a short perpendicular wall, median length always greater than that next the eyes. Face strongly convex; clypellus comparatively slender. Pro- notum wider than head, rather short; posterior margin broadly and shallowly incised; lateral margins comparatively long, distinctly carinate, somewhat curved and diverging posteriorly. Scutellum large. Forewing long and slender, with one closed, elongate, tri- angular anteapical cell. Appendix of forewing either very small or entirely absent. Type of the genus, Neocoelidia rubrolineata Baker, 1898 (p. 290), from Brazil. Other South American species included in this genus are Neocoelidia bimaculata Baker and A^. modesta Baker, 1898, and N. infiata Osborn and N. croceata Osborn, 1923. Neocoelidia smithii Baker, 1898, and probably N. punctata Osborn, 1923, and iV. ornata Osborn, 1924, should be referred to the genus Chinaia Bruner and Metcalf. Chinaia Bruner and Metcalf (PI. XXXV, figs. 7, 7A) Chinaia Bruner and Metcalf, Bull. Brooklyn Ent. Soc., vol. 29, No. 3, pp. 120-121, 1934. Most closely related to Neocoelidia Gillette and Baker and Jassus Fabricius by the broad genae, narrow head, long antennae, broad, short pronotum, and the reduced venation of the forewing. 398 The University Science Bulletin Differing from Neocoelidia in having four apical cells in the hind wing and from Jassus by the venation of the forewing and the nar- row costal area of the hind wing. Crown raised above eyes and separated from them by a short, perpendicular area, rounded downward anteriorly to the front. Crown extending but slightly anterior to the eyes. Ocelli small, dis- tant from the eyes about one third the distance from eyes to median line of vertex. Clypellus strongly constricted near base. Antennae extremely long, usually longer than entire body, basal segments large. Pronotum short, lateral margins diverging posteriorly and feebly carinate, posterior margin broadly and shallowly incised. Forewing long and slender, basal two thirds opaque, appendix small or absent, one closed anteapical cell present. Type of the genus, Chinaia bella Bruner and Metcalf, 1934. The genus is represented in South America by Neocoelidia smithii Baker, 1898. A^. punctata Osborn, 1923, and N. ornata Osborn, 1924, prob- ably belong here also. Chinaia was originally described as a member of the Bythosco- pidae because of the position of the ocelli. However, as indicated earlier in this paper, that character, used alone, is unreliable as a criterion for separation of the higher groups. The structure of the head and thorax, as pointed out above, identify Chinaia as a relative of Jassus and related genera. The nymphal stages also bear a close resemblance to the very distinctive nymphs of Jassus. Jassus Fabricius (PI. XXX, figs. 9, 9A; PI. XXXV, figs. 8, 8A) Jassus Fabricius, Syst. Rhyng., p. 85, 1803. Coelidia Germar, Mag. d. Ent., vol. 4, pp. 38, 75, 1821. (Type, Coelidia venosa Germar, 1821, from Brazil.) Comparatively large, robust leaf hoppers with simple venation of forewing, narrow crown, and long face. Head narrower than thorax, crown unusually narrow, usually about as wide as one eye. Crown comparatively short, disk raised above level of eyes and separated from them by a short perpen- dicular area. Ocelli large, situated on crown margin about as near to the median line as to the eye. Crown usually rounded to the front, occasionally separated by a distinct margin. Face unusually long, clypeus of about equal width throughout, clypellus usually broadened distally, apex extending beyond genae. Genae reaching clypellus as broad sclerites, lateral margins sinuated. Pronotum Oman: Bythoscopinae and Jassinae 399 short, lateral margins diverging posteriorly and distinctly carinate; carinae about even with costal margins of forewings. Scutellum large. Forewing broad, usually broader apically than medially or basally, appendix distinct, with only one closed anteapical cell which is petiolate basally and truncate distalty. Apical cells usually al- most parallel margined. Type of the genus, Jassus nervosus Fabricius, 1803, an American species. J. nervosus is not known to the writer but the generic de- scription here presented is based on numerous American species be- lieved to be congeneric with nervosus. Coelidia Germar appears to pertain to the longer headed American forms and may be found to be worthy of subgeneric or generic rank when the species are more fully known. A large number of species from South America have been de- scribed in this genus, and since most of them appear to have been correctly placed generically they are not listed below. Terulia Stal Terulia Stal, Bidrag till Rio Janeiro -Traktens Hemipter-fauna, Pt. II. K. Svensk. Vet. Akad. Handl., vol. 3, p. 50, 1860. Similar to Jassus Fabricius but larger and with the anterior and middle tibiae flattened but not broadened. Otherwise as in Jassus. Type of the genus, Terulia jerruginea Stal, 1860, a Brazilian species. The writer has not seen examples of ferruginea, and the above description is based on a specimen which appears to belong to the genus. Gabrita Walker Gabrita Walker, List. Homop., Suppl., p. 254, 1858. Petalopoda Spangberg, Ofv. Vet. Akad. Forh., vol. 36, p. 18, 1879. (Type, Petalopoda annulipes Spangberg, 1879, from South America.) Closely related to Jassus Fabricius in all important structural characters except that the fore femora and tibiae are broad and foliaceous. Type of the genus, Gabrita annulivena Walker, 1858, from South America. Neither annulivena nor annulipes are known to the writer from authentically determined material, but the characterization is based on a single specimen which appears to be Petalopoda annulipes Spangberg. Petalopoda is placed as a synonym of Gabrita on the basis of the descriptions of the two. 400 The University Science Bulletin LITERATURE CITED Baker, C. F. 1898. Notes on Jassini, with some new species. Can, Ent., vol. 30, pp. 289-292. 1923. Comparison of neotropical and paleotropical insect Faunae. Philippine Jour. Sci., vol. 23, pp. 531-532. Ball, E. D. 1931. Some new North American genera and species in the group formerly called Platymetopius (Rhynchota, Homoptera). Can. Ent., vol. 63, pp. 216-222, 224-228. 1932. The food plants of the leaf hoppers formerly included in the genus Platymetopius Burm. Can. Ent., vol. 64, pp. 251-255. Berg, C. 1879. Hemiptera Argentina, enumeravit speciesque novas descripsit. 316 pp. (Reprinted from An. Soc. Cient. Argentina, vols. 7-8, 1878-79.) 1881. Sinonimia y descripcion de algunos hemipteros de Chile, del Brasil y de Bolivia. An. Soc. Cient. Argentina, vol. 12, pp. 259-272. 1884. Addenda et emendanda ad Hemiptera Argentina. 213 pp. (Re- printed from An. Soc. Cient. Argentina, vols. 16-17, 1883- '84.) 1895. Hemipteros de la Tierra del Fuego. Coleccionados por el Senor Carlos Backhausen. An. Mus. Nac. Hist. Nat. Buenos Aires, vol. 4, pp. 195- 206. Davidson, R. H., and DeLong, D. M. 1935. A Review of the North American species of Balclutha and Agellus (Homoptera: Cicadellidae) . Proc. Ent. Soc. Wash., vol. 37, pp. 97-112. DoziER, H. L. 1926. Some new and interesting Porto Rican leaf hoppers. Jour. Dept. Agr. P. R., vol. 10, pp. 259-265. Fabricius, J. C. 1803. Systema Rhyngotorum. 314 pp. Fallen, C. F. 1826. Hemiptera Sueciae, Cicadariae. London. Fitch, A. 1851. Catalogue with references and descriptions of the insects col- lected and arranged for the state cabinet of natural history. (From Fourth Annual Report of the Regents of the University of the State of New York en the State Cabinet of Natural History.) Germar, E. F. 1817-1847. Fauna Insectorum Europae. Halae. Fasc. 3-24. Gmelin, J. F. 1788. In Linnaeus' Systema Naturae, 13th ed., vol. 1, pt. 4. Herrich-Schaeffer, H. S. 1838. Faunae Insectorum Germaniae initia. Regens- burg. Heft 154. (Fortsetzung von Panzer, Faunae Insect. Germ.) Jensen-Haarrup, a. C. 1922. Hemipterological notes and descriptions II. Ent. Medd., vol. 14, pp. 11-16. KiRKALDY, G. W. 1906. Leaf hoppers and their natural enemies. (Pt. IX. Leaf hoppers — Hemiptera.) Hawaiian Sugar Planters' Assoc. Expt. Sta., Div. Ent. BuJ. 1, pp. 271-479, pis. 21-32. 1907. Leaf Hoppers — Supplement (Hemiptera.) Hawaiian Sugar Plant- ers' Assoc. Expt. Sta., Div. Ent. Bui. 3, 186 pp., 20 pis. Lawson, p. B. 1932. The genus Spangbergiella (Homoptera, Cicadellidae) in America north of Mexico. Jour. Kans. Ent. Soc, vol. 5, No. 4, pp. 116-122. Linnaeus, C. 1761. Fauna Suecica. Holmiae. Ed. 2. McAtee, W. L. 1934. Genera and subgenera of Eupteryginae (Homoptera; Jassidae). Proc. Zool. Soc. London, pp. 93-171, 5 pis. Oman: Bythoscopinae and Jassinae 401 OsBORN, H. 1923. Neotropical Homoptera of the Carnegie Museum, parts 1 and 2. Ann. Cam. Mus., vol. 15, pp. 8-79. 1924. Neotropical Homoptera of the Carnegie Museum, parts 3 and 4. Ann. Cam. Mus., vol. 15, pp. 383-462. Provancher, L. 1872. Description de plusieurs Hemipteres noveaux. Nat. Canad., vol. 4, pp. 73-79, 103-108, 319-320, 350-352, 376-379. 1889. Petite faune entomologique du Canada, vol. 3. Quebec. Say, T. 1830-'31. Descriptions of new North American hemipterous insects, belonging to the first family of the section Homoptera of Latreille. Jour. Acad. Nat. Sci. Phila., vol. 6, pp. 235-244, 299-314. SiGNORET, V. 1879-'80. Essai sur les Jassides Stal, Fieb., et plus particulier- ement sur les Acocephalides Puton. Ann. Ent. Soc. France, ser. 5, vol. 9, pp. 259-290, vol. 10, pp. 41-70. Snodgr-ass, R. E. 1935. Principles of insect morphology. 667 pp., illus. Spangberg, J. 1879. Homoptera nonnula americana nova vel minus cognita descripsit. Ofv. Kong. Vet.-Akad. Forh., vol. 36, No. 6, pp. 17-26, 2 pis. Stal, C. 1859. Hemiptera. K. Svenska Fregatten Eugenies resa omkring Jorden under befiil af C. A. Virgin aren 1851-1853, vol. 2, Zoologi, pt. 4, Insekter, pp. 219-298. 1860. Bidrag till Rio Janeiro-Traktens Hemipter-faima. Part 2. K. Svenska Vetensk. Akad. Handl., vol. 3, No. 6, 75 pp. Thunberg, C. p. 1782. Acta Upsala, vol. 4. Uhler, p. R. 1877. Report upon the insects collected by P. R. Uhler during the explorations of 1875, including monographs of the families Cydnidae and Saldae, and the Hemiptera collected by A. S. Packard, Jr., M. D. Bui. U. S. Geol. Geog. Surv. Terr., vol. 3, pp. 355-475, 765-801. 1880. Remarks on a new form of Jassid. Am. Ent., vol. 3, pp. 72-73. 1895. An enumeration of the Hemiptera-Homoptera of the Island of St. Vincent, W. I. Proc. Zool. Soc. London, 1895, pp. 55-84. Van Duzee, E. P. 1890. New California Homoptera. Ent. Amer., vol. 6, pp. 35-38, 49-52, 77-80, 91-96. 1891. New North American Homoptera — III. Can. Ent., vol. 23, pp. 169-172. 1892a. New North American Homoptera — IV. Can. Ent., vol. 24, pp. 113-117. 1892b. The North American Jassidae allied to Thamnotettix. Psyche, vol. 6, pp. 305-310. 1892c. A synoptical arrangement of the genera of the North American Jassidae, with descriptions of some new species. Trans. Amer. Ent. Soc, vol. 19, pp. 295-307. 1894. New North American Homoptera, No. VIII. Can. Ent., vol. 26, pp. 136-139. 1907. Notes on Jamaican Hemiptera: A report on a collection of Hemiptera made on the island of Jamaica in the spring of 1906. Bui. Buffalo Soc. Nat. Sci., vol. 8, No. 5, 79 pp. Walker, F. 1858. List of the specimens of homopterous insects in the col- lection of the British Museum. Supplement. 369 pp. 26—6037 402 The University Science Bulletin PLATE XXVII FiQ. 1, forewing (X 7); lA, hind wing (X 7); IB, face (X 15); IC, head and thorax (X 15); and ID, left intermediate tibia and tarsus (X 30), of Chlorotettix unicolor (Fitch). Fig. 2, forewing; and 2 A, hind wing, of Bythoscopus lanio (Linnaeus) (X 7). Fig. 3, forewing of Pachyopsis laetus Uhler (X 7). Fig. 4, hindwing of Oncopsis distinctus (Van Duzee) (X 15). Oman: Bythoscopinae and Jassinae 403 PLATE XXVII H r> CM CU. COSTAL UAPCIM P -RADIUS M -MCOIA CU -CUBITUS CS-CLA^AL SUTURE CV-CLA^AL yClNS CL -COMMISSURAL LIH£ C -COSTAL AR£A BA.BASAL CELL DC-DISCAL CELLS aH-BflACHIAL CELL OA-OUTER ANTEARICAL CA-CEHTRAL ANTEARICAL I A. INNER ANTEARICAL XI -FIRST CROSS-VEIN Xl-SECOND CROSS-VEIN POSITION AI-riRST APICAL CELL Ai-SECOND APICAL CELL Al-TNIRD APICAL CELL AA-FOURTH APICAL CELL AP-APPEMOIX CM-COSTAL MARem R -RADIUS M -MEDIA CI -CUBITUS I Cl-CuaiTUS i PC-POSTCUBITUS IV-f'iRST VANNAL !V- SECOND VANNAL SV -UJBMARCINAL VEIN M -HAMULUS A -APICAL CELLS CP-CLVPEUS CL-CLVPELLUS O OCELLOCULAR AREA E -COMPOUND EVES e -CENAE L -LORI c-CRoru O-OCELLI P-PRONOTUH SSCUTSLLUIt ID 3-BYTHOSCOPUS J-PACHrOPSIS 4- ON CO PS IS 404 The University Science Bulletin PLATE XXVIII Fig. 1, hindwing of Brasa rugicollis (Dozier) (X 15)- Fig. 2, forewing of Bythonia rugosa (Osborn) (X 7). Fig. 3, hindwing of Nionia palmeri (Van Duzee) (X 15). Fig. 4, forewing, and 4A, hindwing, of Xestocephalus pulicarius Van Duzee (X 30). Fig. 5, forewing of Arrugada rugosa (Osborn) (X 8). Fig. 6, same of Egenus acuminatus Oman (X 7). Fig. 7, same of Spanghergiella vulnerata (Uhler) (X 15). Fig. 8, same of Bolarga boliviana (Osborn) (X 15). Fig. 9, same of Luheria constricta Osborn (X 15). Oman: Bythoscopinae and Jassinae 405 PLATE XXVIII 4-XESTOCEPHALUS 3-NIONIA 6-EGENUS 7 -SPA NG BERG I ELLA B-BOLARGA 9-LUHERIA 406 The University Science Bulletin PLATE XXIX Fig. 1, forewing of Scaphytopius elegans (Van Duzee) (X 15). Fig. 2, same of Sanctanus san-cttis (Say) (X 15). Fig. 3, same of Scaphoidida dngulata Osbom (X 15). Fig. 4, same of Bonamus lineatus Oman (X 15). Fig. 5, same of Bahita injuscata (Osborn) (X 8). Fig. 6, same of Kanorba reflexa Oman (X 15). Fio. 7, same of Garapita garbosa Oman (X 15). Fig. 8, same of Caphodus maculatiis Oman (X 15). Fig. 9, same of Agudus typicus Oman (X 15). Fig. 10, same of Hecaloidia nervosa Osbom (X 15). Oman: Bythoscopinae and Jassinae 407 PLATE XXIX hSCAPHYTOPIUS 2-SANCTANUS 4-BONAMUS 5-BAHITA 6-KANORBA 7-GARAPITA 9-AGUDUS lO-HECALOIDIA 408 The University Science Bulletin PLATE XXX Fig. 1, forewing of Eusceloidia nitida Osborn (X 15). Fig. 2, same of Acinopterus acuminatxis Van Duzee (X 15). Fig. 3, same of Neobala pallida (Osborn) (X 15). Fig. 4, forewing; and 4A, hindwing, of Balclutha punctata (Thunberg) (X 15). Fig. 5, forewing of Exolidia picla Osborn (X 15). Fig. 6, same of Cortona minuta Oman (X 30). Fig. 7, same of Idiotettix fasciatus Osborn (X 8). Fig. 8, same of Coelidiana rubrolineata (Baker) (X 8). Fig. 9, forewing; and 9A, hindwing, of Jassus olitorius (Say) (X 7). Oman: Bythoscopinae and Jassinae 409 PLATE XXX l-EUSCELOIDIA 2-ACINOPTERUS 4-BALCLUTHA 3-NEOBALA 7-IDlOTETTIX 8-COELIDIANA 9-JASSUS 410 The University Science Bulletin PLATE XXXI Fig. 1, head and thorax of Bythonia rugosa (Osbom) (X 15). Fig. 2, same of Nionia palmeri (Van Duzee) (X 30). Fig. 3, same of Xestocephalns pulicarius Van Duzee (X 30). Fig. 4, same of Cariancha cariboba Oman (X 15). Fig. 5, same of Brincardorus laticeps Oman (X 15). Fig. 6, same of Cerrillus notatus (Osborn) (X 35). Fig. 7, head and thorax; and 7A, face, of Arrugada rugosa (Osborn) (X 15)- Fig. 8, head and thorax of Luheria conslricta Osborn (X 15). Fig. 9, same of Bonamus lineatus Oman (X 15). Fig. 10, same of Egenus acuminatus Oman (X 15). Oman: Bythoscopinae and Jassinae 411 PLATE XXXI l-BYTHONIA 3-XESTOCEPHALUS 6-CERRILLUS 4-CARIANCHA 7-ARRUGADA 2-NlONIA S-BRINCADORUS 7A e-LUHERiA 9-BONAMUS lO-EGENUS 412 The University Science Bulletin PLATE XXXII Fig. 1, head and thorax of male; and lA, of female, of Spangbergiella vulnerata (Uhler) (X 15). Fig. 2, head and thorax of Bolarga boliviana (Osborn) (X 30). Fig. 3, head and thorax ; 3A, front ; and 3B, head in profile, of Scaphytopius elegans (Van Duzee) (X 30). Fig. 4, head and thorax of Scaphoidula cingulata Osborn (X 15). Fig. 5, same of Sanctanus sanctus (Say) (X 30). Fig. 6, same of Osbornellus auronitens (Provancher) (X 15). Fig. 7, same of Garapita garbosa Oman (X 30). Fig. 8, same of Portanus stigmosus (Uhler) (X 15). Fig. 9, same of Agudus typicus Oman (X 15). Oman: Bythoscopinae and Jassinae 413 PLATE XXXII l-SPANGBERCIELLA 2-BOLARGA 4'SCAPHOIDULA lA 5-SANCTANUS 7-GARAPITA a-SCAPHYTOPIUS 6-OSBORNELLUS 9-AGUDUS 8-PORTANUS 414 The University Science Bulletin PLATE XXXIII Fig. 1, head and thorax of Cumora angulata Oman (X 30). Fig. 2, same of Kanorha reflexa Oman (X 30). Fig. 3, same of Hecaloidia nervosa Osborn (X 15). Fig. 4, same of Caphodus maculatus Oman (X 30). Fig. 5, head and thorax; and 5A, face, of Baroma reticulata Oman (X 15). Fig. 6, head and thorax of Menosoma cincta (Osborn and Ball) (X 15). Fig. 7, same of Atanus de7itatus (Osborn) (X 30). Fig. 8, same of Bahita injuscata (Osborn) (X 15). Fig. 9, same of Hegira brunnea Oman (X 15). Fig. 10, head and thorax; and lOA, face of Exitianus obscurinervis (Stal) (X 30). Oman: Bythoscopinae and Jassinae 415 PLATE XXXIII 9-HEGIRA 10 A lO-EXITIANUS 416 The University Science Bulletin PLATE XXXIV Fig. 1, head and thorax; and lA, head in profile, of Eusceloidia nitida Osborn (X 30). Fig. 2, head and thorax of Faltala bmchyptera Oman (X 30). Fig. 3, same of Brazosa pichirella (Baker) (X 30). Fig. 4, same of Alaca longicauda Oman (X 15). Fig. 5, same of Acinopterus acuminatns Van Duzee (X 15). Fig. 6, same of Cortona minuta Oman (X 30). Fig. 7, same of Onura eburneola Oman (X 30). Fig. 8, same of Chlorotettix unicolor (Fitch) (X 15). Fig. 9, head and thorax; and 9A, face, of StireUus bicolor (Van Duzee) (X 30). Fig. 10, head and thorax of Benala tumida (Osborn) (X 30). Oman: Bythoscopinae and Jassinae PLATE XXXIV 417 l-EUSCELOIDIA 2-FALTALA 3-BRAZOSA 6-CORTONA 9-ST/RELLUS 27—6037 4-ALACA 7-ONURA S-ACIN0PTERU5 e-CHLOROTETTIX lO-BENALA 418 The University Science Bulletin PLATE XXXV Fig. 1, head and thorax of Balclutha punctata (Thunberg) (X 30). Fig. 2, same of Nesosteles hebe Kirkaldy (X 30). Fig. 3, head and thorax; and 3A, face, of Idiotettix fasciatus Osbom (X 15). Fig. 4, head and thorax of Exolidia picta Osbom (X 15) . Fig. 5, same of Conala fasciata (Osborn) (X 30). Fig. 6, same of Coelidiana rubrolineata (Baker) (X 15). Fig. 7, head and thorax; and 7A, face, of Chinaia bella Bruner and Metcalf (X 15). Fig. 8, head and thorax; and 8A, face, of Jassus olitorius (Say) (X 15). Oman: Bythoscopinae and Jassinae 419 PLATE XXXV l-BALCLUTHA 2-NESOSTELES 3-IDIOTETTIX 4-EXOLIDIA 6-COELIDIANA 5- CON ALA 7-CHINAIA 8-JASSUS BA 420 The University Science Bulletin INDEX Numbers refer to pages on which the genera are described. Synonyms and homonyms are italicized, valid names are not. Aceratagallia Kirkaldy 350 Acinopterus Van Duzee 388 Agallia Curtis 351 Agalliana Oman 350 AgaUiopsis Kirkaldy 351 AgaUiota Oman 351 Agelina Oman 392 Agdlus De Long and Davidson 394 Agudus Oman 371 AJaca Oman 386 Amplicephalus De Long 373 Arrugada Oman 362 Atanus Oman 381 Bahita Oman 379 Balclutha Kirkaldy 393 Baldulus Oman 392 Bandara BaU -: 3S0 Baroma Oman 377 Batracomorphus Lewis 353 Benala Oman 390 Bergallia Oman 350 Bergiella Baker 364 Bolarga Oman 366 Bonamus Oman 365 Brasa Oman 352 Brazosa Oman 386 Brincadorus Oman 361 Bythonia Oman 358 Bythoscopus Germar 353 Caphodus Oman 375 Cariancha Oman 360 Cerrillus Oman 362 Chinaia Bruner and Metcalf 397 Chlorotettix Van Duzee 389 Cieadula Zetterstedt 388 Cloanthus Ball 367 Codidia Germar 398 Coelidiana Oman 397 Conala Oman 396 Convelinus Ball 368 Cortona Oman 390 Cumora Oman 374 Deltocephalus Burmeister 373 Deltopinus Ball 367 Dixianus Ball 377 Egenus Oman 363 Eugnathodus Baker 393 Euragallia Oman 351 Eurinoscopus Kirkaldy 354 Eusceloidia Osborn 384 Exiti anus BaU 382 Exolidia Osborn 395 Faltala Oman 385 Gabrita Walker 399 Galboa Distant 352 Garapita Oman 369 Gargaropsis Fowler 353 Gnathodus Fieber 393 Haider us Oman 373 Hecaloidia Osborn 378 Hegira Oman 383 Idiotettix Osborn 395 Ionia Ball 350 I owanus Bali 377 Jassus Fabricius 398 Kanorba Oman 374 Luheria Osborn 363 Macropsis Lewis 352 Menosoma Ball 379 Nasutoideus Ball 368 Neobala Oman 396 Neopsis Oman 353 Nesosteles Kirkaldy 394 Nionia Ball 358 Oncopsis Burmeister 353 Onura Oman 387 Opsius Fieber 381 Osbornellus Ball 369 Pachyopsis Uhler 354 Faraphlepsius Baker 377 Pediopsis Burmeister 352 Pendarus Ball 377 Penestirellus Beamer and Tuthill 391 Petalopoda Spangberg 399 Phlepsius Fieber 376 Platymoideus Ball 368 Portanus Ball 370 Sanctanus Ball 371 Scaphoidula Osborn 368 Scaphytopius Ball 367 Spangbergiella Signoret 364 Stirellus Osborn and Ball 391 Stragania Stal 353 Straganiopsis Baker 354 Terulia Stal 399 Texananus Ball 377 Thamnotettix Zetterstedt 388 Thamnus Fieber 388 Xestocephalus Van Duzee 359 Zioninus Ball 377 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 17. A Contribution to the Taxonomy of the Subfamily Issinae in America North of Mexico (Fulgoridae, Homoptera) KATHLEEN C. DOERING, Department of Entomology, University of Kansas Part I Abstract: This paper comprises the first part of a monograph dealing with the taxonomy of the subfamily Issinae (Fulgoridae, Homoptera) in America, north of Mexico. This section of the study includes only the genus Dictyssa, which is one of the few large genera in the subfamily. Of the ten established species the following five species of Melichar's are redescribed and compara- tive notes and drawings of structural details are given: D. areolata, D. mutata, D. fusca, D. clalhrata and D. marginepunctata. Comparative notes and draw- ings are added for four other species; namely, D. ohliqim Ball, D. ovata Ball, D. fcjiestrata Ball, and D. trayisvcrsa Van Duzee. Material of Dictyssa mira Van Duzee was not available for study, but this species is included in the key. Dictyssa semivitrea Provancher is reduced to synonymy under Dictyobia permutata Uhler. The characters of most value for classification are the male genitalia, the shape of the tegmina and the venation of the wings. The follow- ing species are described as new: D. maculosa, D. monroviana, D. quadravitrea, D. beameri and D. balli. INTRODUCTION THE purposes in writing this paper are several: First, to assem- ble under one article all our knowledge to date concerning the taxonomy of the previously described species in this subfamily which occur in America, north of Mexico, since many species have been added to the group after the publication of Melichar's mono- graph in 1906; second, to make comprehensive keys to the genera and the species ; third, to describe a number of new species and rede- scribe some of the older doubtful species; fourth, to study and figure the male genitalia of all available species, which, as far as the writer knows, has not hitherto been done in the Fulgoridae; lastly, (421) 422 The University Science Bulletin to present visual aids, such as the drawings of lateral views of the different species, which may help others in identifying the species. Family Characteristics Antennae placed on the side of the head below the eyes, very vari- able in shape and size, but usually consisting of two segments termi- nated by a very fine hair or bristle, rarely with basal appendages (Otiocerus) ; sense organs on antennae numerous and generally of a complex nature and mostly situated on the second segment; middle coxae articulated considerably apart, allowing for considerable range of movement, as opposed to those of the Cicadoidea, which are closer together and allow for very limited movements ; hind coxae immobile and exterior part coalesced with metathorax; no filter present be- tween midintestine and crop; ovipositor, with the exception of part of the Cixiinae, greatly reduced or incomplete, anterior and middle pairs of valves not fastened together by tongue and groove arrange- ments as in the Cicadoidea. Prothorax normally developed and unarmed. Note. — The classification of the family Fulgoridae is today a controversial question. Some Homopterists believe that the family should remain as one family, which in turn is then subdivided into some ten or eleven subfamilies. Others believe that many of these subfamilies should be raised to family rank. Until we have completed more detailed morphological and taxonomic work of the entire group, especially in regard to wing venation and male genitalia, the writer does not feel that it is advisable to make too radical a departure from the generally accepted older view of keeping the family Fulgoridae intact. If this view is held, the family characteristics are the same as the superfamily Fulgoroidea. Subfamily Characteristics Head not narrower, or only slightly so, than pronotum. Clypeus convex, but not greatly produced beyond plane, its lateral margins not keeled. Pronotum posteriorly subquadrate, anteriorly roundingly produced. Scutellum usually much longer than pronotum. Tegmina in texture either coriaceous, somewhat corneus, or vitreous, either entirely or with vitreous patches; held rooflike over body, their anal margins usually meeting, but sometimes widely separated. Hind wings very variable, being either absent, rudimentary, slight and narrow, or large and ample where they are notched at sides so that they can be folded in three. Hind legs with two to four spines on hind tibiae. Doering: Subfamily Issinae 423 Technique and Terminology The only special technique which might be of interest to describe is that used in studying the male genitalia. Drawings of these struc- tures were made from microscope slides, made in the following way. The specimens were relaxed either in a relaxing jar or by boiling them for a minute or two in hot water. The tip of the abdomen was carefully removed and this placed in 10 percent caustic potash which was heated to the boiling point and sometimes boiled for a moment or two (the exact time required should be tried out by the operator) . After boiling in the caustic potash the tip of the abdomen was next placed in glycerine or water for dissection. The dissected parts were then allowed to stand in 95 percent alcohol for a few seconds. From this medium they were immediately mounted in diaphane on the slide. Drawings of the lateral views of the aedeagus, the harpagones and the anal flaps were made from these slides. All other drawings were drawn from the specimens. The terminology for most of the morphological characteristics was obtained from Snodgrass' textbook "Principles of Insect Morphol- ogy" (1936) or Muir's paper, entitled, "On the Classification of the Fulgoroidea" (1923). Special terms used in the descriptions or keys are labeled or indicated on the first drawings of each plate. THE GENUS DICTYSSA Description of the Genus Small, robust insects with hemispherical tegmina which are held vertical to the body and whose greatest width is at tip of clavus or posterior to tip. The apical margin of the tegmen strongly round- ing and the costal margin expanded. Vertex extremely short through middle, not produced much beyond eyes, usually depressed through middle and with elevated margins. Pronotum extremely narrow at sides where it tapers to almost a point; its anterior margin deeply emarginate into region of the vertex; its posterior margin shallowly concave. Mesonotum triangular, with or without a median carina present. Frons quadrangular, usually more or less parallel mar- gined, truncate posteriorly, deeply emarginate anteriorly for the in- sertion of the postclypeus. Postclypeus moderately inflated. Ante- clypeus and labrum small. Main veins of tegmen coarse, a network of finer veins between them; median cells or bands of cells fre- quently hyaline, which contrast sharply with the opaque or darker 424 The University Science Bulletin outer cells. Wing venation, based on Muir, Tillyard and Snodgrass, show the following general characteristics: vein Sc divided, Sc^ ( = costal vein of Metcalf) running along costal border for at least half the length of the wing; Scg and R united at base and then running more or less parallel with each other to apex where they are lost in the apical reticulation; vein R thus unbranched, as is typical for many Homoptera; vein M typically two branched, sometimes three branches present; vein Cu^ divided into two branches, Cu^a and Cu^b; vein Cug forming the claval suture as in all Homoptera. Hind wings usually absent or rudimentary. Two spines present on hind tibiae. History of the Genus To date the following eleven species have been described: D. semivitrea Provancher (1889), D. areolata Mel. (1906) , D. clathrata Mel. (1906), D. jusca Mel. (1906), D. marginepunctata (1906), D. mutata Mel. (1906), D. fenestrata Ball (1910), D. ovata Ball (1910), D. obliqua Ball (1910), D. transversa Van Duz. (1914), and D. mira Van Duz. (1928). In the opinion of Mr. E. D. Ball (University of Arizona, Tucson) and Mr. Paul Oman (Homopterist, U. S. National Museum) Dic- tyssa semivitrea Provancher is Dictyobia 'permutata Uhler (1889). The author agrees with this viewpoint, as Provancher's written de- scription fits fairly accurately the species known as Dictyobia per- mutata Uhler, and cannot possibly be apphed to any known species of Dictyssa. Melichar in his revision of the Issinae evidently mis- took Dictyonia obscura Uhler (1889) for Provancher's semivitrea. Of the ten known species in the genus the writer has studied types or paratypes of all species except mira Van Duz. To this list is being added in the present paper five new species. Key to the Species 1. Tegmen with no hyaline cells or oblique band across center of corium dis- tinctly separated from darker outer cells 2 Tegmen with two or more large hyaline cells or an oblique band of hyaline cells across middle of corium, distinctly separated from the other opaque cells 4 2. (1) Entire tegmen translucent with occasional brown spots, elongate; smallest species in the genus ; greatest width of tegmen in line with apex of clavus transversa Van Duzee, 440 Tegmen opaque, greatest width at least two thirds of length and posterior to apex of clavus 3 3. (2) Tegmen entirely fuscous, semiopaque; vein M1 + 2 peparatrd at oentpr of tegmen; tegmen broadest just posterior to apex of clavus. . ./usco Melichar, 428 Tegmen opaque black, with a marginal border of circular hyaline spots; angle in vein R nearer to M1 + 2; vein M1-I-2 simple, marginepwnctata Melichar, 431 Doering: Subfamily Issinae 425 i. (1) Tegmen with hyaline band across corium composed of numerous cells, at least more than five 5 Tegmen with three to five conspicuous hyaline cells across corium, in an oblique band 7 5. (4) Tegmen with width subequal to length, cells small; greatest width of teg- men in line or slightly anterior to apex of clavus ; vein Mi4-2 divided just beyond middle clathrata Melichar, 430 Tegmen elongate, their greatest width either in line or posterior to apex of clavus ; smaller insects 6 6. (5) Tegmen with large, angular cells, the hyaline band across corium broad in proportion to opaque part; vein M1+2 unbranched and running straight across corium; base of wing greatly narrowed quadravitrea, n. sp., 441 Tegmen dark blackish -brown ; hyaline band of cells composed of many small ones; vein Mi-|-2 branched beameri, n. sp., 443 7. (4) Larger insects, 4.5-5 mm. in length ; elytra with oblique bands of fuscous and tan in addition to the vitreous cells 8 Smaller insects, usually under 4.5 mm. ; bicolored tegmina with vitreous and fuscous cells strongly contrasting 9 8. (7) Tegmina thickened, greenish-white with oblique fuscous and pale vittae, Ttiira Van Duzee, 427 Tegmina not particularly thickened, pale, clouded with fuscous; distinct large hyaline cells across corium ; greatest width beyond apex of clavus ; angle of R strongly pronounced so that vein R at this point comes nearer vein Mi+2 than Sco areolata Melichar, 425 9. (7) Tegmina elongate, the large central hyaline cell of corium at least twice as long as its width ; angle in vein R acute, at which point R is nearer vein Mi+2 than Sco 10 Tegmina not elongate, the central hyaline cell subquadrate, its length ap- proximating its width; vein R sinuate, making a less distinct angle at end of the central hyaline cell 11 10. (9) Tegmina distinctly elongate, its length approximately twice its width, the hyaline cells of corium extremely long, running more lengthwise of wing than in other species balli, n. sp., 445 Tegmina not twice as long as wide ; the central hyaline cell large, extending posteriorly beyond middle to about base of apical fourth, two other conspicuous hyaline cells placed between it and costal border; angle in vein R very acute monroviana, n. sp., 448 11. (9) Similar to areolata in shape of tegmina; central hyaline cell of corium al- most circular, the oblique hyaline band abbreviated, not reaching costa; costal border broad ; angle in vein R acute and near center of tegmen. ovata Ball, 438 Species elongate; the oblique hyaline band with cells irregular in size, reach- ing completely to costal border; costal border narrow 12 12. (11) Brown in general color with two transverse bands of hyaline spots behind middle of tegmina 13 Brown with only one transverse band of hyaline spots, the conspicuous hyaline cells across apex lacking 14 13. (12) Head and pronotum dark; one transverse row of hyaline cells behind middle in corium of tegmen; hyaline cells across apex of corium large; angle of vein R nearer to vein M1 + 2 than vein Sc2 fenestrata Ball, 436 Head and pronotum bright yellow ; discs of many cells in corium hyaline, the apical hyaline cells not any larger or more conspicuous than others ; two transverse rows of hyaline cells beyond middle; angle in vein R equidistant between veins Mi +2 and Sc2 maculosa, n. sp., 450 14. (12) Dark tegmina with a yellow or light-colored head and thorax; angle in vein R at about base of apical third of tegmen ; vein R at this point equi- distant between veins Mi-|-2 and Sc2 mutata Melichar, 433 With a dark-colored head and thorax; angle in vein R located just slightly posterior to middle, vein R at this point nearer to vein M1+2 than Sc2, obliqua Ball. 439 Dictyssa areolata Melichar, 1906 Melichar, Leopold. Monographic der Issiden (Homoptera). Abh. k. k. Zool.-Bot. Ges. Wien, III, pt. 4, 1906. Size. Length of bociy to tip of tegmen, 3.6 mm. to 4.8 mm. Length of tegmen, 3.5 mm. to 3.9 mm. ; width of tegmen, 2.7 mm. to 2.8 mm. This is one of the largest species in the genus. 426 The University Science Bulletin Color. This is a variegated species in brownish-yellow and dark brown. Head and thorax from above uniformly golden tan or some- times washed in fuscous. Below thorax light yellow. Abdomen tan, the genitalia frequently brownish. Tegmina variegated in coloring, the oblique hyaline band made up principally of 5 to 6 very large cells with the two central cells subequal and larger than the others and those touching the costal border the smallest of all ; these hyaline cells bordered with opaque white and studded around entire margins by dark spurs of veins; anterior to the hyaline band the tegmen is golden brown shaded to very dark brown at base, the costal border at base whitish-yellow crossed by dark, elevated veinlets; posterior to hyaline band is a very dark brown area, occupying apical half of clavus, then extending on to corium where it ends as the posterior dark border of the large central cell; after this dark band occurs a much lighter band, starting just posterior to apex of clavus as a cluster of about 5 small white cells, then becoming a transverse band of light brown and ending at the more posterior hyaline discal cell ; finally apex of corium pitch brown, studded all around apical border with semicircular and elongate white, subhyaline spots and some- times in addition a cluster of three small round hyaline cells in the region of cell M^+o anterior to the border. Structural characteristics. Vertex extremely broad and short, de- pressed through middle, its lateral margins anteriorly converging, its length on median line approximately half length of pronotum at mid- dle. Greatest width of eyes less than half the width of vertex. Frons with lateral margins subparallel although bulging slightly next to eyes, all its margins sharply elevated, median carina very distinct but disappearing before reaching clypeus. Postclypeus with posterior margin angularly produced into the frons for a distance equal to about one fourth the length of the latter. Pronotum with anterior margin sharply elevated, roundingly pro- duced and its posterior margin very shallowly concave; its length through middle approximately one half the length of the mesonotum at middle; a median carina only faintly visible if at all; two small round depressed spots on disc, one on each side of median line. Mesonotum with only a very faint median carina present, a trans- verse groove following anterior margin but stopping at either side before reaching lateral margin. Tegmina one fourth longer than wide, their greatest width posterior to apex of clavus, hemispherical in outline, with oblique hyaline band across disc made up of 4 to 5 Doering: Subfamily Issinae 427 large cells, the largest one being the central one located between veins M and R, another one almost equaling it in size, posterior and ventrad of it between R and Sc, a large one in the anal region extend- ing to the clavus, and one or two small ones touching costal border; across apex of tegmen large elongate hyaline spots alternating with small semicircular ones; the costal border expanded at base with round or rectangular spots in the cells between the veinlets; vein R making almost right-angled bends, extending very close to M^^, so that the space between veins R and Sc^ is about 3 times the distance between R and M^+o! vein Mj^., single. Male genitalia. Anal flap (10th abdominal segment) in width about twice as wide as long, its posterior margin truncately emargi- nate. Eleventh segment scarcely visible, its dorsal stylus long and conspicuous. Harpagones (genital styli), visible externally as two-pointed plates exceeded by anal tube for a short distance. From a flattened lateral view (see drawing 25, plate XLI) each harpago is roughly quadrangular, broadest through apical third, the dorsal margin of this region extended dorsad into a short, recurved hook, another small broad hook or flap just anterior to the latter. The aedeagus as viewed from the right side is a long tube partially sclerotized, bearing a short recurved lateral hook attached slightly posterior to middle, and two large ventrad-curving basal hooks, the apical halves only of which extend beyond the theca. On the left side a short blunt hook is attached at middle. The theca in this species extends over about the basal third of the aedeagus; on the right side its posterior margin is truncate and on the left side posteriorly it is extended into a heavily sclerotized, strongly recurved hook which simulates the appearance of the thecal hooks. Female genitalia. Anal flap (10th abdominal segment) broad, roundingly pointed posteriorly. The external valves of the ovi- positor widest at a point about midway of their length, their com- bined width greater than their length. Comparative notes. This species is an easily recognized species externally by both color and structure. The variegated or mottled tan and brown of the tegmina and the cluster of small white spots in the region of cell M^^o in addition to the usual oblique and apical bands separate it very distinctly. It is the largest species in the genus, excepting D. mira Van Duzee. It differs from all other species in the genus by having vein R bend at almost an acute angle and coming so close to M^+o that the space between veins R and 428 The University Science Bulletin Scg is about three times the distance between R and Mj_^2- The male genitalia are characteristic and distinctive for the species. (See plate XXXVIII, drawings 4a and 4b.) Notes on distribution. This species was described from males col- lected at Los Angeles, Cal. The type is located in the museum at Washington. Since this species has been wrongly determined, drawings of the type are figured on plate XXXVI as well as that of a specimen recently collected. A large series of this species has been collected by R. H. Beamer from Monrovia, Cal., in July. Dictyssa fusca Melichar, 1906 Melichar, Leopold. Monographie der Issiden (Homoptera). Abh. k. k. Zool.-Bot. Ges. Wien, III, pt. 4, 1906. Size. Length of body to tip of tegmen, 3.3 mm. Length of tegmen, 2.85 mm. ; width of tegmen, 2.4 mm. Color. Most of body and tegmina uniform testaceous brown, oc- casionally with one or two small hyaline cells in corium of tegmen. Thorax light tan shaded with brown. Abdomen dark testaceous brown. Legs tannish to brown. Structural characteristics. Vertex characteristic for the genus, very broad and short, length through middle approximately one half of pronotum at middle. Greatest width of eyes slightly over one third the width of the vertex. Front, parallel-margined, its greatest width through middle; the median carina distinct. Pronotum ex- ceeded at sides by eyes. Mesonotum over one third longer than pronotum, a distinct transverse groove near anterior end and a faint median carina present but not reaching apex. Tegmina semiopaque, not much longer than wide, their greatest width just posterior to apex of clavus at which region they are distinctly inflated; the veins coarse, outlining small cells, the costal margin at base considerably reflexed; venation characteristic for the species with vein R not angled but merely rounding, distinctly nearer Sc than M, vein M forked very near base, vein M^^g separating into M^ and Mg slightly posterior to middle of tegmen, vein M3+4 simple, curving more toward Cuja than in most species except D. fusca. Male genitalia. Anal flap which is an external ventral extension of the tenth abdominal segment, not quite twice as long as broad, slightly bilobed at tip. Eleventh abdominal segment scarcely visible, its dorsal stylus long and slender. Harpagones (genital styli of authors) visible externally on ventral side as two broad, platelike lobes, but which appear to be dovetailed into abdomen more than in most species. From a flattened lateral DoEpiNG: Subfamily Issinae 429 view (see drawing 21, plate XLI) each harpago is subquadrangular and has its posterior dorsal corner prolonged dorsad into a slender process recurved at tip around aedeagus, a short broad spine or hook placed externally at the base of this dorsal extension. The aedeagus is distinctly different from clathrata and other species. Unfortu- nately the drawing is a composite one made from a poor slide, yet the differences are so distinct that it leaves little doubt as to this being a distinct species. The aedeagus as viewed from the right side is a long slender tube bearing one long recurved hook midway of its length and a two- hooked extension attached at its extreme base, these hooks showing beyond the theca as two equal, adjoining and heavily sclerotized hooks which curve only slightly dorsad. As viewed from the left side the tips of the three hooks only are visible. The theca is a semimembranous tube, covering about one half of the aedeagus at its base. As viewed from the right side it has a truncate, posterior margin and from the left side has its ventral region extending more caudad into a broad truncate flap. (See drawings 2a and 2b, plate XXXVIII.) Female genitalia. Anal flap (10th segment) as drawn attached to specimen about twice as long as broad with the genital stylus long and slender. The external valves of the ovipositor as viewed from the ventral side are broadest at a point from base which is about three-fourths of the total length of the valves, their combined widths at this point greater than the length of the valves. Comparative notes. This species is distinguished externally from all other species by the uniform dark-brown color, the small size of the cells of the tegmen, the characteristic broadness of the tegmen and the forking of vein M3+4. It more closely resembles D. clathrata than any other species, but it differs externally from this species by lacking the hyaline cells in the corium and by having vein M separated near base of wing and vein R distinctly nearer to Sco than M. The male genitalia easily distinguish this species from all other species and are especially significant in separating it from D. clathrata. Data on distribution. Melichar lists this from California. The data on paratype material assigns this species to Placer county, Cali- fornia, collected in October. Perhaps the lateness of the season ac- counts for the scarcity of specimens. It appears to be the rarest species in the genus. Type specimens are located in the U. S. National Museum at Washington, D. C. 430 The University Science Bulletin Dictyssa clathrata Melichar, 1906 Melichar, Leopold. Monographie der Issiden (Homoptera). Abh. k. k. Zool.-Bot. Ges. Wien, III, pt. 4, 1906. Size. Length of body to tip of tegmen, 3.3 mm. to 3.6 mm. Length of tegmen, 2.7 mm. to 3.15 mm.; width of tegmen, 2.25 mm. to 2.7 mm. Color. Body pitch brown except the ventral side of thorax, which is light brown. Margins and faint median line of vertex tan. Mar- gins of pronotum darker brown than disc. Margins and carinae of mesonotum tan. Median carina of frons and clypeus tan, rest of frons pitch brown, clypeus somewhat lighter. Tegmina typically pitch brown except for oblique band of hyaline or milky, semi- transparent cells across clavus and corium and 5 small, round, whitish, semitransparent cells along extreme apical margin. Legs pitch brown. Note. — Variation in color occurs in this species to the extent of the head and thorax being golden brown as opposed to pitch brown of the tegmina or sometimes the latter also become golden brown likewise. Structural characteristics. Vertex broad and short, length through middle one half the length of pronotum at middle. Great- est width of eyes approximately one half the width of the vertex. Frons with lateral margins parallel, its length along middle equal to the length of the postclypeus; the median carina distinct, running into a faint transverse carina which does not reach to lateral mar- gins. Mesonotum not quite twice as long as pronotum on median line; a distinct transverse groove along anterior border which does not reach to the lateral margins; a faint median carina present and two lateral carinae faintly indicated. Tegmina semiopaque, some- what inflated, approximately one ninth longer than wide, their great- est width in line with apex of clavus; the veins coarse, outlining small cells, the costal margin narrowly expanded on basal half; venation characteristic of the species with vein R not angled but merely rounding, distinctly nearer to M^ than Sc,, vein M forked only slightly anterior to middle, vein M^+a separated posterior to middle, vein M3+4 curving more toward Cu^a than in most species. Male genitalia. Anal flap (10th segment) not quite twice as broad as long, slightly notched at tip. The eleventh segment slightly visible, its dorsal stylus approximating it in length. Harpagones (genital styli) visible externally on ventral side as two slenderly pointed flaps, the rest of the abdomen showing con- siderably beyond them. From a flattened lateral view (see draw- ing 20, plate XLI) each harpago is roughly quadrangular but has Doering: Subfamily Issinae 431 its posterior, dorsal region extended into a slender process, truncate at tip and only slightly curved, the whole process being shorter than that of D. jusca. At the base of this dorsal extension is a slender, pointed ventrad curving external hook. The eadeagus as viewed from the right side is a curved tubular structure bearing a short slender pointed hook midway of its length and a broad, well sclerotized hook attached near its base but ex- tending well beyond the theca almost to the tip of the aedeagus. On the left side the aedeagus bears no hooks. The theca as viewed from the right side extends a little beyond the basal third of the aedeagus and is truncate anteriorly. From the left view it appears as an elongate broad flap, extending almost to apex of the aedeagus. Female genitalia. Anal flap (10th segment) as drawn on speci- men broader through basal half and tapering slightly to a truncate apex. The external valves of the ovipositor very broad in propor- tion to length, their greatest width being just slightly posterior to the middle. Comparative notes. This species is separated externally from the majority of species by its very broad and characteristically shaped tegmina, and the small cells in the wing and especially those making up the hyaline or light band. It clearly resembles D. jusca Melichar in size and shape of tegmina. It is separated externally from the latter by the presence of the oblique band of hj^aline cells and the apical row of five round white cells as well as by the fact that the greatest width of each tegmen is anterior to the apex of the clavus in clathrata but posterior in jusca, and that vein R is distinctly nearer to vein M than Sca and vein M is separated only slightly anterior to middle. The male genitalia distinctly separate this species from every other member of the genus. Notes on distribution. Melichar lists this species from California. Recently specimens have been taken by Mr. Paul Oman and R. H. Beamer from Redding, Cal., in June, and from Sacramento county, California, by L. D. Anderson in August. The author had available for study 8 specimens. Apparently this species, like ju^ca, is not a common one. Dictyssa marginepunctata Melichar, 1906 Melichar, Leopold. Monographie der Issiden (Homoptera). Abh. k. k. Zool.-Bot. Gea. Wien. Ill, pt. 4, 1906. Size. Length of body to tip of tegmen, 3.15 mm. to 3.5 mm. Length of tegmen, 2.7 mm. to 2.85 mm.; width of tegmen, 1.8 mm. to 2.1 mm. 432 The University Science Bulletin Color. This is the darkest species in the genus. Body color is dark brown except for light brown or yellow on margins and median stripe on vertex, median carinae and lateral and posterior margins on pro- notum and mesonotum, median carina of frons and two spots on gena, one dorsad of antenna and one ventrad. Tegmina uniform blackish-brown, the conspicuous veins lighter or even reddish and a conspicuous row of semitransparent white spots, numbering about twelve to fifteen, sometimes practically uniform in size, in other specimens alternately small and large, those in the costal margin rectangular and crowded together. Legs blackish-brown. Note. — One color variation is noted in some specimens, namely a bright yellow head or thorax contrasting sharply with the pitch-black tegmina. Structural characteristics. Vertex characteristically broad and short, slightly more than twice as broad as greatest width of eye, length through middle about half length of pronotum. Frons broad, subequal in length and width, with a distinct, well-elevated median carina. Pronotum with a faint median carina and laterad of it on either side a small circular depression. Mesonotum with a faint median carina and in center of each lateral third a shallow depres- sion. Tegmina coriaceous with distinct elevated veins, their greatest width on a line even with apex of clavus, their posterior margins evenly rounding, the costal margin somewhat expanded at base; venation characteristic of the species with the angle in vein R in- dicated but not pronounced and vein R at this point closer to vein Mi+2 than Scg, mainly due to the fact that the latter runs nearer middle of tegmen; vein M1+2 single. Male genitalia. Anal flap (10th segment) slightly broader than one half its length, with its apical margin shallowly emarginate. The eleventh segment not always visible, its dorsal stylus an elongate fingerlike lobe. Harpagones (genital styli) from an external ventral view broad at base and tapering at their apices, not much of abdomen visible be- yond them. From a flattened lateral view (see drawing 19, plate XLI) each harpago is roughly quadrangular with its posterior dorsal corner extended into a short, sharply pointed process, at base of which is a broad, shallow, small external hook. The aedeagus as viewed on the right side is a long, curved tubular structure bearing a sharply pointed sclerotized hook attached at a point from apex of aedeagus, which equals in distance about one fourth the total length of the aedeagus and shows the apices of two heavily sclerotized, sharply pointed basal hooks, one of which ex- Doeeing: Subfamily Issinae 433 tends beyond the theca a little more than twice as far as the other does. On the left side the aedeagus bears an apical hook which is attached at a point from apex of aedeagus that equals in distance about one third the total length of the aedeagus. The two basal hooks show again from this view. The theca on the right side ex- tends over the aedeagus not quite half its length, and is angularly produced on posterior margin. On the left side the theca is extended posteriorly into a finely tapering spinelike process. (See drawings 11a and lib, plate XXXIX.) Female genitalia. Anal flap (10th segment) broadest through middle, suddenly tapering to a roundingly pointed apex. The ex- ternal valves of the ovipositor very broad in proportion to length, their greatest width being just slightly posterior to the middle. Comparative notes. In size and shape of tegmina very similar to the obliqua and miitata group. It differs from these species by the uniform dark-brown tegmina with the sharply contrasting white apical and costal spots and the small cells throughout the length of the tegmina. It differs from other species by the characteristic shape of the tegmina. The male genitalia definitely distinguish it from other species. Notes on distribution. Melichar lists this species from California. It is described from a female specimen which is in the Signoret col- lection in Wien, Austria. The author studied nine specimens col- lected in San Diego county, California, by Paul Oman, and at Campo, Beaumont and Jacumba, Cal., by R. H. Beamer in July and August. Dictyssa mutata Melichar, 1906 •fe*- Melichar, Leopold. Monographie der Issiden (Honioptera). Abh. k. k. Zool.-Bot. Ges. Wien, III, pt. 4, 1906. Size. Lenth of body to tip of tegmen, 3 mm. to 3.3 mm. Length of tegmen, 2.4 mm. to 2.7 mm.; width of tegmen, 1.8 mm. Color. Head, pronotum and scutellum uniform pale-yellowish or brownish-yellow. Thorax tawny yellow. Abdomen somewhat darker yellow washed in bro^vTi, especially at lateral carinae and on the genitalia. Legs, tawny yellow. Tegmina with wing pattern characteristic of the mutata-obliqua group. In this species the vitreous, oblique band of cells starts at base of clavus, runs poste- riorly slightly beyond middle, then bends abruptly ventrad, ending in the costal margin. The central hyaline cell located between veins M and R is much larger than any other cell in wing, is distinctly 28—6037 434 The University Science Bulletin ovate in shape, its posterior boundary situated two thirds of the wing length from the base of the wing, its margins studded with 5 to 9 short, dark spurs of veins. The second largest hyaline spot lies dorsad and cephalad of the central cell, a group of small, roundish hyaline cells, varying in number from 2 to 5, or else one large cell, adjoin the central cell at its caudo-ventral angle, and beyond these on the costal border is a median-sized elongate pellucid cell. The entire apical and costal margins of the tegmen are studded by pel- lucid disclike or semicircular spots which are not uniform in size but which frequently alternate between large and small ones. Structural characteristics. Vertex broad and short, length through middle one half the length of the pronotum. Greatest width of eyes one half the width of vertex. Pronotum with anterior margin roundingly produced and its posterior margin shallowly concave; its length through middle one half the length of mesonotum at middle; the median carina not present or only faintly discernible; two small, depressed, round spots on disc, one on each side of median line. Mesonotum with no median carina distinctly visible; a shal- low depression in disc on each side of median line and a transverse groove at base of apical prolongation. Frons, subequal in length and width, parallel-margined, the median carina very distinct but fading away before reaching postclypeus. Postclypeus with pos- terior margin angularly produced into frons for a distance equal to about one third of the total length of the latter. Tegmina approxi- mately one fourth longer than wide, their greatest width posterior to apex of clavus showing a great contrast between the hyaline cells and the opaque brown cells, both in texture and size; the largest cell in the wing, located on disc of corium between veins R and M, is in length approximately about one third the total length of the teg- men, and its width is approximately two thirds of its length, although its size is subject to some variation; veins of the tegmen coarse, outlining cells of varying sizes; the costal margin moderately ex- panded; wing venation characteristic of the species with a distinct angle in vein R, located more posteriorly than in some species, at a point approximately at base of apical third of wing, the area be- tween R at this point and vein M^.a equal to space between R and SCo; vein M^+a single. Male genitalia. Anal flap (10th abdominal segment), its width approximately two thirds of its length, its posterior margin truncate. Eleventh segment scarcely visible, its dorsal stylus a short finger- like projection. Doering: Subfamily Issinae 435 Harpagones (genital styli) visible externally as two triangular plates, not much of abdomen visible beyond their apices. From a flattened lateral view (see drawing 22, plate XLI) each harpago is subquadrangular and has its posterior dorsal corner prolonged dorsad into a sharply pointed short process, at base of which is a recurved short broad external flap. The aedeagus as viewed from the right side is a long slender tube bearing a slender, sclerotized, sharply pointed projection attached at the base of its apical third and two basal hooks, the apical half only of which shows beyond the theca. These latter two hooks are of unequal size, the dorsal one very slender and sharply pointed, the ventral one broad at base and gradually tapering to a recurved slen- der tip. On the left side the aedeagus bears an apical projection also, but this one is attached nearer to the middle of the aedeagus. The theca envelops the aedeagus at its base not quite half its length. On the right side it is truncate posteriorly. On the left side it ex- tends posteriorly as a sharply pointed, slightly sclerotized spine, which very easily can be missed, since a process of the aedeagus usually covers it. Female genitalia. Anal flap as drawn while attached to specimen approximately one third longer than wide, tapering posteriorly to a roundingly pointed apex. The eleventh segment and dorsal stylus approximately very small. The external valves of ovipositor broad- est at a point about midway of their length, their combined width much greater than their length. Comparative notes. This species is quite easily separated from the majority of the species in the genus by the presence of the large central hyaline cell. Other species in the genus having a large central cell likewise are D. areolata, D. obliqua, D. fenestrata, D. ovata and D. halli. From D. areolata it is easily separated by the much larger size of body of areolata and by the fact that vein R in the latter is so far removed from Sc. From D. fenestrata it is like- wise easily separated by lacking the second transverse band of hya- line cells across apex and by having the angle of vein R not as close to Mi+2 as in that species. D. halli is easily distinguished from other species by its elongate form and very long central cell between veins R and M. In D. ovata the hyaline band does not reach the costal margin and the angle in vein R is very near the middle of the wing, both of which characters are distinctly different from mutata. From D. obliqua and D. monroviana, mutata is not so easily distinguished. In fact, these three species seem to be repre- •436 The University Science Bulletin sentatives of a rather unstable group in the genus, for specimens of various localities show any number of variations in the cells and veins of the tegmina. The writer believes that D. mutata and D. obliqua are both good species, since their genitalia are distinctly- different. Externally the only distinguishable characters are the color of the head and thorax, which is yellow in mutata and dark in obliqua, and the angle of vein R being at base of apical third of tegmen in mutata but nearer the middle in obliqua, while vein R is equidistant from vein Sco and vein M^^, in "the former and nearer Mi+2 in the latter. Any number of gradations from these typical types can be found. The writer took only the form differing most widely from either of these forms, both in genitalia and external characters, and described it as a new species, namely monroviana. If typical examples of all three species are compared monroviana shows a noticeable difference in the tegmina by having much larger hyaline cells on both the discal area and apical margin. It is sepa- rated, furthermore, from obliqua by having the angle in vein R located as in mutata at base of apical third. It is separated from mutata by having R nearer vein M^+o than vein Sc, as in obliqua. Notes on distribution. The species was described from a male taken in Los Angeles county, California, and one at Claremont, Cal., collected by Baker. The former type is in the U. S. National Mu- seum and the latter in Melichar's collection. This seems to be a common species. Specimens are at hand for study from San Ber- nardino, Cal., collected by Coquillet, and the following places in California, collected by R. H. Beamer in August: Big Bear Lake, San Jacinto Mountains, Beaumont and Orange county. Dictyssa fenestrata Ball, 1910 Ball, E. D. New Genera and Species of Issidae. Proc. Biol. Soc. Wash. XXIII, pp. 41-46, 1910. Comparative notes. D. fenestrata belongs in the mutata-obliqua group, which it resembles in shape of tegmina and size. Its measure- ments are as follows: length from tip of head to apex of tegmina, 3 mm. to 3.4 mm. ; length of tegmen, 2.7 mm. to 2.9 mm. ; width of tegmen, 1.8 mm. to 1.9 mm. This species is separated from other species by the following characteristics: the elongate tegmina, which are widest at a point some little distance back of apex of clavus, the unifonn brown color of tegmina with lighter veins, the extremely large hyaline cells and spots across apex and in the oblique band of clavus and corium, and the presence of a transverse band of hyaline cells, numbering 5 or 6, just back of middle of Doering: Subfamily Issinae 437 corium which no otlier species possesses; an acute angle is present in vein R just posterior to middle, at which point this vein is much closer to M^+o than Scg, vein M^^^ sometimes is crowded toward clavus so that it coalesces with or obliterates the tip of vein Cu^a, and vein Cu^ is usually two-branched. Male genitalia. Anal flap (10th abdominal segment) twice as long as wide, only slightly emarginate at apex. Eleventh segment small and usually invisible, its dorsal stylus large and conspicuous. Harpagones (genital styli) visible externally as two pointed tri- angular plates, the anal flap visible a short distance beyond their apices. From a flattened lateral view (see drawing 15, plate XLI) each harpago is subquadrangular in shape, narrowed at base, bulg- ing at middle, and with the posterior, dorsal angle prolonged dorsad as a short, bluntly pointed projection, at base of which is a short, recurved, external hook. The aedeagus is a slender, tubular structure, its apex curving to- ward its base in such a way as to almost outline a complete circle. On the right side it bears a flattened, very sharply tapering project tion attached at about base of apical third and which extends to apex of the aedeagus. At base the latter bears also two heavily sclerotized hooks, of which only the apices show beyond the theca, the dorsal one of these hooks curving slightly dorsad, the ventral one almost straight and extending beyond theca for only half the distance that the other one does. On the left side the aedeagus bears another flattened, tapering projection which is attached at base of apical third and extends to apex of aedeagus. The theca envelops the aedeagus for approximately one third its length. On the right side the posterior margin is truncate. On the left side its posterior dorsal angle extends posteriorly as a sharply pointed process closely adpressed to the side of the aedeagus. Female genitalia. Anal flap partially covered at base by ninth abdominal segment, roundingly pointed at apex. The eleventh segment inconspicuous and bearing a slender dorsal stylus. The external valve of ovipositor broadest through middle, the length ap- proximately one and one half times its width. Notes on distribution. Doctor Ball described this species from six specimens collected at Tia Juana, Cal. The author greatly appre- ciated the gift of one type for study and from which the drawings have been made, Mr. Paul Oman of the United States National Museum collected a series of twenty-two specimens from Del Mar, Cal, on June 2, 1935. 438 The University Science Bulletin Dictyssa ovata Ball, 1910 Ball, E. D. New Genera and Species of Issidae. Proc. Biol. Soc. Wash. XXIII, pp. 41-46, 1910. Comparative notes. This is one of the broader species in the genus, with the following measurements: from tip of head to apex of tegmina, 3.3 mm. to 3.6 mm.; length of tegmen, 2.85 mm. to 3.15 mm.; width of tegmen, 2.1 mm. to 2.5 mm. It is easily separated from other species by the following characteristics; the circular shape of the tegmen with the very broadly expanded costal area, the abbreviated oblique hyaline band which does not reach the costal margin, and the almost circular central hyaline cell of this band be- tween veins R and M, caused by vein M curving toward anal region and vein R toward the costal margin ; a distinct angle present in vein R very near middle of wing, at which point R is much nearer to vein Mi+2 than vein Scg; vein Sc2 breaking up into a network of veins at a point midway of the length of the central cell. Male genitalia. Anal flap (10th abdominal segment) very long, more than twice as long as wide, its posterior margin shallowly emarginate. Eleventh segment, short and inconspicuous, the dorsal stylus proportionally long and prominent. Harpagones (genital styli) visible externally as two-pointed tri- angular plates, the anal flap visible for considerable distance beyond their apices. From a flattened lateral view (see drawing 17, plate XLI) each harpago is subquadrangular in shape, much more broad- ened across apical region than in most species, the posterior dorsal angle drawn out into a recurved projection, ventrad of which is a short, recurved, external hook. The aedeagus is a much curved, tubular structure, bearing on the right side a sharply tapering flattened process which is broadly at- tached at base of apical third and two basal heavily sclerotized hooks, whose ventrad curving apices show considerably beyond the theca, the ventral one of the two about two thirds as long as the dorsal one. On the left side the aedeagus bears a sharply pointed flattened process attached slightly posterior to middle. The theca envelops the aedeagus for about half its length. On the right side its posterior margin is truncate. On the left side its posterior dorsal angle extends caudad as a sharply pointed projection to a point at about base of apical third of the aedeagus. Female genitalia. Anal flap broad at base, roundingly pointed at apex. Eleventh segment inconspicuous, bearing a slender dorsal Doering: Subfamily Issinae 439 stylus. External valves of ovipositor longer than in most species, their combined width not much greater than their length, their great- est width at base of apical third. Notes on distribution. Doctor Ball described this species from eight examples from Tia Juana, Mexico, and Tia Juana, Cal. He very kindly gave the author a type male and female for study. This seems to be a very rare species. Dictyssa obliqua Ball, 1910 Ball, E. D. New Genera and Species of Issidae. Proc. Biol. Soc. Wash., pp. 41-46, 1910. Comparative notes. This species is a trifle larger than D. mutata, measuring 3.3 mm. to 3.6 mm. from apex of head to tip of tegmina, and each tegmen 2.7 mm. to 3 mm. long and 1.8 mm. to 2.1 mm. wide. It is separated from the various species in the genus by the following characteristics; a dark-brown head and thorax, margined in yellow; its tegmina about one fourth longer than their width, their greatest width at a point even with apex of clavus or just slightly posterior, the hyaline oblique band of cells across corium and the apical row of semicircular or elongate ones arranged as, and similar in size to, those in D. mutata; the venation of these two species, however, dif- fering in that D. obliqua has the angle of R more anterior than D. mutata, located just slightly posterior to middle and vein R is usually nearer to vein M^^, than Sc^, while in D. mutata it is about equidistant from veins R and Sc,; vein M.^^^ single. For further comparative notes see the discussion under this heading in the de- scription of D. mutata. Male genitalia. The genitalia are distinctive for this species. Anal flap (10th abdominal segment) elongate, not quite twice as long as width, its posterior margin shallowly emarginate. Eleventh segment inconspicuous, its dorsal stylus small. Harpagones (genital styli) visible externally as two pointed tri- angular plates, the anal flap visible for some distance as viewed from the ventral side, beyond their apices. From a flattened lateral view (see drawing 26, plate XLI) each harpago is subquadrangular in shape, its basal half somewhat narrowed, the posterior dorsal angle prolonged dorsad into a short, sharply pointed projection, at base of which is a short, broad, recurved external hook. The aedeagus as viewed from the right side is a long, much curved tubular structure, bearing a sharply pointed flat projection attached at approximately the base of its apical fourth and two basal hooks. 440 The University Science Bulletin whose apices show beyond the theca, the ventral one shorter and curved slightly dorsad, the other one extending beyond the thecal margin twice as long as the other one and curved ventrad at tip. On the left side the aedeagus bears a pointed flat projection attached at about base of apical third. The theca envelops the aedeagus at base for not ciuite half of its length. On the right side it is truncate posteriorly. On the left side its posterior dorsal angle extends caudad to a point a little beyond middle of aedeagus, but does not form the long slender hook as in D. mutata or D. monroviana. Female genitalia. Anal flap approximately one third longer than wide, tapering posteriorly to a roundingly pointed apex. The eleventh segment small, showing very little, bearing a slender dorsal stylus. External valves of ovipositor broadest at a point midway of their length, their combined width greater than their length. Notes on distribution. The type specimens are in Dr. E. D. Ball's collection, Tucson, Ariz. He collected twelve specimens from Tia Juana, Cal., and Tia Juana, Mexico. The writer had a long series of specimens to study collected from the following places in Cali- fornia: Alpine, Campo, San Diego county and Claremont county, by R. H. Beamer, in July and August. Dictyssa transversa Van Duzee, 1914 Van Duzee, E. P. A Preliminary List of the Hemiptera of San Diego county, California. Trans. San Diego Soc. of Nat. Sci., 2, p. 41, 1914. Comparative notes. This is the smallest species in the genus, with the following measurements: length from apex of head to tip of tegmina, 2.5 mm. to 2.75 mm.; length of tegmen, 2 mm. to 2.25 mm.; width of tegmen, 1.1 mm. to 1.25 mm. It is easily distinguished from other members of the genus by the whitish hyaline tegmina, reticulated with heavy brown nerves and crossed by two brownish fuscous bands, one located just anterior to apex of clavus and the other at base of apical fourth of the corium; the wing venation is characteristic; vein Sc is simple; vein R is not angled but broadly sinuate, extending nearer to vein M^+a than Sq^', veins M and Cu^ both branched about the same distance from base at a point just anterior to middle; all veins rather straight and running more or less parallel. Male genitalia. Anal flap (10th abdominal segment) long, slender with lateral margins parallel and posterior margin rather broadly concave, its length a little over twice its width. The eleventh seg- ment inconspicuous and its dorsal stylus short and broad. Doering: Subfamily Issinae 441 Harpagones (genital styli) visible externally as triangular plates, which are twice as broad at base as at their tips; the anal flap scarcely visible beyond their apices; each harpago is quadrangular with dorsal and ventral margins parallel, except for the posterior dorsal angle which is prolonged dorsad into a slender process which is slightly recurved at tip and at base of which is located a short, broad external hook. The aedeagus is a curved tubular structure. On the right side it bears a small, sinuately curved, well-sclerotized spine attached at middle, and two sclerotized hooks or processes attached near base and which extend beyond theca for only part of their length. The dorsal one of these is almost straight and extends beyond theca only about one third as far as the other; the ventral one, curving to a sharp ventrad curving point, extends almost to middle of the aedea- gus. On the left side the aedeagus bears a short projection attached at apex of its basal third. The theca on the right side envelopes the aedeagus for only about one fourth the length of the latter and is truncate posteriorly. On the left side its dorsal angle extends cau- dad as a sharply pointed, partially sclerotized projection for a dis- tance equaling about two thirds the length of the aedeagus. Female genitalia. Anal flap short, and in length about one fourth greater than wide. The eleventh segment is barely visible and possesses a long dorsal stylus. The external valves of the ovi- positor have their greatest width at a point just posterior to the middle ; the length of each valve is approximately one third greater than the width. Notes on distribution. Mr. Van Duzee in the original description gives the following notations: "Described from numerous specimens mostly taken on the slopes of Mt. Soledad at La Jolla, from Sept. to Nov. Also taken at Alpine in Mar. and Oct. and at Torrey Pines in June. Like most of its congeners it lives on Artemisia." Dr. R. H. Beamer collected a small series of these at San Diego, Cal., in August, 1935. Dictyssa quadravitrea, n. sp. ORIGINAL DESCRIPTION Size. Length of body from apex of head to tip of tegmen, 2.7 mm. Length of tegmen, 2.1 mm. ; width of tegmen, 1.5 mm. Color. General color of body yellowish-brown ; tegmina also yel- lowish-brown with large hyaline cells through middle. Vertex, pro- notum and mesonotum from above uniformlv vellowish-brown. 442 The University Science Bulletin Eyes dark brown. Disc of frons yeilowish-brown ; margins dark, outlined in brown. Postclypeus and genae likewise uniformly yel- low-brown. Clavola of antennae dark brown. Thorax from under side slightly lighter than rest of body. Legs tan or yellow-brown as well as most of abdomen except for median margins of ovipositor valves, which are dark. Tegmina yellowish-brown except for the following variations: a dark spot at base; an oblique hyaline band, which starts on clavus at its middle, runs posteriorly to a point just beyond middle of corium, then bends ventrad and extends as a transverse band to the costal border; numerous ovate whitish hya- line spots in the apical and expanded costal borders. Structural characteristics. Vertex somewhat produced cephalad beyond eyes, its anterior margin quite straight, its lateral margins slightly converging anteriorly, the length at this point equal to length of pronotum at middle. Greatest width of eyes approximately one third of the width of the vertex. Pronotum with anterior margin roundingly produced and its posterior margin shallowly concave; its length through middle approximately two thirds of the length of the mesonotum; the disc somewhat depressed, in each half of which just laterad of median line is a faint round depression. Mesonotum with a faint median carina present, laterad of which on either side the disc is somewhat depressed; a faint transverse groove across base of apical projection. Frons slightly wedge-shaped, being broader at its extreme posterior end due to lateral margins converging slightly anteriorly; lateral margins considerably elevated; length and width subequal. Postclypeus dovetailed into frons a short distance only. A distinct median carina present on both frons and clypeus. Teg- mina narrowed greatly at base where their width is only about one third of their width at apex; their greatest width beyond apex of clavus at a point which is at base of apical third ; the costal bor- der narrowly expanded; the cells large, very angular, many of them rectangular or hexagonal in shape, which together with the hyaline texture gives a characteristic look to the wing of resembling panes of glass in a window. Venation of the tegmen as follows: vein R mostly sinuate with a slight angle located just posterior to middle, at which point vein R is equidistant from Scg and Mi^.2; vein M^+g straight, running lengthwise at about median line of tegmen; vein M3+4 simple, but making a bend dorsad to meet Cuja- Female genitalia. Anal flap (10th abdominal segment) two thirds as wide as long, its lateral margins parallel for two thirds of the length, from whence they taper to a rounded apex. The eleventh Doeking: Subfamily Issinae 443 segment only slightly visible and bearing a short dorsal stylus. Ex- ternal valves of ovipositor with their length one third greater than their width and their greatest width through middle. Comparative notes. This species was described from only one fe- male specimen but, since it was distinctly different from any other species in the genus or even in the subfamily, it seemed advisable to describe it even though lacking a series of specimens. The follow- ing structures are the most distinguishing characters: the somewhat anteriorly produced vertex, the wedge-shaped frons, the extreme narrowness at base of the tegmen, and the presence in the tegmen of extremely large angular cells and the broad oblique band of hyaline cells across the corium. Location of types. The holotype female is in the National Mu- seum at Washington, D. C. The labels on the specimen state that it is from the P. R. Uhler collection and was taken in Placer county, California, in October. Dictyssa beameri, n. sp. ORIGINAL DESCRIPTION Size. Length of body from tip of head to apex of tegmina, 2.38 mm. to 2,75 mm. Length of tegmen, 1.875 mm. to 2.13 mm.; width of tegmen, 1.395 mm. to 1.5 mm. This is one of the smallest species in the genus. Color. General color dark brown variegated with yellow on body and many whitish hyaline cells on tegmina. Vertex brown except for a yellow median line and a spot at each lateroposterior corner. Pronotum brown except for a narrow median line, a large spyot oc- cupying each lateral half of disc and all the margins yellow. Meso- notum dark brown through central half, each lateral fourth light yel- low with a brownish spot in center and all margins light. Frons brown except for pale yellow on median carina, a narrow border in- side brown outer margins and an irregular longitudinal streak down middle of each lateral half which converges with the one from oppo- site side below the abbreviated median carina. Postclypeus brown except for anterior border and median carina, which are somewhat lighter. Gena above and around antenna brown; just below a broad pale band. Underside of thorax pale yellow. Abdomen of male pitch brown. Abdomen of female yellowish-brown; genital ap- pendages dark brown. Legs dark brown. Tegmina pitch brown with small whitish hyaline areas in the following places: on clavus in the center of each small cell; on corium, four elongate cells follow- 444 The University Science Bulletin ing the claval suture and a cluster of cells just caudad of claval apex; an oblique band of many cells starting at a point midway of clavus and extending across corium to a point on costal border at about base of apical third of tegmen; in apical border small, round, uniform spots, numbering six or seven; costal border with spots between the brownish veinlets and two clusters of small cells just mesad of costal vein. Structural characteristics. Vertex very broad and short, its lateral margins elevated and slightly converging anteriorly, somewhat de- pressed across disc. Greatest width of eye less than half the width of the vertex. Pronotum with anterior margin rounding and con- siderably produced forward; its length through middle about twice that of vertex at middle; a median carina only faintly discernible; two small, depressed, round spots on disc, one on each side of median line. Mesonotum wdth its anterior margin roundingly produced; in length twice as long as pronotum at middle ; a distinct median carina present and a transverse groove just posterior to anterior margin but not reaching to the sides; a transverse groove at base of apical pro- longation; shallow depression in center of each lateral half. Teg- mina broadest at a point in line with apex of clavus, the apical margin evenly rounding, the costal border moderately expanded; veins prominent, cells small ; veins M and Cu^ branching at a point just anterior to apex of clavus, vein Mi^.o running through center of tegmen and branching at base of apical third, vein R with no acute angle, broadly sinuate, almost equidistant from veins M^ and Sco. Male genitalia. Anal flap (10th abdominal segment) shorter than in most species, its width about three fifths of its length; its lateral margins slightly concave through middle; its posterior margin dis- tinctly notched at middle. The eleventh segment only slightly visi- ble externally and bearing a short dorsal stylus. Harpagones (genital styli) visible externally as two triangular plates, which are broad at base but taper to slender apices beyond which is visible a small portion of the anal tube. Each harpago as viewed from a flattened lateral position (see drawing 16, plate XLI) is roughly rectangular, broadest through apical third, with the ven- tral margin outwardly curved and the dorsal posterior angle pro- longed dorsad into a flat hook with a recurved, pointed apex. At the base of the dorsal projection is a ventrad curving flat external hook. The aedeagus as viewed from the right side is a short tubular structure bearing a flat, pointed projection attached at middle and extending caudad to base of apical fourth. Another well-sclerotized Doering: Subfamily Issinae 445 hook attached to base of aedeagus, whose apex only shows beyond theca as a dorsad-curving stout hook. On the left side the aedeagus is practically covered by the membranous theca, which is prolonged caudad more than in most species as a flat plate. On the right side the theca covers the aedeagus for only the basal third of the latter and has its posterior margin truncate. Comparative notes. This little species is readily distinguished from other species by the following characters: the tegmina are reticulated into many small cells, of which many are whitish hyaline and contrast strongly with the deep fuscous or dark-brown coloration of the rest of the wing; in fact, the pattern of the wing is very similar to that of the genus Dictyobia; the wing venation is distinctive in that the veins all run more lengthwise of the wing; vein R is not angled but only broadly sinuate, and therefore equidistant from veins Sc^ and M^ and vein M^+o divides before apex, which it does not do in any other species. The male genitalia is quite distinct from those of other species (see drawings 7a and 7b, plate XXXIX) . Location of types. This species was described from a holotype male, collected at Carson City, Nev., August 9, 1929, and an allo- type female, same date, by R. H. Beamer. These types are in the Francis Huntington Snow Entomological Collection at the University of Kansas. Dictyssa balli, n. sp. ORIGINAL DESCRIPTION Size. Length of body from apex of head to tip of tegmen, 3 mm. to 3.1 mm.; length of tegmen, 2.25 mm. to 3 mm.; width of tegmen, 1.5 mm. to 1.6 mm. Color. General body color fulvous, washed in brown. Vertex dark brown with a faint yellow median line. Pronotum and mesono- tum brown with a broadish white median line. Frons and clypeus uniform brown. Under side of thorax and abdomen yellow. Legs yellow with brown-tipped spines. Tegmina uniformly pitch brown with a greatly contrasting band of whitish hyaline cells extending lengthwise across corium from base of clavus to a point somewhat anterior to apical margin of corivmi; the entire apical margin studded by pellucid disclike or semicircular spots which are practically all of equal size. Structural characteristics. Vertex broad and short, its lateral margins tapering anteriorly, its length through middle about one half the length of the pronotum. Greatest width of eyes about one 446 The University Science Bulletin half the width of the vertex at its anterior margin. Pronotum with its anterior margin rounding and its posterior margin very shallowly emarginate; two small depressed, round spots on disc; its length through middle one half the length of the mesonotum at middle. Mesonotum with a transverse crease following posterior border of pronotum but not reaching to lateral margins ; a faint median carina present; a shallow depression in disc on each side of median line and a transverse groove at base of apical prolongation. Frons with lateral margins outwardly bulging ; length and width subequal ; me- dian carina distinct, not quite reaching clypeus. Postclypeus moder- ately inflated, dove-tailed into frons to a point about one fourth of the greatest length of the latter. Tegmina differing greatly from other members of the genus by their slenderness; each tegmen in length two fifths longer than wide, its greatest width at a point in line with apex of clavus ; the apical margin sloping gently from apex of clavus, thence evenly rounding to the costal border; the costal border moderately expanded and crossed by numerous dark-brown veinlets; an oblique hyaline band present, made up of a spot on base of clavus and four cells on corium, the largest one of which is an elongate central cell between veins R and M and which is approxi- mately twice as long as wide and has its border studded by 8 to 9 spurs of veins ; the second largest cell lying posterior to this, usually angular in shape and about one fourth or one third of the size of the central cell ; veins Sc and R running parallel and very close together for about two thirds of the length of the tegmen, at which point vein R makes a distinct bend toward vein Mi^,, which makes it in this region nearer to the latter vein than to vein Sco; M^^, simple; vein Cui branching somewhat more anteriorly than M. Male genitalia. Anal flap (10th abdominal segment) narrow at extreme base, then broadening through basal third, from whence it narrows to a truncate apex ; its length twice its width ; a short dorsal stylus of the eleventh segment present, but none of the latter show- ing externally. Harpagones (genital styli) visible externally as broad, triangular, fiat plates whose apices are sharply pointed; the anal flap not visible to any extent beyond their apices. From a flattened lateral view (see drawing 24, plate XLI) each process is characteristically sub- quadrangular, broadest through apical third and with its posterior Doeking: Subfamily Issinae 447 dorsal corner prolonged dorsad into a short, pointed, slightly re- curved process, at base of which is a short, broad, recurved external hook. The aedeagus is a curved tubular structure. On the right side it bears a long, flat, sharply pointed projection attached midway of its length and extending posteriorly to about the base of the apical sixth of the aedeagus ; also two hooks attached near base which are covered by the theca at their bases, the dorsal one extending beyond the theca about twice as far as the other, with its extereme apex curved ventrad and sharply pointed, the ventral one almost straight and bluntly pointed. On the left side the aedeagus bears also a flat, pointed projection which is attached somewhat posterior to the middle and extends almost to apex of the aedeagus. No additional basal hooks are found on this side. The theca on the right side covers the aedeagus at base for about one third the length of the latter; its posterior margin is shallowly concave. On the left side the theca is about the same as on the right side, except that its dorsal posterior angle is extended posteriorly as far as the middle as a slen- der, sharply pointed projection which resembles an aedeagal hook and its posterior margin seems to be divided at middle. Female genitalia. Anal flap about one third longer than wide, sharply tapering to a blunt, narrowed apex. The eleventh segment scarcely visible, its dorsal stylus moderately long. The external valves of the ovipositor broadest through middle, in length about one third longer than wide. Comparative notes. This species is easily recognizable by its elongate form. Dr. E. D. Ball, recognizing this species as new, was preparing to call this species elongata, but when the writer started revising this genus he very graciously relinquished any prior claim to it and told the writer to describe it. For this generosity and mag- naminity of spirit the writer preferred to name the species in his honor, although elongata would have been a very descriptive name. It is much the longest and narrowest species in the genus. It can be recognized furthermore by the fact that the hyaline band on tegmen runs more nearly lengthwise than in other species, is abbreviated be- fore reaching apex and has the central clear cell about twice as long as wide. For further comparative notes see the discussion under this heading in the description of mutata. The male genitalia are distinctive. 448 The University Science Bulletin Notes on distribution. A long series of this species was taken by R. H. Beamer in August at Campo, Cal. Location of types. In the Francis Huntington Snow Entomo- logical Collection at the University of Kansas. Holotype, allotype, and paratypes from Campo, Cal, August 10, 1935, collected by R. H. Beamer. Dictyssa monroviana, n. sp. ORIGINAL, DESCRIPTION Size. Length of body from tip of head to tip of tegmen, 2.4 mm. to 2.9 mm. Length of tegmen, 2 mm. to 2.3 mm.; width of tegmen, 1.5 mm. to 1.6 mm. Color. A fuscous brown species marked with hyaline on the teg- mina. Vertex uniformly yellowish-brown with a faint median yel- low line. Pronotum and mesonotum, uniformly yellowish or fus- cous brown. Frons and clypeus uniformly brown. Underside of thorax and legs yellowish-brown. Abdomen dark brown. Tegmina with the oblique hyaline band very conspicuous and proportionally long; large, round or hemispherical whitish hyaline cells, alternat- ing with small ones across apical margin and a cluster of three to four small hyaline cells just beyond apex of clavus in cells Cu^a and Cuibi rest of wings dark fuscous with veins frequently lighter in color. Structural characteristics. Vertex slightly narrowed anteriorly, depressed through middle, its length through middle about one half the length of the pronotum. Greatest width of eye less than one half the width of vertex. Pronotum with all its margins elevated, usually no median carina present; two small faint depressed spots on disc. Mesonotum with median carina lacking or at most only faintly indicated; a transverse groove following anterior margin, but stopping either side before reaching lateral margins; length through middle twice that of pronotum. Frons with lateral margins parallel, considerably elevated ; median carina distinct, but abbreviated before reaching apex; a broad depression on each side between lateral mar- gin and median carina. Postclypeus with posterior margin angu- larly produced into the frons for a distance equal to about one third the length of the latter. Tegmina about one fourth longer than wide, broadest through base of apical fourth; its posterior margin truncately rounding, the costal margin broadly expanded ; cells large with the hyaline area large in proportion to the opaque regions, the oblique hyaline band starting on clavus where it is very broad and Doeeing: Subfamily Issinae 449 continuing across coriiim as about five very large angular cells, the largest one being on the disc between veins R and M and which is elongate oval in shape, in size at least twice as long as wide, and with its margins studded by 9 to 10 spurs of veins; two other large cells ventrad to this, the one on coastal border especially so; typi- cally four to five large round or oval cells present on apical border, with smaller ones in between ; vein R at apex of central hyaline cell making an acute angle which is at a point nearer to vein M^+a than Sc2 and as in mutata more posterior in the wing, usually at base of apical fourth; vein M^^g simple. Male genitalia. Anal flap (10th abdominal segment) about twice as long as wide and with an evenly rounding apex. Eleventh seg- ment scarcely visible, its dorsal stylus very long and slender, reach- ing almost to apex of the flap. Harpagones (genital styli) visible externally as triangular plates, broad at base and with tapering apices. Each harpago, from a flattened lateral view (see drawing 23, plate XLI), is rectangular in outline and has its posterior, dorsal corner prolonged dorsad into a slenderly pointed projection at the base of which is a slender, re- curved, external hook. The aedeagus is a curved, tubular structure, shorter than in many species. On the right side it bears a flat, sharply pointed process, attached at base of apical third and extending almost to apex. At- tached to its base are two well-sclerotized hooks which are covered at their bases by the theca but extend beyond it, the ventral one to a point beyond middle of aedeagus and the dorsal one somewhat anterior to it. The ventral hook is more slender and has its pointed apex curved ventrad. The dorsal hook is blunt and nearly straight. On the left side the aedeagus bears no basal hooks. There is present a flattened, sharply pointed projection attached at base of apical third which almost reaches the apex. The theca on the right side covers slightly over one third of the aedeagus at base and has the posterior margin somewhat triangularly produced caudad. On the left side the theca also covers the basal third of the aedeagus, but has its dorsal angle prolonged caudad as a long, sharply pointed flap which in typical forms reaches almost to apex of the aedeagus. Comparative notes. This species is recognized by the large hya- line cells and spots of the tegmina, especially that of the very large central cell, which is at least twice as long as broad. It more nearly resembles D. mutata and D. obliqua in size and shape. It resembles D. obliqua by having the head and thorax fuscous or darkish, but it 29—6037 450 The University Science Bulletin differs from it by having the angle of vein R located more poste- riorly, approximately at base of apical fom-th. It differs externally from D. mutata by having the head and thorax darkish or washed with fuscous, while in the latter the head is bright yellow; also, be- cause vein R in monroviana approaches nearer to vein M^+o than vein Sc, The aedeagus is more similar to D. mutata than that of any other species. Typical examples of the two species show the following differences: in D. monroviana the thecal hook is longer and is always visible, extending to middle at least and in some cases almost to tip of aedeagus, while in D. mutata the thecal hook is transparent, not readily distinguishable and does not extend beyond middle ; as a general rule the dorsal aedeagul hook is proportionally smaller in D. mutata than in D. monroviana. Many gradating forms of the aedeagus between these two have been found, however, but all dark-headed specimens with the large hyaline cells have been placed under monroviana. For further discussion see notes under this heading in the description of mutata. Location of types. Holotype male, allotype female and eleven paratypes, Monrovia, Cal., August 27, 1935, collected by R. H. Beamer, in the Snow Entomological Collection at the University of Kansas. In the same collection other paratypes from Laguna Beach, August 25, 1933, from Claremont, Cal., Aug. 29, 1935, and a long series from Orange county, California, collected by R. H. Beamer in August, 1929, and two paratypes from San Jacinto mountains, California, collected by L. D. Anderson and R. H. Beamer in August, 1929. Dictyssa maculosa, n. sp. ORIGINAL DESCRIPTION Size. Length of body from apex of head to tip of tegmen, 2.5 mm. Length of tegmen, 2.13 mm.; width of tegmen, 1.38 mm. Color. General color similar to D. mutata in that both species have dark-brown tegmina with bright-yellow heads and pronota. Vertex bright yellow with margins etched in dark brown. Eyes reddish-brown. Pronotum uniformly bright yellow except darker at extreme lateral margins. Mesonotum bright yellow except fuscous at lateral corners and in the depressed area either side of the median carina. Front uniformly yellow in the type female with margins faintly etched in dark brown; in the paratype female a group of dark spots speckled along lateral margins. Postclypeus bright yel- low with traces of reddish or brownish oblique stripes on each side. Gena and rest of head bright yellow except pedicel of antenna, which Doering: Subfamily Issinae 451 is fuscous. Underside of thorax bright yellow. Legs bright yellow with carinae of femora and tibiae and tips of tarsi and tarsal claws fuscous. Abdominal segments mostly yellow, a little darker at sides. External valves of ovipositor brown with their median mar- gins much darker. Tegmina blackish-brown and very opaque ex- cept for the numerous whitish hyaline spots which in the main occur in the following regions: an oblique band starting on clavus where it occupies the basal third, then extends across coriimi to just beyond middle, at which point it joins a transverse band of hyaline cells, about equal in size and which extends from a point just posterior to apex of clavus across to the costal border; between the larger transverse band and the apical margin a second but more abbreviated transverse band made up of four ovate spots, the apical margin studded with ovate or round spots, the larger ones, of which there are four, alternating with either one or two small ones; the costal border with numerous white spots between the dark margined veinlets; a group of three to four uniformly ovate spots in the cells between veins R and Sc. Structural characteristics. Vertex not greatly produced beyond the eyes; its anterior margin almost straight; its lateral margins distinctly converging anteriorly ; all margins greatly elevated ; length through middle a little over one half the length of pronotum at middle; a very shallow round depression in the mesoposterior corner of each lateral fourth. Greatest width of eyes one third the width of the vertex. Pronotum with anterior margin strongly roundingly emarginate; posterior margin shallowly concave; all margins greatly elevated; a median carina only faintly indicated, laterad of which on each side in the disc is a faint round depressed spot. Mesonotum triangular; the disc depressed; a transverse ridge across middle half, running parallel with anterior margin ; a median carina present and a faint groove indicated across base of apical extension; length about twice that of pronotum. Frons with anterior margin almost straight and lateral margins slightly bulging outwardly, all margins considerably elevated; a distinct median carina present on posterior two thirds but completely gone from apex. Postclypeus dovetailed into frons for not more than one sixth of the length of the latter and moderately inflated. Tegmina with apical and costal margins rounding, the latter expanded somewhat but not greatly reflexed; greatest width of tegmen near base of apical fourth ; the largest cells of wing found in the oblique hyaline band, consisting of one large elongate cell about one third to one half longer than wide, whose margins are studded with triangular dark-brown spurs of veins num- 452 The University Science Bulletin bering about eight to nine and whose apex reaches not far beyond middle of the wing; a second large hyaline cell somewhat anterior to the central cell, which is angular in shape, and a third bilobed and somewhat smaller posterior cell which extends into the transverse row of small, ovate hyaline cells. Wing venation as follows: veins Sc and R united for only a short distance from base of wing, vein R broadly sinuate with a slight angle indicated at apex of the cen- tral hyaline cell, at which point R is equidistant from veins Scg and M1+2; vein Mi+2 makes the characteristic bend towards the costal border; vein M3+4, separated at base of apical third or else the tip of CUia, which seems to be lost in the apical network, has secondarily united to the stem of M3+4, thus giving that vein a two-branched appearance. Female genitalia. Anal flap (10th abdominal segment) broad at base, tapering to a bluntly rounding apex; in length one-third longer than wide. The eleventh abdominal segment inconspicuous, but bearing a slender, tubular dorsal stylus. The external valves of the ovipositor conspicuous, their length not quite twice the width, their greatest width at middle. Comparative notes. This species is easily separated from other species in the genus by the bright-yellow head and thorax, which contrasts greatly with heavy pitch-brown tegmina, by the large number of ovate or round whitish hyaline cells which are present not only as an oblique band across corium but are arranged in two transverse rows through apical third of tegmen and also in apical region between the costal vein and vein R. Location of types. Holotype female, collected at Anza, Cal., Aug. 6, 1935, by R. H. Beamer, and one paratype female, collected at Idyllwild, Cal., Aug. 8, 1935, by R. H. Beamer, in the Francis Hunt- ington Snow Entomological Collection at the University of Kansas. 454 The University Science Bulletin PLATE XXXVI Figure 1. Diagi-am of a tegmen. 2. Lateral view of Dictyssa fusca. 3. Lateral view of Dictyssa clathrata. 4. Lateral view of Dictyssa monroviana. 5. Lateral view of Dictyssa ovata. 6. Lateral view of Dictyssa maculosa. 7. Lateral view of Dictyssa obliqua. 8. Lateral view of Dictyssa marginepunctata. 9. Lateral view of Dictyssa beameri. 10. Lateral view of Dictyssa quadravitrea. IL Lateral view of Dictyssa fenestrata. 12. Lateral view of Dictyssa mutata. 13. Lateral view of Dictyssa transversa. 14. Lateral view of Dictyssa balli. 15. Lateral view of Dictyssa areolata — type specimen. 16. Lateral view of Dictyssa areolata — perfect specimen. Doering: Subfamily Issinae 455 PLATE XXXVI 4. D.monroviona 7.Di obllqua 10 b quodrovifrea S. D.ovala 8 .O.marginepunctata II.D.fcnestrala 13. D transversa SD.elolhrata 6.0. maculosa 9.D.beameri l2.D.mijtala K.D.boIli IS.D.areolota --type le.D.areoIata 456 The University Science Bulletin PLATE XXXVII Figure 1. Dorsal view of head and thorax of Dictyssa clathrata. 2. Dorsal view of head and thorax of Dictyssa ficsca. 3. Dorsal view of head and thorax of Dictyssa ovata. 4. Dorsal view of head and thorax of Dictyssa transversa. 5. Dorsal view of head and thorax of Dictyssa mutata. 6. Dorsal view of head and thorax of Dictyssa obliqua. 7. Dorsal view of head and thorax of Dictyssa areolata. 8. Dorsal view of head and thorax of Dictyssa beameri. 9. Dorsal view of head and thorax of Dictyssa marginepunctata. 10. Dorsal view of head and thorax of Dictyssa maculosa. 11. Dorsal view of head and thorax of Dictyssa balli. 12. Dorsal view of head and thorax of Dictyssa quadravitrea. 13. Dorsal view of head and thorax of Dictyssa fenestrata. 14. Dorsal view of head and thorax of Dictyssa monroviana. 15. Cephalo-ventral aspect of head of Dictyssa clathrata. 16. Cephalo-ventral aspect of head of Dictyssa fusca. 17. Cephalo-ventral aspect of head of Dictyssa areolata. 18. Cephalo-ventral aspect of head of Dictyssa balli. 19. Cephalo-ventral aspect of head of Dictyssa transversa. 20. Cephalo-ventral aspect of head of Dictyssa monroviana. 21. Cephalo-ventral aspect of head of Dictyssa jenestrata. 22. Cephalo-ventral aspect of head of Dictyssa maculosa. 23. Cephalo-ventral aspect of head of Dictyssa quadravitrea. 24. Cephalo-ventral aspect of head of Dictyssa ovata. 25. Cephalo-ventral aspect of head of Dictyssa beameri. 26. Cephalo-ventral aspect of head of Dictyssa mutata. 27. Cephalo-ventral aspect of head of Dictyssa obliqua. 28. Cephalo-ventral aspect of head of Dictyssa marginepunctata. Doering: Subfamily Issinae 457 PLATE XXXVII I.O.clafhratQ 2.D. fusca 5.D. mutato 4D. transverso 7. D.areolato 8.D.beameri 9.D.marginepunctata |0D. maculosa 13. D. fenestrato 2I..D.fenestrata 22.D.maculosa 23. D.quadravitreo 24D.ovata 25.D. beameri 26.D.mutata 27. D.obliqua 28.D.marginepunctato 458 The University Science Bulletin PLATE XXXVIII Figure la. Lateral aspect as viewed from left side of male genitalia of Dictyssa clathrata. lb. Lateral aspect as viewed from right side of male genitalia of Dictyssa clathrata. 2a. Lateral aspect as viewed from right side of male genitalia of Dictyssa fusca. 2b. Lateral aspect as viewed from left side of male genitalia of Dictyssa fusca. 3a. Lateral aspect as viewed from right side of male genitalia of Dictyssa ovata. 3b. Lateral aspect as viewed from left side of male genitalia of Dictyssa ovata. 4a. Lateral aspect as viewed from left side of male genitalia of Dictyssa areolata. 4b. Lateral aspect as viewed from right side of male genitalia of Dictyssa areolata. 5a. Lateral aspect as viewed from left side of male genitalia of Dictyssa balli. 5b. Lateral aspect as viewed from right side of male genitalia of Dictyssa balli. Doering: Subfamily Issinae 459 PLATE XXXVIII 460 The University Science Bulletin PLATE XXXIX Figure 6a. Lateral aspect as viewed from right side of male genitalia of Dictyssa fenestrata. 6b. Lateral aspect as viewed from left side of male genitalia of Dictyssa fenestrata. 7a. Lateral aspect as viewed from left side of male genitalia of Dictyssa beameri. 7b. Lateral aspect as viewed from right side of male genitalia of Dictyssa beameri. 8a. Lateral aspect as viewed from left side of male genitalia of Dictyssa transversa. 8b. Lateral aspect as viewed from right side of male genitalia of Dictyssa transversa. 9a. Lateral aspect as viewed from left side of male genitalia of Dictyssa obliqua. 9b. Lateral aspect as viewed from right side of male genitalia of Dictyssa obliqua. 10a. Lateral aspect as viewed from left side of male genitalia of Dictyssa monroviana. 10b. Lateral aspect as viewed from right side of male genitalia of Dictyssa monroviana. 11a. Lateral aspect as viewed from left side of male genitalia of Dictyssa marginepunctata. lib. Lateral aspect as viewed from right side of male genitalia of Dictyssa marginepunctata. 12a. Lateral aspect as viewed from left side of male genitalia of Dictyssa mutaia. 12b. Lateral aspect as viewed from right side of male genitalia of Dictyssa mutata. Doering: Subfamily Issinae 461 PLATE XXXIX 462 The University Science Bulletin PLATE XL Figure 1. Dorsal aspect of the tenth abdominal segment (anal flap) of the male of Dictyssa ovata. 2. Same for Dictyssa transversa. 3. Same for Dictyssa monroviana. 4. Same for Dictyssa balli. 5. Same for Dictyssa beameri. 6. Same for Dictyssa fusca. 7. Same for Dictyssa clathrata. 8. Same for Dictyssa ohliqua. 9. Same for Dictyssa mutata. 10. Same for Dictyssa fenestrata. 11. Same for Dictyssa areolata. 12. Same for Dictyssa marginepunctata. 13. Ventral aspect of the tip of the abdomen of the male of Dictyssa beameri. 14. Same for Dictyssa transversa. 15. Same for Dictyssa balli. 16. Same for Dictyssa marginepunctata. 17. Same for Dictyssa monroviana. 18. Same for Dictyssa fenestrata. 19. Same for Dictyssa obliqua. 20. Same for Dictyssa clathrata. 21. Same for Dictyssa ovata. 22. Same for Dictyssa mutata. 23. Same for Dictyssa fusca. Doering: Subfamily Issinae 463 PLATE XL /= ^?ii -■,? \^ eOfusco 2.D tronsverso ' 4D. bafli 3D/nonroviano 5 0 beomen 7 O.clothToto IC ovota 2I.D ovato ZZ.O.Riutato 23.D.fusca 464 The University Science Bulletin PLATE XLI Figure 1. Dorsal aspect of the tenth abdominal segment of the female of Dictyssa ovata. 2. Same for Dictyssa transversa. 3. Same for Dictyssa marginepunctata. 4. Same for Dictyssa monroviana. 5. Same for Dictyssa mutata. 6. Same for Dictyssa obliqua. 7. Same for Dictyssa fusca. 8. Same for Dictyssa clathrata. 9. Same for Dictyssa jenestrata. 10. Same for Dictyssa balli. 11. Same for Dictyssa areolata. 12. Same for Dictyssa quadravitrea. 13. Same for Dictyssa maculosa. 14. Same for Dictyssa beameri. 15. Lateral aspect of harpago (male genital stylus) of Dictyssa jenestrata. 16. Same for Dictyssa beameri. 17. Same for Dictyssa ovata. 18. Same for Dictyssa transversa. 19. Same for Dictyssa marginepunctata. 20. Same for Dictyssa clathrata. 21. Same for Dictyssa jusca. 22. Same for Dictyssa mutata. 23. Same for Dictyssa monroviana. 24. Same for Dictyssa balli. 25. Same for Dictyssa areolata. 26. Same for Dictyssa obliqua. Doering: Subfamily Issinae 465 PLATE XLI 18.D- tronsversa 19. D morglnepunctata ZO.D.clathrato 2I.D.fusco E2X)mutoto 23.0.monroviana 24 £i ball 30—6037 466 The University Science Bulletin PLATE XLII Figure 1. Ventral aspect of the tip of the abdomen of the female of Dictyssa marginepunctata. 2. Same for Dictyssa quadravitrea. 3. Same for Dictyssa monroviana. 4. Same for Dictyssa jenestrata. 5. Same for Dictyssa beameri. 6. Same for Dictyssa areolata. 7. Same for Dictyssa balli. 8. Same for Dictyssa clathrata. 9. Same for Dictyssa maculosa. 10. Same for Dictyssa niutata. 11. Same for Dictyssa jusca. 12. Same for Dictyssa transversa. 13. Same for Dictyssa obliqua. 14. Same for Dictyssa ovata. Doering: Subfamily Issinae 467 PLATE XLII l2.D.traiKversa IS.D.obliquo l4.D.ovafa THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 18. Description of a new Mexican Subspecies of Sceloporus spinosus Wiegmann (Lacertilia) HOBART M. SMITH Department of Zoology, University of Kansas Abstract: Sceloporus spinoaus caeruleopunctattis ssp. nov., is described from Cerro de San Luis, Oaxaca, Mexico. Types at University of Kansas. AMONG the specimens collected in Mexico during the summer of 1935 by Dr. Edward H. Taylor and myself are four which represent an undescribed, distinct subspecies of Sceloporus spinosus Wiegmann, which may be described as follows:* Sceloporus spinosus caeruleopunctatus ssp. n. Holotype. EHT & HMS 8467, collected on the slopes of Cerro de San Luis, about 15 miles N. of Oaxaca, Oaxaca, August 5, 1935, by Hobart M. Smith. Paratypes: EHT and HMS 8464-6, from the type locality, same collector. Diagnosis. Closely related to spinosus spinosus, differing in average scale characters and dorsal coloration. Dorsal scales from occiput to base of tail, 31 to 34 (30 or less in s. spinosus) ; ventrals, 47 to 52 (average, 45.6 in s. spinosus) ; femoral pores 9 to 13; lorila- bials usually reduced to one row below subocular (rarely in s. spi- nosus) ; supraoculars, five to seven (rarely six, usually four in s. spi- nosus) ; prefrontals usually in contact medially (seldom in s. spi- nosus) ; some of the dorsal scales dark blue, coloration otherwise much as in s. spinosus. Description oj holotype. Head scales smooth, slightly convex, pitted; interparietal pentagonal; parietals single on either side, about two thirds size of interparietal; frontoparietals single on either side, more or less square, separated from each other medially by contact of frontal and interparietal; posterior section of frontal * I am greatly indebted to Dr. Edward H. Taylor for making possible the study of these specimens, and for other courtesies and assistance. The study of the genus Sceloporus has been forwarded by grants from the Graduate Research Fund of the University of Kansas. (469) 470 The University Science Bulletin three fourths or four fifths size of anterior section; prefrontals in contact medially; three frontonasals, the lateral scales in contact with both canthals, median frontonasal distinctly larger than lateral frontonasals, and slightly larger than prefrontals; two pairs of in- ternasals, the anterior pair separated from the rostral by a row of scales continuous with lorilabials; both pairs of internasals sepa- rated from first canthal by a small scale; supraoculars five-five, the anterior the smaller, the fourth slightly larger than others; a single, complete row of small, usually keeled scales separating supraoculars from median head scales; one complete and another incomplete row of scales separating supraoculars from superciliaries (except the first supraocular, which is in contact with first superciliary) ; five superciliaries visible from above; two canthals, the first not touch- ing lorilabials nor forced above canthal ridge, the second forming a small portion of the superciliary series; subnasal present, approxi- mately size of first canthal; loreal present, more or less square; pre- ocular entire; subocular moderate in size, followed posteriorly by two small, strongly keeled postoculars; lorilabials in two rows, not reduced to one below subocular (usually reduced to one row) ; su- pralabials four, infralabials five-six to a point below middle of eye. Mental pentagonal, with a labial border about three fifths that of rostral; mental followed posteriorly by three pairs of postmentals, the scales of only the anterior pair in contact medially; outer row of labiomentals separated from mental by partial contact of first postmental and first infralabial; inner row of labiomentals terminat- ing below third infralabial; gular scales smooth, with one or two apical notches (usually one) ; anterior gular scales rounded, reduced in size; posterior gular scales subequal in size; gular scales below ear weakly keeled; scales in gular fold region with two or three apical notches. Auricular lobules three, the upper the longest, but not reaching across ear; auricular lobules larger than preceding temporal scales; temporal scales weakly keeled, weakly mucronate toward eye, more strongly toward ear, somewhat smaller than largest scales between ear and lateral nuchal fold; ridge of skin overlapping lateral nuchal fold surmounted by strongly keeled, very strongly mucronate scales, which continue in an irregular row to a point below ear; scales be- tween ear and insertion of foreleg keeled, rather strongly mucronate, strongly denticulate. Dorsal scales keeled, rather strongly mucronate, denticulate, in parallel rows, the scales toward occiput reduced in size; median Smith: Scelopokus Spinosus 471 lateral scales of somewhat the same general character as the dorsal scales, somewhat smaller than dorsal scales, somewhat more strongly keeled, in oblique rows; lateral scales toward axilla and groin be- coming smaller, those in axilla almost granular, those in groin one half as large as median lateral scales, imbricating; ventral scales about one half or three fifths size of dorsal scales; scales on chest slightly larger than those on belly; interfemoral and preanal scales not or very slightly smaller than scales on belly; some of preanal scales rather porelike; all ventral scales smooth, with a single apical notch ; dorsal scales on rump very slightly reduced in size ; basal caudals on dorsal surface as large as largest dorsal scales on body; dorsal caudals strongly keeled, strongly mucronate, not or weakly denticulate, becoming smaller and more strongly keeled toward tip of tail; subcaudals smooth, rounded, becoming keeled distally on tail; enlarged postanals present, separated from each other medially by two small scales. Dorsal scales of foreleg keeled, mucronate, denticulate, those on upper foreleg about one third size of dorsal scales on body, some- what larger than those on lower foreleg; scales at elbow reduced in size; ventral scales of lower foreleg about as large as dorsal scales of same member, keeled except at extreme anterior edge; ventral scales on upper foreleg smooth (keeled anteriorly), usually notched, about one half or two thirds size of ventrals of lower forelegs, be- coming smaller near axilla; lamellar formula for fingers 8-12-16- 16-9 (8-12-16-16-10). Dorsal scales of hind leg rather strongly keeled, mucronate, weakly denticulate, those on thigh somewhat larger than those on upper foreleg; dorsals on shank somewhat larger than those on thigh; ventral scales of shank smooth, notched, as large as dorsal scales of same member; scales on anterior surface of thigh nearly smooth, acuminate, with a single notch on either side of apex, the scales be- coming cjuite smooth, with a single apical notch, and much smaller toward series of femoral pores; scales in a median area on posterior surface of thigh keeled, mucronate, much smaller than preanal scales; no postfemoral dermal pocket; lamellar formula for toes 8-13-16-19-12 (8-13-17-19-12). Color. General dorsal coloration brownish-yellow; a dorsolateral light line on each side, about one and one half scales wide, extending from temporal region onto base of tail; below this the ground color is darker, sometimes concentrated into broad, indistinct dark bars separated by narrow lighter areas, not passing onto ventral surface 472 The University Science Bulletin of abdomen; between the dorsolateral light lines are a number of dark-blue scales (the color usually fades in preservative) apparently- arranged in two irregular rows ; the heads of the two males are slate (rather dark in type), with numerous white flecks; among these the most distinct are: a light spot on each parietal, a light pineal spot with a small, dark, central spot, a light spot on each side at the posterior edge of interparietal, a light spot on each frontoparietal, a light, narrow bar across posterior section of frontal, and a light spot at the posterior, medial edge of the third supraocular; in the males an indefinite, brownish band crosses the head in front of the eyes, and is preceded by a darker brown area extending to the inter- nasals; the internasal area, to the rostral, is light brown; the labia are irregularly marked with dark and very light brown ; a dark spot in front of arm, extending dorsally a short distance, with a few scales of dark-blue color; the black shoulder spots are narrowly continuous with each other across neck; in the type, the anterior part of the gular area is white, the posterior part blue ; in the male paratype, the whole throat is dark blue; in both specimens black, or dark blue, convergent bars are present in the gular region; the chest, a narrow area down middle of abdomen, prenanal region and the posterior part of the ventral surface of the thighs are white ; on each side of belly is a broad area of blue, bordered medially by a narrow area of black or dark blue; the groin is black or dark blue, the color extending onto thighs; the chest has a few irregular spots of black. In the females two rows of irregular, dark-brown spots are pres- ent between the dorsolateral light lines; the throat has indistinct bluish bars, and the sides of the abdomen have a bluish tinge. The following notes from the field catalogue are pertinent: ^' Large male. Lighter parts of labial region and lighter parts of back tinged with reddish-orange, more distinct on head. Posterior gular region dark blue; sides of belly lighter, tinged with light green. Some of the scales of the back between the dorsolateral light lines dark blue. Smaller male. Lacks orange color of head. Female. In life, when shot, the blue spots on the back were very distinct. A few hours later, before preservation, the color had disappeared." Habits and habitat. All specimens were found running about on the ground at low elevations, in more or less open areas, usually near rocks. They always ran into rocks or bushes at the first indica- tion of danger, and were rather difficult to collect. Smith: Sceloporus Spinosus 473 Remarks. According to my notes, taken hurriedly on a recent short, trip to eastern museums, the following specimens apparently belong to this subspecies: AMNH 18804-14 (Tlacolula to San Pablo Mitla, Oaxaca, July 11-13, 1920, Paul D. R. Rlithling) ; FMNH 1011 (Oaxaca, Oaxaca, June, 1901, S. E. Meek) ; USNM 47534-5, 47396-7 (Oaxaca, Oaxaca, June 14, 1894, Nelson & Goldman). Measurements (in mm.) and Scale Counts Number 8465 8466 8464 8467 Snout to vent 80.0 15.0 16.9 54.0 20.7 10.1 15.9 0.2 19-20 12-13 9 100.0 34 51 42 82.0 15.0 18.4 58.7 22.4 9.7 16.8 5.1 20-21 10-12 cf 115.0 31 47 38 82.5 15.0 18.2 53.5 20.5 9.2 14.6 6.2 21-22 11-12 9 103.0 33 52 43 94.0 Snout to occiput 17.0 Snout to ear 20.5 Hind leg 61.6 Fourth toe 23.0 Fifth toe 11.0 Tibia 17.0 Scales to head length 6.0 Lamellae, fourth toe 19-20 Femoral pores 9-12 Sex (^ Tail 126.0 Dorsals 31 Ventrals 52 Scales around body 40 THE UNIVERSITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 19. Notes on the Herpetological Fauna of the Mexican State of Sonora EDWARD H. TAYLOR, Department of Zoology, University of Kansas Abstract: The report is based largely upon a collection of reptiles and amphibians, now at the University of Kansas, secured by Edward H. Taylor in Sonora during parts of the summer of 1934. An ecological classification is given of the species discussed (five of amphibians, eighteen of lizards, sixteen of snakes, and two of turtles). Cneniidophorus burli sp. nov. is described from La Posa, near Guaymas. An annotated bibliography of literature concerning the herpetological fauna of the state is appended. THE fauna of Sonora is of special interest to students of Ameri- can herpetology, because of the proximity of this region to the states of Arizona and New Mexico, and the fact that the southern boundary of the state is not far from the southern limit of distribu- tion of the species characteristic of the Sonoran faunal region, at least of those forms that are adapted to a lowland habitat. At Mazatlan in southern Sinaloa, this fauna has been largely replaced by other species, a large number of which show strong Central American affinities. Although much of the state has not been explored herpetologically, a few collections have been made, chiefly in the coastal region about Guaymas and along the highway between Nogales and Guaymas, A few specimens w^ere obtained by members of the Mexican Boundary Commission between the years 1852 and 1854, and it appears that certain of the "Sonora" localities were incorporated in Arizona and New Mexico after the Gadsden Purchase. Specimens from the commission are in the National Museum. Prof. J. T. Lovewell and Mr. Heiligbrodt collected in Sonora at a much later date. This collection, which was presented to Washburn College, Topeka, Kan., was made the basis of a special report by Cragin (1884). A collection made by Dr. Gustav Eisen and Walter Bryant in April and May, 1892, is in the California Academy of Sciences. It was the subject of a report by Van Denburgh (1897). (475) 476 The University Science Bulletin Certainly the most important collection made prior to my own is one made by Morrow J. Allen, Jean Piatt and John Scofield, spon- sored by the Museum of Zoology, University of Michigan. The collection numbered 326 specimens, 59 amphibians and 267 reptiles, and was obtained chiefly at Puerto, Noria, Hermosillo and Guaymas in June and July, 1932. Allen (1933) has published data on this collection, listing four amphibians, twelve lizards, six snakes (the report on the two species of rattlesnakes is accredited to H. K. Gloyd) and one turtle. I spent the time between June 19 and July 16, 1934, in Sonora studying the herpetological fauna. A considerable collection was made, chiefly in the localities previously visited by Allen, Piatt, and Scofield. A brief itinerary from my diary follows: June 19. Entered Sonora at Nogales. Collected a few specimens along the road, and made a late afternoon camp, 53 miles south of Nogales. June 20 and 21. Collected along the road, and camped at Noria, the evening of the 20th. On both nights some time was spent in the field collecting with a light. June 22. A stop was made 30 miles south of Noria, and later I pitched camp about five miles southwest of Hermosillo, in a large boulder field near low mountains. June 23 to 25. Collected at Hermosillo, spending a part of each night in the field. I broke camp the afternoon of the 25th and made a camp 51 miles south of Hermosillo, remaining there until noon of the following day. June 26. Spent the morning afield, then drove to Guaymas. Dur- ing the evening I collected near the beach at Miramar, a small resort three miles northwest of Guaymas. June 27. Collected about Miramar and later made camp at LaPosa, about ten miles northwest of Guaymas. This place con- sists of three small houses and a nearby well, situated within a quarter mile of the beach. June 28 to July 16. With the La Posa camp as a base, collect- ing was carried on within a five-mile radius. Several hours were spent collecting in the field with a light on most of the nights. In the immediate vicinity of La Posa are a few low mountains rising from sea level. To the north about one and one half miles there is a chain of low mountains, and tw^o miles to the west are other low peaks in the vicinity of San Carlos Bay. The intervening terrain is gravelly and covered with sparse bush and cacti, with occasional Taylor: Fauna of Son or a 477 stunted trees. At this season the country was arid and none of the small streams carried any water. With the aid of friendly vaqueros of La Posa two small springs were located about two miles to the north of my camp. Here a few hylas and the ubiquitous Rana pipiens were found by the springs; elsewhere only a single am- phibian was taken, this, a large Bufo alvarius, obtained at the La Posa well, at night. During the period I spent at this camp, a few specimens were also collected at Miramar and along the road to Guaymas. Two trips (one during the day, one at night) were made to Empalme, ten miles southeast of Guaymas. On June 16 I journeyed south into Sinaloa, and later into Nayarit to continue collecting. On my return to the north, I spent a few days, August 4-8, in the general vicinity of Guaymas (Miramar and La Posa). A few specimens were taken along the highway on the return to the border. I arrived at Nogales August 10. In the general region about Guaymas the mountains come down to the sea, but at various points there are flat, sandy beaches limited by a dyke of sand built up by the waves. The line where this dyke and the shrub met was an especially fine locality for col- lecting. This terrain offered shelter for numerous rodents, insuring a constant food supply for the snakes. An ecological classification would include four general habitats where collecting was done: beach line; low gravelly flats; low moun- tains; and springs. The fauna of each is recorded below. Beach Line Callisaurus inusitatus Heloderma suspectum Masticophis flagellum frenatum Phyllorhynchus decurtatus decurtatus Crotalus cinereous Springs Ilyla arenicolor Rana pipiens Thamnophis eques Ficimia desertorum (accidental) Low Mountains Phyllodactylus homolepidurus Sauromaulus hispidus Ctenosaura hemilopha Uta taylori Cnemidophorus burti Constrictor constrictor imperator Masticophis piceus Crotalus molossus molossus Crotalus tigris Gopherus agasaizii Low Brushy Flats Coleonyx variegatus Dipso-saurus dorsalis sonorievsis Holbrookia elegans thermophila Uta ornata lateralis Uta stansburiana stejnegeri Sceloporus clarkii clarkii Sceloporus magister magister PhrynoKoma solare Cnemidophorus melancstethus Constrictor constrictor imperator Hypsiglena ochrorynchus Masticophis flagellum frenatum Thamnophis eques Trimorphodon lambda Tantilla hobartsmitld Crotalus cinereous Kinostemon sonoriense Bufo punctatus Bufo alvarius Scaphiopus couchii 478 The University Science Bulletin AIVIPHIBIANS The amphibian collection makes no claim to completeness. Most of the time spent in Sonora was prior to the advent of smiimer rains, when much of the amphibian population was still in aestivation. On my return to the United States in August I obtained a few amphibians in Guaymas and at Noria. In the latter locality I failed to find Pternohyla fodiens Boulenger and Microhyla olivacea (Hallowell), species which Allen (1933) found to be plentiful. Bufo woodhousii Girard and Bufo marinus (Linnaeus) have been re- ported by Kellogg (1932). The record for Bufo compactilis Wieg- mann from this locality may be regarded as doubtful. Scaphiopus couchii Baird A single specimen (No. 1151) was captured late at night on dry sand near Noria. It appeared to have been attracted by my lan- tern light, and approached close to my camp. Four were taken near Guaymas on a flat during a light rain. Bufo alvarius Girard One specimen (No. 14) was taken at night at La Posa, ten miles northwest of Guaymas. Four typical, immature specimens (Nos. 1102-1105) were collected on the night of August 4, on a barren fiat four miles north of Guaymas. Bufo punctatus Baird and Girard One specimen (No. 132) was captured with the four above-men- tioned alvarius. Hyla arenicolor Cope A series of eleven specimens (Nos. 368-378) was obtained from the immediate vicinity of a spring, about two miles north of La Posa. Here there is a sudden break in the low range, and within the cleft is a small basin where numerous palm and fig trees grow luxuriantly. The frogs emerged at twilight from among the dead palm leaves, which hung suspended about the trunk of the palms, and approached the pools below the spring. Here they were cap- tured as they sat in the edge of the water or on the banks. All appear to be half grown. They are marked with very distinct rounded spots on a lighter, grayish ground color. Taylor: Fauna of Sonora 479 Rana pipiens Schreber Five recently transformed specimens (Nos. 379-383) were en- countered in the same pools mentioned above. A few tadpoles, not yet transformed, were seen in the pools. LIZARDS Phyllodactylus hornolepidurus Smith Phyllodnctylus homolepidnrus Smith, Univ. Kansas Sci. Bull., XXII, Apr. 15, 1935, pp. 121-125, pi. XXV, fig. 2a and text fig. 1, A. A series of twenty-six was collected at the following localities: twelve (including the type), five miles southwest of Hermosillo; eleven, from a point ten miles northwest of Guaymas; three, four miles southeast of Guaymas on the Empalme road. Smith {loc. cit.) has given a thorough account of these specimens. Coleonyx variegatus (Baird) Stenodactylus variegatus Baird, Proc. Acad. Nat. Sci., Phila., 1858, p. 254 (type descrip- tion ; type locality, Rio Grande and GUa Valleys). Coleonyx variegatus Smith, Trans. Kansas Acad. Sci., XXXVI, 1933, pp. 301-314. Thirty-eight specimens of Coleonyx variegatus (Baird) were col- lected in Sonora, and with one exception, in which the specimen was found hidden under a rock, they were discovered at night run- ning about over gravelly soil near the beach and in the mountains. These specimens agree with the characters noted by Smith {loc. cit.) as being diagnostic of this species. The preanal pores vary from six to nine, and the supranasals are invariably in contact medi- ally behind the rostral. The cloacal bones agree with those typical of variegatus save there is a suggestion of a groove or notch at the exposed end in certain specimens. In markings, a number of the specimens show a tendency for the transverse bars to break medially and a suggestion of a median dorsal line is often evident. The bars on the tail do not extend ventrally as is typical in many specimens of brevis. The breaking up of the juvenile color pattern seems to begin a little later in life than is typical, since only the largest specimens (snout to vent, 60 mm.) show this condition. The specimens of this species usually run with the tail lifted, often curved over the back. The bright light from my lantern tended to bewilder them, and they were caught at night with little difficulty. 480 The University Science Bulletin I obtained specimens from the following Sonoran localities: Nos. 123-124, 152-153, 176-177, five miles southwest of Her- mosillo, June 22 to 24; Nos. 260, 267, 293-295, 301, 327-333, 396, 436-437, 458, 461-462, 383, 385, La Posa, ten miles northwest of Guaymas, June 29 to July 16; 1142-1146, 1178-1182, near La Posa, Guaymas, August 5 to 7; 1183-1187, five miles southwest of Her- mosillo, August 8 and 9. Ctenosaura hemilopha (Cope) Ctevosaura hemilopha Smith, Univ. Kansas Sci. Bull., XXII, No. 6, Apr. 15, 1935, pp. 140-142, pi. XXIII, fig. 1. Two specimens were collected; one from a locality ten miles northwest of Guaymas is figured by Smith. The second specimen is from five miles southwest of Hermosillo. Smith has discussed these specimens {loc. cit.). Dipso-saurus dorsalis sonoriensis Allen Specimens were taken at Hermosillo and Guaymas as follows: Topotypes, Nos. 180, 180a, five miles southwest Hermosillo, June 25; Nos. 296, 341, 362a, 419, 463, and 494-497 (skeletons), between July 2 and 12 in the general region about Miramar and La Posa; 1128-1129, northwest of Guaymas, August 7; 1127-1128, 1148-1149, near Miramar, August 7 and 8. These specimens agree with the details of the type description. The two or three scale rows between the nostril and rostral (as occurs in the typical subspecies) is reduced in this form to a single series. This appears to be constant. The brown reticulation on the throat enclosing round or oval cream areas, the less distinct rows of dark spots forming caudal annuli, and the tendency to lose the ocellated spots on the neck and shoulders, are all characters that tend to separate these from the more northern forms. The labials are fewer and smaller. All the specimens come within the limits of variation assigned to the form by Allen. Crotaphytus collaris baileyi Stejneger One male specimen (No. 1162) was taken about sixty-one miles south of Nogales. The throat is lavender, becoming purplish pos- teriorly and covered with cream reticulations. The spots in the temporal region are a light chocolate brown; the neckbands are widely broken below; body with slate bands separated by only Taylor: Fauna of Sonora 481 slightly lighter interspaces, the whole covered with tiny light-cream spots; tail with indistinct brown bands. The hind leg brought for- ward, the tibiotarsal articulation reaches the shoulder, the longest toe to a point somewhat in advance of the eye. Ventral surface of limbs, abdomen and tail, cream-white. Snout to vent, 98 mm.; tail, 188 mm.; hind leg, 85 mm.; femoral pores, 16-19. Sauromaulus townsendi Dickerson Sauromaulus townsendi Schmidt, Bull. Amer. Mus. Xat. Hist., XLVI, Dec. 7, 1922. p. 643; Belding, West Amer. Scientist, III, 1S87, pp. 97-99. I collected three specimens of this species: one (No. 198) fifty- four miles southwest of Hermosillo; one (No. 222) at Miramar, three miles northwest of Guaymas; and one (No. 346) five miles north of Guaymas. All were discovered in rock crevices from which localities they were removed with considerable difficulty. These specimens have been identified with Dickerson's species on the basis of the character of the interauricular scales and the large size of the caudal scales. Schmidt {loc. cit.) has pointed out that the species is, in several respects, intermediate between &. obesus and S. hispidus but actually distinct from both. Measurements (Nos. 198 and 222, respectively [in millimeters]): Snout to vent, 126, 155; tail, 140, 122 (regenerated); head length, 28, 32; head width, 26, 32; body width, from edges of lateral folds, 54, 62; foreleg, 52, 61; hind leg, 78, 88. The number of scales in a single whorl about the tail at widest part, 64, 62; scales from gular fold to anus, 153, 143. Color. No. 198. Above, the specimen is yellowish-tan and brown, the darker color forming five, rather indefinite, broad bands across the body. The tail with five, unequal, broad, dark-colored bands and five lighter bands, the terminal band narrow. The dorsal sur- face is marked with small spots, those on the dark bands larger than those on the lighter interspaces. Below, the body is a dirty cream with some lavender marbling on the throat. The ventral surface of the thigh has brown, punctate spots. The edges of the gular fold and the preauricular lobules are cream. The head is brown above. The larger specimen (No. 222) is darker above than the preced- ing specimen. The general pattern of dark bands with lighter inter- spaces can be discerned with difficulty. There is only a slight con- trast between the light and dark bands on the tail. The sides of the body have numerous small brown spots. 31—6037 482 The University Science Bulletin Callisaurus inusitatus Dickerson Callisaurus inusitatus Dickerson, Bull. Amer. Mus. Nat. Hist., XLI, 1919, p. 465; Allen, Occ. Papers Mus. Zool. U. of Mich., No. 259, Apr. 3, 1933, pp. 7-8. The following specimens were collected: Nos. 51, 52, twenty- five miles south of Nogales, June 20; No. 60, fifty -three miles south of Nogales, June 20; Nos. 83, 84, near Noria, June 22; Nos. 131, 132, and 178, five miles southwest of Hermosillo, June 23; Nos. 186, 188, fifty-four miles southwest of Hermosillo, June 28; Nos. 207, 208, Miramar, near Guaymas, July 29; Nos. 283-286, 290, 339-340, 347, 363-364, 396, 399-407, 423, 452-459, 464, 465, 486, 487, 504-506, 521, La Posa, June 29 to July 10; Nos. 1111-1117, 1121-1126, 1150, 1188-1190, La Posa and Miramar, August 4 to 6. If Linsdale* is correct in his surmise that there is but a single species of Callisaurus on the peninsula of Lower California and the western United States, the form here considered would be Calli- saurus draconoides inusitatus. I lack sufficient comparative mate- rial to either concur in, or disprove, his conclusions, Holbrookia elegans thermophila (Barbour) Holbrookia elegans thermophila Smith, Univ. Kansas Sci. Bull. XXII, No. 8, Apr. 15, 1935, pp. 194-195, pi. XXVII, fig. 1; pi. XXVIII, fig. 4. (Also, pp. 191-193.) Sixty-eight specimens were collected at various points in Sonora, as follows: Fifty -three miles south of Nogales, two specimens; two, eight miles south of Magdalena; five near Noria; one, thirty miles south of Noria; three, five miles southwest of Hermosillo; five, fifty- four miles south of Hermosillo; fifty, ten miles northwest of Guay- mas. Smith (loc. cit.) has discussed this lot, and has given a series of measurements. Uta taylori Smith Uta taylori Smith, Univ. Kansas Sci. Bull., XXII, Apr. 15, 1935, pp. 158-166, pi. XXVI, fig. 3. Thirty-one specimens, including the type, were collected as fol- lows: Nos. 252-258, June 30; 280-284, July 1; 300, 304, July 2; 320, 321, 322, 323a on July 3; 335, July 4; 393, July 6; 474, 474a, July 12; 500, July 13; all from La Posa, ten miles northwest of Guaymas. Nos. 199-202, fifty-four miles southwest of Hermosillo, June 26; Nos. 448-450, from a locality twelve miles northwest of Guaymas, July 10. * Linsdale, Amphibians and Reptiles from Lower California. Univ. Cal. Publ. Zool., XXXVIII, No. 6, June 24, 1932, pp. 357-359. Taylor: Fauna of Sonora 483 Uta stansburiana stejnegeri Schmidt Uta stansburiana stejnegeri Smith, Univ. Kans. Sci. Bull., XXII, Apr. 15, 1935, pp. 166-167. A series of seven (Nos. 501-503, 514-517a) from near Empalme. See Smith {loc. cit) for comments on these specimens. Uta ornata lateralis (Boulenger) Uta ornata lateralis Smith, Univ. Kansas Sci. Bull., XXII, April 15, 1935, p. 179. Thirty-nine specimens were taken at the following localities: Eight miles south of Magdalena; thirty miles south of Noria; five miles southwest of Hermosillo; La Posa, ten miles northwest of Guaymas. Smith {loc. cit.) discusses this series. Sceloporus clarkii clarkii Baird and Girard Sceloporus clarkii Baird and Girard, Proc. Acad. Nat. Sci., Philadelphia, VI, Aug., 1852, p. 127. (Type de.scription ; type locality, "Province of Sonora." John H. Clark, collector.) (Not improbably from southern Arizona or southern New Mexico.) This species was found to be present in most of the localities where I collected. In this region the species was encountered, al- most without exception, in trees. In New Mexico, in certain lo- calities, it becomes adapted to a rock habitat along deep arroyos. In eastern Arizona, magister magister also occurs and is the more terrestrial form, while clarkii clarkii is chiefly arboreal in habit. It may be remarked that magister magister has a range from Nevada to Texas and southern Sonora — a range that includes the smaller range of clarkii clarkii. Nowhere is there any evidence of intergradation. Certain scale differences are constant and other differentiating characters of equal specific importance, suffice to en- able each to maintain complete distinction from the other. The following specimens were taken: Nos. 47-48, twenty-five miles south of Nogales, June 19 ; 54-57, about fifty-three miles south of Nogales, June 19; 65-68, Noria, June 20 (skeletonized); 74, thirty miles south of Noria, June 20; 105-106, thirty miles south of Noria, June 22 (skeletons); 116, 122, five miles southwest of Her- mosillo, June 24; 182, fifty miles south of Hermosillo, June 24; 230, 306, 316-319, 336, 367, 386-391, 325-429, 429a, 429b, in the vicinity of La Posa, ten miles northwest of Guaymas, June 29, July 9 ; 524, Miramar, July 9; 1164, fifty miles south of Nogales, August 10. 484 The University Science Bulletin Sceloporus magister magister Hallowell Sceloporiis magister Hallowell, Proc. Acad. Nat. Sci. Phila., VII, 1854, p. 93 (type descrip- tion ; type locality, Yuma, Ariz.). Eight specimens were collected in Sonora, as follows: Nos. 81-82, near Noria, June 20, in a hole in a tree; 292, yg.. La Posa, ten miles northwest of Guaymas, July 1; 408, July 7, La Posa; 420, La Posa, July 9 (skeleton) ; 466 and 466a, San Carlos Bay, July 12; 1152, La Posa, August 8. This speciesf was usually encountered feeding in trees or on cactus. It appears to have burrows or at least takes refuge in burrows about the roots of plants. The stomachs were filled chiefly with flowers at this season. One specimen (No. 420, skeletonized) contained nineteen well- developed eggs. The head and neck of the specimen was distinctly reddish in life. Phrynosoma solare Gray Specimens of this species were collected as follows: No. 179, southwest of Hermosillo, enroute to Guaymas, June 25; 259, ten miles northwest of Guaymas, July 8; 512, on the beach road near Empalme, July 13; 1110, in a street in Guaymas; 1156-1157, in the highway near Santa Ana, August 10. All of these specimens, save one, were encountered in the morn- ings, along roads. A single one was taken under a low shrub, in partial shade, about 2:30 in the afternoon. Compared with Arizona specimens from Tucson, the Sonoran specimens exhibit the same general color patterns. The horns sur- rounding the back of the head appear to be slightly longer in the males and the scales growing from the edge of the lateral skinfold (below the lateral spine series) are somewhat larger. All the speci- mens have the belly whitish or rusty white, with numerous, punctate, black spots. Heloderma suspectum Cope Three specimens of this species were collected at night near La Posa. Two (Nos. 263, 264) were captured in the sand near the point of meeting of the sandy beach and the shrub. (No. 264 has been skeletonized.) No. 205 was found on the top of the low moun- tain rising from the edge of the sea. This last specimen was captured and placed in a sugar sack. On picking up my "snake stick" which had fallen near the sack, the t More complete data on this form and S. clarkii clarkii will appear in a work by Hobart M. Smith treating of Mexican rnd American Scelopori. Taylor: Fauna of Sonora 485 Heloderma made a quick lunge, and seized the third finger of my right hand, sinking the teeth to the bone. With a quick jerk, I tore my finger from the animal's mouth, thus widely opening the wounds. After sucking the wounds, the fingers were wrapped up and col- lecting continued. There was no ill effect, other than that which any wound of like extent would cause. No effects that could be at- tributed to venom could be observed. Cnemidophorus melanostethus Cope Cnemidophorus melanostethus Cope, Proc. Acad. Nat. Sci., Philadelphia, 1863, p. 104 (type description; type locality "Region of the Colorado of California," H. B. Mollhausen, collector). Cnemidophorus tessellatus aethiops Cope, Ann. Rep. U. S. Nat. Mus., 1898 (1900), p. 582 (type description; type locality, Hermosillo, Son., Mex. Jenkins and Evermann, collectors). Cnemidophorus tessellatus tessellatus Allen, Occ. Papers Mus. Zool. U. of Mich., Xo. 259, 1933, p. 10 (Puerto, Hermosillo, and Guaytnas, Sonora). I observed this species in 1934 at a number of localities along the highway between Nogales and Guaymas, in fact, it was one of the most common of Sonoran lizards. The following are in the collection: Nos. 91-95, 96, 96a, 99-100, June 21, near Noria; 114, thirty miles south of Noria, June 22; 133-136, 161-165, five miles southwest of Hermosillo, June 23-25; 220, 231-232, 232a-232b, 238, 270-271, 297, 313-315, near La Posa, ten miles northwest of Guaymas, June 28 to July 5; 358-362, near Miramar, three miles northwest of Guay- mas, July 5; 404, 475-476a, 507-510a, July 14, near La Posa; 1118- 1119, 1130-1134a, near Guaymas, August 5-7, 1934. Several other specimens have been skeletonized. Cnemidophorus burti sp. nov. (Plate XLII, fig.. 2.) Holotype. No. 269, collected near La Posa, ten miles northwest of Guaymas, Sonora, July 4, 1934; E. H. Taylor, collector. Para- types Nos. 239, 268, 311 (skeleton), 312, 392, 442, 443, all from the region about La Posa, collected from June 20 to July 10. Diagnosis. Related to C. perplexus, but with a tendency toward a reversal of the typical color pattern. The young are five-lined; brownish or tan dots on a dark-brown background between much widened, bright, cream-colored, lateral lines, and between the dorso- lateral and the broad, lavender, median lines. Most of the older specimens lose practically all trace of the dots and the pair of lat- eral lines on each side are strongly intensified, while the median becomes dim lavender to reddish in color. The ground color be- comes dark or light reddish-brown. 486 The University Science Bulletin Description of the type. Rostral typical, the part visible above longer than the nasal suture, but shorter than the frontonasal; frontal single, once and one fourth as long as broad; two fronto- parietals, their greatest combined width equal to two thirds the combined width of the parietal series; three parietals, followed by a group of scales, the one bordering the outer lower edge of the outer parietals largest; four supraoculars, the anterior triangular and of about the same area as the last, the first three touching the frontal; last three supraoculars separated from superciliary series by a single row of small granular scales, and the third supraocular partially and the fourth wholly separated from the frontoparietals. Six-seven superciliary scales; eyelid with a row of seven small plates higher than long; enlarged upper labials 5-5; enlarged lower labials 7-8, the anterior much reduced. Mental rounded, followed by a single undivided postmental; five pairs of chinshields (sublabials), only the first pair in contact, the last separated from the mouth angle by two pairs of moderately enlarged scales only partially in contact with the labials; postnasal large, much higher than wide, followed by a much larger loreal, and a very small scale segmented from the lower posterior corner of the loreal; two preoculars, each keeled along the upper, inner border; subocular elongate, with a low keel, followed by two post- suboculars; a series of enlarged temporals, continuous with the superciliaries, flanked above and below by somewhat smaller scales. Auricular opening large, D-shaped; enlarged anterior gular series extending across the ventral surface, separated from the first gular fold by fourteen rows of subequal scales; three rows of smaller scales in first fold ; five rows of scales between the anterior and pos- terior folds, becoming much enlarged posteriorly; eleven rows of granular scales in the posterior fold. Venter with six complete series and two outer incomplete series of much widened ventral scutes, the two median somewhat narrower than others; about thirty-six transverse rows from gular fold granules to the femoral pores. Upper arm with five, lower with three rows of enlarged scales (brachials and antebrachials) on front of arm; the postantebrachials distinctly enlarged; femoral pores, 18-19; tail elongate, tapering; scale rows around body, 98 (not in- cluding enlarged ventrals). Color in life. Above blackish-brown with five longitudinal light stripes from head to groin or to base of tail; the dorsolateral and median lines extend onto the tail some distance ; the light stripes are Taylor: Fauna of Sonora 487 distinctly more than half the width of the intervening dark stripe; the median light line forks on the neck and is lilac to lavender in color. The lateral stripes are cream anteriorly, becoming light lav- ender posteriorly. There is no trace of spots. Ventral surfaces cream, immaculate. Top of head uniform olive. Measurements (in mm.) of holotype and paratypes of Cnemidophorus burti, sp. nov. Number Sex Snout to vent Tail Width of head, temporal Length of head, rostral to back of parietals Width of head, across eyelids Snout to ear Snout to foreleg Axilla to groin Foreleg Hind leg 268 442 392 312 9 269 443 yg. 85 85 85 80 78 68 240 233 235 197 13.5 13.2 14 12 13 9 17 19.5 19 17 19 15 9 9 9 7.5 9 7 19 19.5 19.8 18 18 16 32 35 31 28 29.5 22 39 38 38 42 38 34 30 29 30 30 27 26 61 58 56 58 55 53 239 yg- 64 183 10 14 6.5 14.5 24 30 23.5 45 Variation. In color the youngest specimen (No. 239) shows a well-defined series of spots on the darker background between the light lines; these have practically disappeared in No. 443, which is a few millimeters larger. Otherwise, the color pattern is prac- tically unvaried in the remainder of the series, save that the ground color tends towards a red-brown in most of the specimens. Scale rows around the body vary between 93 and 99, the average being about 97. Remarks. Burt (Bull. U. S. Nat. Mus., No. 154, 1931, p. 132) mentions a specimen of this form from near Guaymas, Sonora, under the name of C. sexlineatus perplexus, and Allen (Occ. Papers Mus. Zool., U. of Mich., No. 259, 1933, p. 10), likewise mentions one of this species from Guaymas. The coloration is, as suggested by Burt, strongly reminiscent of that of hyperythrus. The species is named for Dr. Charles Burt, who has made the genus Cnemidoph- orus his particular field of study. 488 The University Science Bulletin Criemidophorus perplexus Baird and Girard Cnemidophorus verplexus Baird and Girard, Proc. Acad. Nat. Sci., Philadelphia, 1852, p. 128 (type description; type locality — Valley of the Rio San Pedro. Also collected by General Churchill, on the Rio Grande, west of San Antonio, Te.xas, and by Dr. William Gambel on his last journey to California). The proper identity of numerous populations of Cnemidophorus in the southwestern part of the United States is, I believe, still in question. Burt (1931, Bull. U. S. Nat. Mus., No. 154) has the identification problem solved by placing most of the named forms in the synonymy of sexlineatus perplexus and tessellatus tessellatus. Since the modern tendency in systematic taxonomy is the recog- nition by name of variant populations which cover a considerable range, it seems likely that certain of the names now in synonymy will of necessity be resurrected for such variant populations whose differential characters are not wholly due to differences in age or sex. The species here considered as Cnemidophorus perplexus differs from western Texas (type locality) specimens in several characters, not the least important of which is a much larger number of scale rows around the middle of the body. A series of sixteen specimens from western Texas and southeastern New Mexico have a range from 64 to 72 scale rows around the middle of the body, the average being 68. The series from Hermosillo, in the central northern part of Sonora (Nos. 156-160, June 23-25, 1934; E. H. Taylor, collector), vary between 89 and 95, the average being about 92. The femoral pores are 16-19, the average being 18; this number appears seven times (in 12 femora counted), 19 twice, 17 twice, 16 once; a speci- men (No. 189) taken fifty-four miles southwest of Hermosillo has 95 scale rows. The series shows age transitions from a six-lined form to one with a series of twelve rows or rounded, light spots, largest on the sides, less distinct and smaller on the median dorsal region, those on the nuchal region obsolete. The specimen (No. 189) taken fifty- four miles southwest of Hermosillo has the two median lines tend- ing toward a fusion in the middle of the back; they are separate posteriorly and are obscured by the brownish color in the neck region. It is probable that this form should be recognized as a subspecies • of perplexus. It seems that the young lack the bluish ventral and caudal coloration and there is an average difference in the adult color pattern. However, in general, these follow the expected indi- vidual evolution of the color pattern for perplexus (typical) . Taylor: Fauna of Sonora 489 SNAKES Lichanura roseofusca gracia Klauber Lichanura roseofusca gracia Klauber, Trans. San Diego Soc. Nat. Hist., VI, No. 20, Apr. 30, 1931, pp. 305-318, pi. 21, fig. 1; and Copeia, No. 4, Dec. 27, 1933, pp. 214-215. The report of a form of Lichanura (Klauber, 1933) at Guaymas, Sonora, anticipates the capture of my specimen from near Her- mosillo, which lies about seventy-five miles north of Guaymas. The specimen (No. 129) was captured June 29, 1934, just after daybreak in a mass of boulders five miles southwest of Hermosillo. It is a pregnant female with the embryos probably two thirds de- veloped. No fear was shown at my approach and when picked up the snake remained quite docile. The embryos were removed when it was preserved. These are Nos. 166-169. They show the typical striped color pattern of the mother. The measurements (in millimeters) and scale counts are as fol- lows: Snout to vent, 546; tail, 62 (injured); ventrals, 223; sub- caudals, 40; scale rows, neck 40, about body 43,' in front of anus 25. The young measure about 160 to 172 in total length, the tails measuring about 25 mm. Three of the young are males, with the hemipenes still completely extruded; one is a female; the latter has slightly shorter tail than the males. The following table shows the scale counts of these specimens: No. Sex 166 $ 167 ^ 168 $ 169 9 Constrictor constrictor imperator (Daudin) Constrictor constrictor imperator Allen, Occ. Papers. Mus. Zool., U. of Mich., No. 259, Apr. 3, 1933, p. 11 (Hermosillo). Three specimens of this boa were taken in the neighborhood of La Posa. One was skeletonized, the other two were brought back to Lawrence, Kan., alive. All are typical. One was found in the mouth of a small cave, in which was ob- served on several occasions the track of what appeared to be a very large specimen of this species. One specimen, a female, was cap- tured in a tree cactus about five feet from the ground. A third was captured at night crawling along a dry arroyo. Ventrals Subcaudals 223 49 220 47 223 48 227 48 490 The University Science Bulletin The capture of this snake by Allen (1933) at Hermosillo suggests the possibility of its being eventually discovered in some of the mountain ranges of southern and southwestern Arizona. Masticophis flagellum frenatum Stejneger Masticophis flagellum frmiatus Ortenburger, Mem. Univ. Mich. Mus., I, 1928, pp. 112-125, pis. XX, XXI. A specimen collected in a low tree on the edge of the beach at La Posa is referred to frenatus. This specimen (No. 395) is uni- form brown-olive, each scale showing a lighter area on the anterior median part. This lighter area becomes more distinct posteriorly. There is some evidence of spots on the labials, and the loreal and posterior nasal seem to be more yellow than the scales above and below. The anterior ventrals have a double row of median spots which are more evident anteriorly. A second specimen (No. 513), captured near Empalme, is slightly darker than the preceding, but there is no trace of the dark bands. The dark, labial spots are dimly visible, while the anterior ventral and mental spots are quite distinct. The loreal line is dimly visible. A third specimen (No. 1460) was collected near Santa Ana in northern Sonora. This young specimen has the typical coloration, but even in the young the transverse markings are not pronounced. The head markings are, however, typical of frenatus. The anterior ventral spotting is irregular, the spots being in about four rows. A large specimen (No. 1137) of the "red" phase of this species was captured near Guaymas, and brought alive to Kansas Uni- versity. At first the specimen was shy and timid, attempting to es- cape as one approached the cage, striking when a hand was placed in the cage. Later it became very tame, pennitting itself to be force-fed on beef without resistance. Measurements and Scale Counts of Masticophis flagellum frenatum Stejneger Sub- Upper Po.'^t- Total Tail No. Ventrals caudals labials oculars Scale formula length length 513 203 97 8-8 2-2 22 , 17, ,15, ,13, , 12 1810 450 395 201 110 8-8 2-2 22, 17, 17, 13, 12 1292 246 1160 196 99 8-8 2-2 21, 17, 17, 13, 12 935 242 1137 197 106 8-8 2-2 1760 415 Masticophis pdceus (Cope) Masticophis piceus Ortenburger, Mem. Univ. Mich. Mus. Zool., I, 1928, pp. 112-125, pis. XX, XXI; Allen, Occ. Papers Mus. Zool. Univ. Mich., No. 259, Apr. 3, 1933, p. 11. (Hermosillo). Two specimens were captured in Sonora. The first (No. 113) was collected thirty miles south of Noria near the highway. It was dis- Taylor: Fauna of Sonora 491 covered extending its head from a woodpecker hole, in a tree, about thirty feet from the ground, and was shot. This specimen is of a deep, blue-black color, save that toward the tip of the tail the color becomes brownish-black. On the ventral surface on the posterior half of the body there is an infusion of brown, becoming lighter pos- teriorly. On the last fifth of the body there are a few j'ellow flecks. In the anal region these form larger yellow areas. The inner edges of the subcaudals are light colored. Two other specimens seen es- caped into holes in a hollow branch of the same tree. No. 379. The anterior half of the body is black. About the middle of body there are three black-spotted, reddish bands, while the latter third of the body is more or less reddish. The tail is uniformly reddish-brown. The specimen was captured in the heat of the day under a flat rock, near La Posa. These specimens seem to be common in the trees along small ravines. At least eight specimens seen escaped into holes in the boles or branches. Scale Counts and Measurements (in mm.) of Masticophis piceus (Cope) Vev- Sub- Upper Lower Pre- Total Tail No. Sex trals Anal caudals labials labials oculars Scale formula length length 113 $ 205 2 96 8-8 9-9 2-2 19,17,16,13,12 1460* 383 479 $ 200 2 113 8-9 11-12 2-2 21, 17, 17, 12, 12 1468* 395 • Tip of tail missing. Masticophis semilineatus (Cope) Masticophis semilineatus Ortenburger, Mem. Univ. Mich. Mus., I, 1928, pp. 48-57, pis. XI, XII, XIII, and text figs. 6, 7, 8; Allen, Occ. Papers Mus. Zool. Univ. Mich., No. 259, April 3, 1933, p. 11 (Hermosillo). Three specimens of this species were taken. No. 155 was cap- tured five miles southwest of Hermosillo at night in a bush; Nos. 234 and 325 were taken in the morning, at La Posa. No. 425, taken in a deep canyon in the mountains near La Posa, has been skeletonized. These specimens agree well with specimens described and figured by Ortenburger {loc. cit.) and the scale counts are well within the known range of variation. Scale Counts and Measurements (in mm.) of Masticophis semilineatus (Cope) No. Sex Ventrals Sub- caudals Upper labials Lower labials Scale rows Total length Tail length 155t ■f . . . . . . 8-8 10-10 20,17, 17, ? . • • 325 9 203 135 8-8 9-9 20, 17, 16, 13 1127 354 234* $ 202 88 + 8-8 8-9 20, 17, 17, 13 1084 265-1- • Tip of tail missmg. t Posterior part of body wanting. 492 The University Science Bulletin Salvadora grahamiae hexalepis (Cope) Salvadora grahamiae Baird and Girard, Cat. N. Amer. Rept., pt. 1, 1853, p. 104 (Sonora, Mex. ; Col. J. D. Graham, collector). A specimen of S. g. hexalepis (No. 1163) was found dead under a rock at a point on the highway fifty miles south of Nogales. It consisted of a skeleton with dried skin sufficient to identify the species. Phyllorhynchus decurtatus decurtatus (Cope) Phj/llorhynchus decurtatus decurtatus Linsdale, (part) Copeia, No. 4, 1933, p. 222; Klauber, Bull. Zool. Soo. San Diego, No. 12, Sept. 12, 1935, pp. 5-9. Three specimens of Phyllorhynchus were collected in southern Sonora. No. 457 2 was found at La Posa running rather rapidly near some shrubs, about twenty-five meters from the sea, the night of July 10. No. 483 $ was discovered at La Posa by following a snake's trail in loose dry sand on the beach, the night of July 12. A third specimen, No. 1100 $ , was found on a barren fiat near the sea about four miles north of Guaymas, the night of August 4. Data and Measurements of Phyllorhynchus decurtatus decurtatus (Cope) No. Sex : Length Tail Ventrals Sui beau dais Scale rows Upper labials Lower labials 457 9 284 27 165 25 24-19-19 7-6 9-9 483 s 273 49 151 34 23-19-17 6-7 9-9 1100 $ 296 44 152 31 20-19-16 6-6 9-9 No. Oculars Loreals Dorsal spots body Dorsal spots tail 457 8-8 3 -3 42 7 483 8-8 2- -2 25 5 1100 7-6 1- -1 28 5 The first specimen (No. 457) is a female, and the scales are smooth save for a very faint suggestion of keels along the posterior half of the body. No. 483 has the scales strongly keeled on the nine dorsal scale rows, the keels beginning about one centimeter back of the head; No. 1100 has the scales of the nine median dorsal rows even more strongly keeled than the preceding. These three specimens were taken in the same general locality, all less than five miles apart, and two of these, a male and a female, within a few yards of each other. These are, apparently, specimens of the same species, but there are strong differences in markings, perhaps due to sexual dimorphism. Linsdale believes a specimen taken at Caborca, Sonora, having faint keels, should be referred to browni, but as a subspecies of decurtatus. Taylor: Fauna of Sonora 493 The specimens listed here have been studied by L. M. Klauber in his recent review of the genus Phyllorhynchus [loc. cit.). I have followed him in the disposition of the specimens. They present the following characteristics: Snout with a curved band rising on fifth and sixth labials, passing through eye and across the anterior part of the frontal to the opposite side of head ; a second band crosses just behind the parietals and the ends run back varying distances on the side of the neck, on either side of the large nuchal spot; the area between these two bands on the head is variously spotted, or the two bands may be joined by confluent spots. A light spot on the frontal is enclosed wholly or partially by dark color. The first two dorsal spots are confluent in the males but divided in the female. The first few dorsal spots are deep, black-brown. The dorsal spots on the males have darker borders; in the female, this darker border is scarcely evident. In the males the dorsal spots total 30, and 33; in the female, 50. There is an irregular series of lateral spots in the males; in the female these are more numerous and less distinctly outlined. Pituophis sayi affinis (Hallowell) Pituophis sayi aiftnis Stull, Occ. Papers Mus. Zool. Univ. Mich., No. 250, Oct. 12, 1932, p. 4. A single specimen (No. 80) was collected on the morning of June 20 in the shade of a small tree in the arid region near Noria. It remained motionless until I approached and grasped it. It presents the following characters: Four prefrontals; rostral entering between the internasals nearly half their length; loreal elongate; one pre-, three postoculars; 8-8 upper labials, the fourth entering orbit; 12-13 lower labials, six touching the first chinshield; scale rows, 29, 31, 33, 33, 23; ventrals, 235; anal, single; subcaudals, 57; blotches on body, 46; blotches on tail, 12; anterior two thirds of body has the lateral spots more or less confluent with the dorsal blotches and the yellow cream interspaces form small spots occupy- ing 9 to 12 scales on the sides; the light scales have a brown keel. The head is yellowish posteriorly, with small black flecks; a curved band slightly anterior to eyes; a diagonal line from post- oculars to the last labial; labial sutures brown; a series of brown spots on the ventrals and subcaudals, somewhat confluent under tail. 494 The University Science Bulletin Ficimia desertorum Taylor (Plate XLIII, fig. 1.) Ficimia desertorum Taylor, Proc. Biol. Soc. Wash., Vol. 49, May 1, 1936, pp. 51-52. The type specimen was discovered near a spring, about one and a half miles northeast of La Posa, ten miles northwest of Guaymas. It had been injured, and ants had destroyed a few scales. This spring is a few hundred yards farther east than the opening of the canyon in which was a spring mentioned elsewhere. Hypsiglena orchrorhynchus Cope Hypsiglena ochrorhyncha Cragin, Bull. Washburn Lab. Nat. Hist., Sept., 1884, No. 1, pp. 6-8 (Guaymas) ; Allen, Occ. Papers Mus. Zool. Univ. of Michigan, No. 259, Apr. 3, 1933, p. 12 (Herinosillo). The Cragin (1884) record of one specimen from Guaymas is the earliest known for Sonora. It was collected by Prof. J. T. Lovewell. Allen obtained a specimen at Hermosillo. I collected three specimens. No. 120 was taken in the boulder field five miles southwest of Hermosillo. It was found at dusk under the edge of my car step, June 22. No. 266 was caught in camp at La Posa, ten miles northwest of Guaymas, June 30; and No. 281 was discovered under a rock, under an overhanging cliff, near San Carlos Bay, two miles west of La Posa, July 1. The ground color of the three specimens varies from an ash-gray in No. 281 to a brownish-gray in No. 120, and to a lavender-gray in No. 266. The characteristic neck- and head-marking consists of a dark-brown spot which fails to reach the ventrals by three scale rows. From the lower anterior edges, branches, on each side, run across the temples through the eyes to the rostral, but fail to meet anteriorly, widest on the temples; dorsally the blotch has a median forward projection which reaches the parietals. The whole blotch is outlined by an indistinct cream line which becomes a very distinct white on the sides. It follows the upper edges of the anterior labials, passes diagonally across the posterior labials and the angle of the mouth and may be continued back along the neck a short distance. In two specimens the neck band is broken unequally in twain by a cream-white line running longitudinally. The dorsal markings on the body are typical in all. The three specimens agree in the following characters: The nostril is between two nasals; a loreal present; two preoculars, the upper very large, the lower very small and below the anterior part of the eye. Two postoculars, the upper largest. Temporals, 1+2; Taylor: Fauna of Sonora 495 eight upper, nine lower labials (No. 281 has seven upper labials on right side, due to a fusion of the second and third) ; five labials touching the first chinshields; anal divided. Measurements (in mm.) and scale counts of Hypsiglena ochrorhynchus (Cope) No. Sex Total length Tail Ventrals Subcaudals 120 S 325 74 178 60 266 9 339 69 177 54 281 S 226 73 181 63 Thamnophis eques (Reuss) Thamnophia eques Ruthven, Bull. U. S. Nat. Mus., No. 61, 1908, pp. 158-164. Eight specimens were taken, all in the general neighborhood of La Posa, which appear to differ but little from highland specimens from Mexico and the United States. Ruthven (loc. cit.) has suggested that the species is absent from the coastal plain. These specimens demonstrate its presence at sea level, at least in the region about Guaymas. Cragin's (1884) record of Eutaenia cyrtopsis* from Guaymas has been referred to this species. Trimorphodon lambda Cope Trimorphodon lambda Cope, Proc. Amer. Phil. See, 1885, pp. 286-287 (type description; type locality, Gua\'mas, Sonora. Type, No. 13487, U. S. N. M. H. F. Emerich, collector); Cope, Ann. Rep. U. S. Nat. Mus., 1898 (1900), p. 1104. The retention of Cope's Sonoran species of Trimorphodon seems warranted despite the rather striking resemblance of the color pat- tern to certain Mexican and American forms. As yet, no series of sufiicient size is available to properly evaluate the various characters on which most of the species of this genus have been established. Klauber (1928), in his review of Trimorphodon in Baja Cali- fornia, California, and Arizona, recognizes the Arizona specimens from the Huachuca Mountains region as belonging to the species T. lyrophanes, and makes no effort to place lambda in the synonymy of that form. I have no lyrophanes available for making a critical comparison. I obtained three specimens (Nos. 265, 366, and 341) in the general vicinity of Guaymas. The first was captured near La Posa crawling on gravelly ground at sea level, near my tent about midnight. It stopped and remained quiet, allowing itself to be picked up without struggling, and without endeavoring to bite. No. 336 was found at night in similar terrain near the beach. A third was captured in a * I have endeavored to find this specimen in the collection of Washburn College at Topeka, Kan., but it appears to be lost. 496 The University Science Bulletin niche in a low rock cliff near the sea about ten o'clock in the morn- ing. The specimen was tightly coiled and was picked up in this condition, the specimen remaining coiled when captured, making no attempt at resistance. The last mentioned has been skeletonized. My specimens Nos. 265 and 336 present the following characters: sex, $ , $ ; scale counts at various points on body, 28, 19, 22, 22, 22, 16, 16; 29, 19, 21, 22, 22, 17, 15; ventrals, 232, 222; subcaudals, 83, 87; anals, 2, 2; supralabials, 9-9, 9-9; infralabials, 12-13, 11-12; preoculars, 3-3, 3-3; postoculars 3-3, 3-3; temporals, 3, 3, 4, 5; 3, 3, 3, 4; loreals, 2-2, 2-2; total length, 721 mm., 788 mm.; tail, 132 mm., 155 mm.; head length to jaw angle, 20, 25; head width, 14, 15; spots on body, 29, 31 ; spots on tail, 17, 18; spots separated by scales, 3%-4, 31/2-4; labials enter eye, 4-5, 4-5. The diameter of eye equals the distance of eye to nostril, in the first, and very slightly less in the second; the frontal is longer than its distance from the end of the snout; the prefrontals are as long as wide; frontal longer than parietal width, shorter than parietal length ; preocular is not in contact with frontal in either. The type has 234 ventrals; anal divided; subcaudals, 83. It agrees in the remainder of the characters so far as stated save that there are three loreals. This condition is probably anomalous, and occurs occasionally in several other species. The body markings of lambda consist of a series of saddlelike brownish blotches, with a whitish to a light gray transverse center, and bordered completely with the same color. The blotches are about four scale rows wide dorsally, narrowing to a single scale width laterally; they reach to the outer scale row. Between these blotches are other brownish-gray blotches which are about two scale rows wide dorsally, and five scale rows wide laterally; a more or less regular series of distinct, brownish spots on the outer edge of every second or third ventral, which may extend on the adjoining scales; rarely are these spots confluent with the dark dorsal blotches; ventrals with other dim brownish flecks. The head markings con- sist of a series of chevrons, the one across the snout brownish, fol- lowed by a narrow grayish band on the anterior part of the pre- frontals; this is followed by a broader brown band which passes through eye diagonally and terminates on the upper part of the fifth and sixth labials; next to this appears an irregular grayish chevron, followed by another which is broad, irregular, brownish, much lighter laterally, showing two darker spots on the supraoculars. A white dot present on the posterior part of the frontal which is Taylor : Fauna of Sonora 497 confluent with the gray chevron which follows. This chevron forms the light anterior border for the first dorsal blotch, which is grayish- brown with a circular gray medium dorsal spot. In the smaller specimen (No. 265) the markings between the blotches themselves are a little darker and the ventral flecks fewer than in the larger (described) specimen. Tantilla hobarts7?iithi Taylor Taiitilla huhnrtsmitlii Taylor. Tnin>. Kciii. Acad. Sci. XXXIX, 193(i, pp. :^40-84-2, fiir. 2. Only the type was captured. This was taken at a point about two miles north of La Posa, back of the first low range on the bank of a dry stream bed at night. The specimen was running rapidly over gravelly soil under Inrush. Crotalus cinereous Le Conte Caudisona ntrox sonoraciisis, Kennicutt, Proc. Ai-ad. Nat. Sci. Phila., Aug., 1801, pp. 200-207. This species was especially abundant in the southern part of the piovince. Eleven specimens were captured in the neighborhood of Miramar and La Posa, and several which were seen escaped. Sev- eral deca])itated and decaudated specimens w^ere seen, and a few specimens killed by autos were observed. The specimens in this region appear to be distinctly dwarfed as comjiared with the Texas specimens, and may warrant a revival of Kennicott's "sonoraensis." Most of the specimens were captured on the gravelly flats near the sea, and in the sand piled up by the waves where the sandy beach and shrub meet. This area harbors many rodents, insuring a constant food supply. One specimen was captured at Hermosillo. These specimens have been studied by Mv. Howard K. Gloyd. Crotalus tnolossus ntolossu^i Baird and Girard Crotalus molossus Allen (Gloyd), Occ. Papers Mus. Zool. Univ. of Mich., No. 259, April 3, 1933, pp. 13-14. Four specimens of this form were collected, one five miles south- west of Hermosillo, crawling at night among the boulders, and three near La Posa, ten miles northwest of Guaymas. One, an old, ema- ciated male, was taken late at night in a small cave in the mountains near La Posa. One was captured at 3:00 o'clock in the afternoon crawling along the west side of a cliff in a narrow shadow. The colors were very bright, as if it had recently shed. It did not sound 32—6037 498 The University Science Bulletin its rattles on my approach or even while being captured. A fourth was found early in the morning in another small cave near La Posa. This specimen was skeletonized. Data on these specimens have been taken by Mr. Howard K. Gloyd. Crotalus tigris Kennicott Crotulus tigris Allen (Gloyd), Dec. Papers Miis. Zoir iihylogenetic relationshijis. Bull. U. S. Nat. Mus., No. 54, pp. 1-2S6. C)itmidopliorus scxlincatus pcrplcxus (Hermosillo, Nogales, Guaymas, San Pedro Bay) ; Cnemidophorus tessellatus tessellalus (Guaymas, Hermosillo, San Pedro Baj-, Tepoca Bay, Tiburon Island). Cope, E. D. 1863. Description of new American Squamata in the museum of the Smithsonian Institution. Proc. Acad. Nat. Sci. Phila., 1863, pp. 100-106. 1900. Crocodilians, lizards and snakes of North America. Ann. Rept. U. S. Nat. Mus., 1898 (1900) pp. 151-1294. Numerous plates and figures. Lists a few Sonoran specimens from tlio collections of the U. S. National Museum. Cragin, F. W. 1884. Notes on some southwestern reptiles in the cabinet of Washburn College. Bull. Washburn Lab. Nat. Hist., I, Sept., 1884, No. 1, pp. 6-8. Sceloporus clarkii Baird and Girard (Sonora) ; Uta gulans sp. no\'. [=17/0 orimta lateralis, fide Van Denburgh. 1922 (Guaj'mas)]; Eutaenia cyrlopsis Kennicott [z= Thamnuphia eques (Reuss) (Guaymas)]; Elaps euryxanthiis Kennicott [=z Micruroides euryxanthus (Kennicott)] (Sonora). DiCKERSON, M. C. 1919. Diagnosis of twenty-three new species and a new genus of lizards from Lower California. Bull. Amer. Mus. Nat. Hist., XLI, pp. 461-477. Kellogg, Remington. 1932. Mexican tailless amphibians in the L^nited States National Museum. Bull. U. S. Nat. Mus., No. 160, 1932, pp. 1-224. Scaphi- opus hammondi multiplicatus (Cope) (Nogales, Sonora) ; Biifo alvarius 500 The University Science Bvlletix Girard (Guadalupe Canon) ; Bujo compactilis Wiegraann (Sonora [doubt- ful J); Bujo 7narimis (Linnaeus) (Camoa) ; Bujo icoodhousii Girard (Sonora) ; Hyla arenicolor Cope (Nogales, San Jose de Giiaymas) ; Bana pipiens Schreber (San Pedro ri\er, Santa Cruz river; in Sonora, near Lochiel, Pima county, Arizona). Kennicott, Eobt. 1861. On throe new forms of rattle.^nakes. Proc. Acad. Xat. Sci. Phila., 1861, pp. 106-108. Kl.auber, L. M. 1928. The Trimorphodon (lyre snake) of California, with note.s on the .species of the adjacent areas. Trang. San Diego Soc. Nat. Hist., V, No. 11, Ajuil 28. 1928. pp. 183-194, pis. 22-23. 1931. A new subspecies of the California boa with notes on the genus Lichanura. Trans. San Diego Soc. Nat. Hist., VI, No. 20, April 30, 1931, pp. 305-318, pi. 21, fig. 1. 1931. Crotdlus tigris and Crotaliis cnyo. two little known rattlesnakes of the southwest. Trans. San Diego Soc. Nat. Hist., VI, No. 24, pp. 353-370, pi. 23. Reports Crotalus tigris from Caballo, near Guaymas. — 1933. Notes on Lichanura. Copeia, 1933, No. 4, Dec. 27. 1933, pp. 214- 215. Lichanura roscofusca gracia from Guajmias. 1935. Phvllorhvnchus, the leaf-nosed snake. Bull. Zool. Soc. San Diego, No. 12, Sept. 12, 1935, pp. 1-31, one plate, one map. LiNSD.\LE, J. 1932. Ami)hibians and leptiles from Lower California. L^niv. Calif. Puhl. Zoul., 38, No. 6, June 24. 1932, pp. 345-386. 1933. The leaf-nosed snake in Sonora, Mexico. Copeia, No. 4, Dec. 27, 1933, ])i>. 222-223. Phyllorhynchus deci(rt(iti(.-< broirui Stejneger (Alamo Muerto, thirty miles west of Caljorca, Son.). Ortenburger, a. I. 1928. The whiyisnakes and racers: genera Masticophis and Coluber. Mem. Univ. Mich. Mus., I, pp. 1-XVIII, and 1-247. RuTHVEN, A. G. 1908. Variations and genetic relationships of the garter- snakes. Bull. Nat. Mus., No. 61, 1908, pp. 1-201, figs. 1-82, pi. 1. Schmidt, Ivahl P.atteesox. 1922. The amphibians and reptiles of Lower California and the neighboring islands. Bull. Amor. Mus. Nat. Hist., 46, pp. 607-707, pis. 47-57, text figs. 1-13. Smith, Hob.^rt M. 1935. Descriptions of new species of lizards from Mexico of the genus Uta, with notes on other Mexican species. Univ. Kansas Sci. Bull.. XXII, Ainil 15. 1935, No. 7. pp. 157-183. PI. XXVI. Uta taylori described from Gua>'mas. Via ■■-, University of Kansas Abstract: The report is based upon a collection of reptiles and amphibians, now on deposit at the University of Kansas, secin-ed by Edward H. Taylor in the southern part of Siualoa, July 19 to 25, 1934. Two hundred and forty- one specimens were secured: Amphibians, 128 specimens, twelve species; lizards, 94 specimens, ten species; snakes, 16 specimens, seven siu'ci{>s; turtles, three specimens, one species. A list is given of the species known from the state. Bnfo kclloggi, sp. nov., is described from a locality near Mazatlan; and Trimor- ■phodon pancimaculatus, sp. nov., is described from Mazatlan. A bibliography is app(nided. THE basis for this report is a collection of reptiles and amphib- ians which T made in the soiitlieni i)art of Sinaloa during the summer of 19:34. The period from July 19 to July 25 was spent in the region about jNIazatlan and Presidio, and 241 specimens were collected. These are distributed as follows: Amphibia, 128 speci- mens, twelve species; lizards. 94 specimens, ten species; snakes, 16 specimens, seven species; turtles, three specimens, one species. The following extracts from the field book show date and locality of the collections: July 19. Journeyed by "autovia" to a i)oint a few miles beyond Presidio, which is situated on the Rio Mazatlan about fifty miles south of Mazatlan. Returned to Mazatlan at night. Terrain flat, with low forest. July 20. Collected in the general area about two miles east of Mazatlan. A low rock}- hill was visited. Rain pools in this same ^•icinity were visited at night. July 21. Same as previous day. July 22. Xo collecting. July 23. Returned to Presitlio and collected within a three-mile radius of the village. July 24. Collected in the region about Mazatlan, visiting rain pools at night. July 25. Collected during the morning in series of hills two to fi\e miles east and southeast of the city. Departed for Nayarit in afternoon. (505) 506 The University Science Bulletin There \Yere a few heavy rains during this time, making rather ideal conditions for collecting amphibians. The hills near Mazatlan were low; none were visited that reached an elevation above three hundred feet. Consequently there was little variation in the general ecological associations. The rain-pool associations were of a temporary nature. Sinaloa is a region of importance to students of animal distribu- tion, as it is a boundary between two faunal regions. The fauna of the southern part differs to the extent of at least seventy percent from the fauna of Sonora, which lies to the north. There is no strongly defined dividing line crossing the state, since there is an intermingling of the two faunas through a rather wide area, and much more extensive collecting must needs be done before the limits even of individual species can be accurately determined. The following species have been reported from Sinaloa. Almost without exception these records are from tlie southern part of the state in the region about ]\Iazatlan and Presidio, Rosario and Plomosas. A few of the records are ciuestioned. AMPHIBIA '? Lcptodactylus mela)iO)iotns (Hallowell) Eleuthcrodactylus mcxicamis (Brocchi) Bujo siuDis Schmidt Bitfo marmoreus Wiegmann Bujo (lebilis i^Bujo kellof/gl. ?\^. nov.) Pternohyln jodicns Boulenger Din(/l( no spa/iildta (iiinther Agalychnis dacnicolor Cope Hyla arenicolor Cope Hyla baudinii Dumeril and Bibion Hypopachus oxyrjiinus Boulenger Microhyla usta (Cope) Bonn jnrnri Boulenger (^ '? Bonn pipicii.'^ Schreber) TITRTLES Emyt^ or IK it (I Gray Eniys pulchcrrima Gray Kinosternon hirtipr.9 Wagler CROCODILES Crocodi}u,'< ariilxs Cu\'ier LIZARDS 1 Sphiu rodiu-l ylus larqiuil us Straucli (iirobably inti'oduced) Phyllodnctylus land Smitli ? Phyllodfictylus (}d)( rculosu.s \\'ieginann Taylor: Fauxa of Sixaloa 507 Peropus mutilatiis (Wiegmann). Introduced. Coleonyx fascial us (Boulenger) Anolis nebidosus (Wiegmann) Anolis utowanae Barbour ? Callisaurus vcntralis Hallowell Ctenosaura pectinata (Wiegmann) Ctenosaura acanthura (Shaw) Iguana rhinolopha Wiegmann ? Holbrookia maculala approximans Baird ? Holbrookia propinqua Baird and Girard Holbrookia elegans elegans Bocourt Uta ornata lateralis (Boulenger) Uta bicarinata (A. Dumeril) Via tuberculata Schmidt Sceloporns nclsoni Cocliran Sceloporus clarkii boulengeri (Stejneger) Hclodcrvia horridum Wiegmann ? Cnemidophorwi sexlittcalus Linnaeus Eumeces humilis Boulenger Eumeces parvulus Taylor SNAKES Geophis redimita Cope Pseudoficimia frontalis Cope Ficimia qnadrangulans Glinther Rhinochilus antonii Duges Larnpropcltis anuulatus (Kennicott) Drymarchoii corais (Cu\ior) ? Masticophis flagcllum flavigularis (Hallowell) Maslicophis semilinealus (Cope) Drymobius margariliferus (Schlegel) Leptophis diplotropis (Glinther) Xatrix valida (Kennicott) Hypsiglena torquata (Giinther) Tropidodipsas philippii (Jan) Tantilln bimaculatum (Cope) Leptodeira punctata (Peters) Leptodeira maculata (Hallowell) Trimorphodon bi-scutatus (Dumeiil and Bibron) Oxybelis acuminatus (Wied) Micrurus diastema distens (Kennicott) Agkistrodon bilineatus (Giinther) Crotalus basiliscus (Cope) The earliest collecting done in this region was previous to 1868, on which date a collection, made by Ferdinand Bishoff, was received at the Smithsonian Institution. A notable collection was that of Alfonso Forrcr, who collected at Presidio and Mazatlan about 1885. He obtained the types of several 508 The University Science Bulletin forms of amphibians, including Hypopachus oxyrhinus Boulenger, Rana forreri Boulenger, Pternohyla fodiens Boulenger and Diaglena spatulata Gunther, and numerous reptiles, including types of Eu- meces bocourti (= Eumeces himiilis Boulenger), Uta lateralis Boul- enger, and Ficimia quadrangularis Gunther. A small collection was made by J. A. Kusche at "Venodia," Si- naloa, where he obtained a specimen of the rare Diaglena and other important specimens. E. W. Nelson and Edward A. Goldman collected in Sinaloa dur- ing parts of 1897, 1898 and 1899. They journeyed through northern Sinaloa from Agiabampo to Culiacan, and from Altata across the state to Chacala, Durango. They also visited the southern part of the state, collecting at Mazatlan, Rosario and Plomosas. Numerous specimens were collected, including the types of Sceloporus nelsoni Cochran. Paul D. R. Riithling made a small collection at Mazatlan, in 1920. AMPHIBIA Scaphiopus couchii Baird (Plate XLIV, figs. 1-4) Nineteen specimens (Nos. 2902 to 2920) of this species were col- lected on a hill about two miles east of Mazatlan. They were found at night along a path leading up to a deserted gold mine. Kellogg {loc. c;Y.)has reported this species from Acaponeta, Nayarit, about ninety miles to the south of Mazatlan. These specimens ap- l)ear to be tlie first record for Sinaloa. These specimens ai^i^ear to differ from the typical form in having the skin of the occipital and interorbital region almost completely involved in the ossification of the skull, dcsi)ite the fact that none appear to be full grown. There are certain other differences such as the width of the "shovel" metatarsal tubercle, the larger size of the eye and the greater width of the skull in si-)ccimcns of equal snout- to-vent length. Bufo nKtriniis Linnaeus A single specimen (No. 951) was routt'd from under a log of drift- wood in the sandy bed of the Mazatlan riwr at Presidio. It has not jireA-iously been reported in Sinaloa. but Kellogg has reported a sju'cimen from Gamoa, Sonora, far to the north. Biifo piinctdtu.^ Baird and (iirard A single specimen (No. 131) was collected under a rock at the top of a small hill about three miles southeast of ]\Iazatlan. It agrees Taylor: Fauna of Sixaloa 509 with specimens from Sonera and Arizona in color and markings, as well as in most structural characters. Although a small specimen, the supraorbital crests are more strongly developed and the parotoid glands are very much larger (nearly double). The ventral surface has numerous black dots on anterior half. This is apparently the southernmost point in western Mexico where the species has been found. It has not hitherto been reported from the state of Sinaloa. Bnfo valliccps Wiegmann Seven specimens of a toad (Nos. 373-379) found two miles east of Mazatlan have been tentatively referred to this species. There are, however, differences evident when compared with typical valliceps from Oaxaca and Guerrero. The cranial crests are high, relatively narrow, and vdiivil with black. The tympanum is large, equalling more than half the di- ameter of the eye. The subocular crest is obsolete or rei)resented only by a few tubercles. The first finger is longer than the second. There is a slight fold or ridge on snout from lip to between nostrils ; there is only a faint suggestion of a parietal crest. The measurements show that these specimens are larger than typical valliccps, and may belong to the form briefly described by Cope as Bufo argillaccus.'^' Meusuremcnt.s (in nun.) of Btifo vallia pa Wiegmann Number. Sex Snout to vent Length of head to posterior edge of t.\ni|i;uHun . . . Width of head, greatest Width interorbital crests Width upper eyehd Foreleg First finger Second finger Hind leg (from anus) Tibia Foot (to tip of longest toe) 373 9 86 24 2'J 8 7 .50 8. 7 95 30 40 379 9 83 24 30 9 7 48 8 7 92 30.5 41 374 9 24 29 8.5 6.6 48 8.6 7 95 28.5 44 375 9 63 6.5 5.8 77 22 32 * Proc. .\cad. Nat. Sci. Phila. 20, 1868, p. 138. This form is regarded as a synonym of inarmorens by Ivellogg (in;i2), who had the types available for study. 510 The University Science Bulletin I have as yet not compared these specimens with the type. It is probable that if this were done other differences would appear which, together with those mentioned, might warrant giving a specific name to this form. Biifo kclloggi sp. nov. (Plate XLV, figs. 1-3) Holotype. No. 21, EHT & HMS Collection. Two miles east of Mazathln, Sinaloa, July 21, 1934, E. H. Taylor, collector. Para- types, Nos. 15 to 20, and 22 to 40, Mazatlan, Sinaloa. July 20-21, 1934, E. H. Taylor, collector. Diagno:^is. A small species related to Bufo insidior and Biifo debilis, but differing in having rather well-developed supraocular, preocular, postocular, subocular, and canthal crests, all surmounted by conical tubercles, each terminating in a sharp spine; interorbital space strongly concave, becoming more shallow in the intercanthal region; loreal region nearly vertical; the inner palmar tubercle not as large as a subarticular tubercle; median i^almar tubercle round, subglobular; parotoid gland large, but low and inconspicuous, broad as long, extending as low on the side of neck as the angle of the jaws, heavily studded with spines. Entire dorsal and lateral sur- faces of body and limbs studded with large conical tubercles, ter- minating in one or more brown-tipped spines. Description of the type. Snout seen from above narrow, short, sharply truncate, the canthi with sharp canthal ridges surmounted by pointed tubercles, the area between forming a V-shaped trough; nostrils lateral, very near the extreme anterior tip of snout ; loreal region sloping slightly, slightly concave between nostril and eye; the suborbital ridge continued more or less distinctly to below nostril (sometimes tending to connect by a row of tubercles with the continuation of the canthal ridge in front of the nostril) ; from a dorsal view the edges of jaws are not visible; the supraorbital ridge semicircular, continuous with a very strong preocular crest which is nearly vertical, but does not reach the subocular crest; postocular crest not always well developed, while the supratympanic crest is not or barely defined; tympanum moderately distinct, its vertical diameter about one third the longitudinal diameter of the orbit; length of snout from in front of eye about three fifths of the length of the orbit; width of upper eyelid about three fourths the narrowest distance between the supraocular crests; interorbital and intercanthal region concave, with spiny tubercles, the area between Taylor: Fauna of Sinaloa 511 the junction of the supraorbital and canthal crests somewhat in- flated; lower edge of the upper lip with a series of small, low tu- bercles forming a slight ridge; in lateral profile the snout projects much beyond the mouth and slopes back from the tip to the mouth rather gradually; the ventral outline of the upper jaw is distinctly not circular — the sides if projected would form a right angle. Tongue narrow, elongate, rounded but not nicked behind; choanae large, separated by a distance of little less than distance between nostrils, partly concealed by the overhanging jaw (male with a median vocal pouch i. The parotoid gland is large but not strongly salient, attaining its greatest width very near its most anterior point, the dorsal border only slightly notched. Body covered with prominent spinose tubercles on dorsal and lateral surfaces, smaller on latero-ventral abdominal surface, be- coming still smaller medially; on breast the tubercles are more prominent, each with a distinct spine. Limbs, save in postfemoral and posthumcral regions, strongly spinose; first finger slightly shorter than second; inner palmar tubei'cle small , no larger than the subarticular tubeicles; hind leg short, the tibiotarsal articulation not or barely reaching axilla; legs folded at right angles to body, the heels narrowly fail to touch; femur in\'olvcd in the body skin nearly two thirds of its lengtli; toes about one third to two fifths webbed; two small metatarsal tubercles. Color in life. Above yellowish-brown with numerous blackish- brown markings, sometimes tending to form elongate spots; an irregular, light yellowish-brown stripe from labial border to groin, more or less interrupted in parotoid region; legs and arms marked with heavy bars, continuous when limb is folded; foot strongly spotted; below dull brownish-yellow with scattered si)ots of blackish on the lateral abdominal region and across the pectoral region; chin immaculate yellowish (in males the chin is dark graj'ish to blackish). 512 The University Science Bulletin C3i O cq s a3 0) N O) X o Tf ^ t^ o o ■^ — -* 0) CO r,. _. 0 O) CO 00 CO 9 •o I-- ^ ^^ O lO CO 00 s- co c co CO •* CO CO CO b- Ol O) c CO 00 i-H ??°* IM oc lO lO t OC O) T— * lO d Tf CO CO CO Ol CO 10 ■* O) ■* C^ t Ol ^o iC 00 oc 0 1-0 oc 0 (M o ^ LO CO CO CO O] cs CO c 10 _l ■* e^ ■* T— IN CO o 2 o o 'J 05 lO CO 10 ^ ■* LO 5;o ■* o ,_,. O) ■* *y; o CO ,- ■* ^ -* o -t Ol S^ o .^ Ol OC CO ,- O) c^ lO o ,r: CO Tt CO o CO CO r- lO O] ■* " o Tt CM ^°' <-0 00 1-0 ^ O) r^ 10 Ol o ■* o d 10 CO Tf CO CO CO ;- lO ^' ■^ " o -* C4 CO CO o CO „ S^o* CO •r" ^ 10 CO t CO o Ol CO t- 10 Ol T)< " " o Tt OJ g ;a ^ a c3 1 ^ - ^. c ^ V.-. * o C _2 ai 'f bO 'J a -T3 c u c *^ o c i-. c ^ ^ (L c -M c -t- 5 CC 1- £ c L S ^ £ > U "•^ ^ ^ " _. c a. 3 o: a r a. J <^ = " — i~ a. t. > c "2 is "o j c .^ t] 1 ^ i 1 c c C c- c c "a > 1 c ^ ^ H o Taylor: Fauna of Sinaloa 513 Variation. The large series displays a remarkable constancy in color markings, body proportions and the general character of the crests and spiny tubercles. There seems to be some variation in the length of the first two fingers; usually the second is a little longer, but occasionally they are very nearly the same length in some younger specimens. Relationship. This small toad is most closely related to Bufo debilis and Bufo insidior Girard. From Bujo dcbilis (Eastern Texas and Tamaulipas) the present species differs in having a slightly larger orbital diameter; in having the cranial crests better developed and studded with sharply pointed tubercles; in having a shorter leg, the tibio-tarsal articulation reaching only to axilla; in having better developed spiny tubercles on the dorsal, lateral and ventral sur- faces of the body; a less prominent, somewhat differently shaped parotoid; the interorbital width distinctly less; and in a totally different color pattern. (See plate XLV, figs. 4-6.) From Bufo insidior Girard (Kansas, Texas, New Mexico, Chi- huahua, Durango and Zacatecas) it differs in the presence of the cranial crests (lacking or with only an occasional faint trace of crests in insidior) in having shorter hind legs, a shorter snout, larger eye, narrower head and narrower interorbital width, a differently shaped parotoid and a totally different dorsal color pattern. It has very much larger and more numerous spiny tubercles on dorsal and lateral surfaces; the inner palmar tubercle is less developed, as are the metatarsal tubercles; the webbing between the toes is slightly more extensive. The color pattern is entirely different. (See plate XLV, figs. 7-9.) Remarks. The specimens were collected in the daytime under rocks and logs, and at night hopping about on a small flat near a rain pool only a few hundred meters from a tidewater bay. They were not heard calling. The revival of the name insidior for the small toad occupying a very extensive territory extending from Kansas south through New Mexico, Oklahoma, Texas, Coahuila, Chihuahua, Durango and Zacatecas is, I believe, wholly warranted. Specimens from Kansas have the same characteristics, including almost the same identical color pattern as those from the southern part of the range in Zaca- tecas. Large series are at hand for comparison. 33—6037 514 The University Science Bulletin The species is named for Dr. Remington Kellogg of the United States National Museum, whose excellent work, "Mexican Tailless Amphibia," appeared in 1932. Leptodactylus occidentalis Taylor Leptodactylus occidentalis Taylor. Trans. Kansas Acad. Sci., 39, 1936, pp. 349-352. Three specimens of this species were collected near Mazatlan. One had been swallowed and partially digested by a snake, Lepto- deria personata (No. 566). These specimens agree with typical specimens from Tepic, Nayarit, from which locality the species has recently been described. Pternohyla fodiens Boulenger Ptemohyla fodiens Boulenger, Ann. Mag. Nat. Hist., Ser. 5, X, No. 58, 1882, pp. 326, 327. (Type description; type locality, Presidio, fifty miles from Mazatlan, Sinaloa ; Forrer, collector.) A series of fourteen specimens, Nos. 1368-1381, were collected about two miles east of Mazatlan. The specimens were hopping about in the fields, usually not far from rain pools. They were very wary and the approach of my light was a signal for them to make for thick clumps of shrubbery where they were safe. Only very alert action on my part prevented the escape of the specimens taken. All are quite typical. Kellogg reports the species from Mazatlan and Rosario in Sinaloa. Diaglena spatulata (Giinther) Two specimens (Nos. 1423, 1424) of this rare species were col- lected a few miles to the south of Presidio, which is the type locality. Kellogg {loc. cit.) was able to discover only four specimens in col- lections; the three cotypes from Presidio, and a fourth specimen from "Venodio," * Sinaloa. Thus the known distribution is in a very restricted district in southern Sinaloa. Intensive collecting in Nayarit by H. M. Smith and myself, and in Colima by Smith, failed to discover the species in these localities. My Presidio specimens were taken under a small piece of a log on the edge of a shallow stagnant pool. Both were crouched together, and when picked up they remained motionless. The axis of the head was turned nearly at right angles to the body. Whether this is indicative of a phragmotic habit as suggested by Barbourf for * I suspect that this should be Venadillo (pronounced somewhat like "Venodio," which name is not listed in the Directorio General de Correos, but which does list Venadillo, Mazatlan, Mazatl&n, Sinaloa). t Barbour, Reptiles and Amphibians, their habits and adaptations. Houghton Mifflin Co, Boston and New York, 1926. p. 74. Taylor: Fauna of Sinaloa 515 certain related genera, I cannot say. J. Aug. Kusche, who collected a specimen (No. 73266 U. S. N. M.), recorded that it was collected in a termite nest in a tree. Both of my specimens are males. One, obviously, had been call- ing the previous night, as the vocal sac was much distended and folded. In the other the throat showed no external evidence of a sac. The contents of the stomachs were examined. In one there was a single small beetle; in the other were found one beetle and frag- ments of a blattid. In the mouth were four, rather large, brown ants. The eyes of this form appear peculiar in that the eyeball appears to be directed somewhat forward rather than laterally. Both of these specimens have the eye so turned, and Giinther's figure shows somewhat this same condition. I failed to observe the eye in the living specimens. The measurements of Nos. 1424 and 1423 are, respectively (in millimeters) : Snout to vent, 74, 71; length of casque, 29, 27.5; width at eyes, 22, 19; length of arm, 37.5, 36; length of legs, 87-86, 81-83; tibia, 28, 27 ; foot, 37, 35 ; diameter of tympanum, 3, 3 ; diameter of eye, 7.5, 6 ; length of third finger, 19, 18. Agalychnis dacnicolor Cope Phyllomedusa dacnicolor Kellogg, Bull. U. S. Nat. Mus., No. 160, 1932, pp. 143-144. A series of twenty-nine specimens (Nos. 1306-1334, July 21-24) was collected in the general vicinity of Mazatlan, for the most part one to two miles east of the city. All were found in shrubbery or trees near rain pools. All the specimens were grass-green in life, often tending toward bluish-green. When preserved most of them changed to a bluish or violet shade. Eggs were found and pairs were found clasping. They did not appear to be afraid of the light, and were easily collected. Kellogg reports the species from Rosario, Presidio and Mazatlan in Southern Sinaloa. Hyla baudinii Dumeril and Bibron Hyla baudinii Kellogg, Bull. U. S. N. M. No. 160, 1932, pp. 160-163. A single specimen (No. 658) was taken from the throat of Masti- cophis lineatus collected two miles north of Presidio. The frog had apparently been hidden among the dead leaves of a palm tree, from which I shot the snake. The frog is a large female measuring 75 mm. snout to vent. The tibiotarsal articulation reaches the eye. The sides are reticulated with brown, enclosing yellowish spots. The 516 The University Science Bulletin posterior femoral region displays a darker reticulation enclosing lighter yellowish spots. Kellogg reports specimens from "north of Mazatlan" (1) and Plumosas (2). Hyla smithii Boulenger Hyla smithii Taylor, Trans. Kans. Acad. Sci. XXXIX, 1936. A single specimen (No. 2174) was taken on a floating plant in a rain pool about two miles east of Mazatlan. About this same pool were found Pternohyla fodiens, Agalychnis dacnicolor, Micro- hyla olivacea, Bufo kelloggi and Bufo valliceps. Others were heard calling. The specimen in life was canary to lemon-yellow with a few darker dorsal spots among the very numerous minute dark flecks (under the microscope many of the minute spots are star- shaped). The specimen is an adult male with the vomerine teeth well developed. Snout to vent, 22 mm. Microhyla olivacea (Hallowell) Engystnma olivareum Hallowell, Prop. Acad. Nat. Sci. Phila., 1857, p. 252. (Type descrip- tion; type locality "Kansas and Nebraska.") Gastropliryne olivacea Smith, Amer. Mid. Nat., XV, 1934, No. 4, pp. 501-505. I have tentatively associated with this species three small micro- hylids (Nos. 1236 to 1238) collected about two miles east of Ma- zatlan under rocks at the base of a small clay hill. When com- pared with Texas specimens of equal size they differ in having a narrower head, the snout a little more projecting and more flattened. They are somewhat darker and on the side, from snout to groin, the pigment tends to form a darker broken line. There is a slight difference in the shape of the foot and the metatarsal tubercle is slightly more salient. They differ somewhat less from a series of specimens collected by Hobart Smith and David Dunkle at Cone- jos, Durango. None of the three specimens approaches the maxi- mum size of olivacea. A larger, more representative series may demonstrate that these and perchance other characters warrant a specific designation for the coastal form. If properly associated with olivacea these records extend the known range some 200 miles farther to the southwest. Rana jripiens Schreber Four specimens were collected: No. 2926, three miles east Ma- zatlan, July 20; 2927-2928 near Presidio, July 24; 2929 near Ma- zatlan, July 25. No. 2926 is a female measuring 125 mm. snout to vent. The ovaries are packed with ripe eggs. Kellogg {op. cit.) reports specimens from several localities in Sinaloa. Taylor: Fauna of Sinaloa 517 Phyllodactylus lanei Smith Phyllodactylits lanei Smith, Univ. Kansas Sci. Bull., XXII, April 15, 1935, pp. 125-132, plate XXV, fig. 3 (photograph of type). (Type description; type locality, Tierra Colorada, Guerrero. E. H. Taylor and H. M. Smith, collectors.) Three specimens (Nos. 534, 535, and 704, July 21-24, 1934) were collected about two miles east of Mazatlan, Sinaloa. H. Smith has studied these specimens and has referred them to this species^ All are immature, but they agree well with juveniles of the series from Guerrero. Coleonyx fasciatus (Boulenger) (Plate XLVI, fig. 2) Eublepharis fasciatus Boulenger, Cat. Liz. Brit. Mus., 2d Ed., I, 1S85, p. 234. (Type de- scription; type locality "Ventanas [Durango] ; A. Forrer, collector); Giinther, Biologia Cen- trali- Americana Reptilia and Batrachia, April, 1893, p. 84, pi. xxxi, fig. a (entire animal natural size and head). Coleonyx fasciatus Taylor, Univ. Kans. Sci. Bull., XXII, Apr. 15, 1935, pp. 203-205. Elsewhere {loc. cit.) I have reported on this specimen collected about ten miles south of Presidio, Sinaloa, June 19, 1934. It has the following measurements: Snout to vent, 59 mm.; tail, regen- erated, 53 mm.; width of head, 10 mm.; head length, to angle of jaw, 15 mm.; foreleg, 17 mm.; hind leg, 22.5 mm. Ctenosaura pectinata (Wiegmann) Ctenosaura pectinata Bailey, Proc. U. S. Nat. Mus., 73, 1928, pp. 24-27; Smith, Univ. Kansas Sci. Bull., XXII, April 15, 1935, pp. 134-137. The following specimens were taken: Nos. 525-527 ten miles south of Presidio, July 19; No. 583 near Mazatlan, July 20, and Nos. 654-656 near Presidio, July 21. Smith {loc. cit.) discusses these specimens. Iguana rhinolopha Wiegmann Iguana rhinolopha Smith, Univ. Kansas Sci. Bull., XXII, April 15, 1935, pp. 134-137. Four specimens, Nos. 650-653, were collected at Presidio, on the edge of the Rio Mazatlan, July 22, 1934. Smith {loc. cit.) reports on these specimens. Holbrookia elegans elegans Bocourt Holbrookia elegans Bocourt, Miss. Sci. au Mexique; Rept. Batr., Liv. 3, 1874, pi. XVII bis, fig. 8, 8a, dorsal and ventral view of head. (Type description; type locality Mazatlan, Sinaloa.) Holbrookia elegans elegans Smith, Univ. Kansas Sci. Bull., XXII, April 15, 1935, pp. 191, 194, pi. XXVII, fig. 2 (photograph dorsal view), and XXVIII, fig. 5 (femoral pores). A series of six specimens, Nos. 643-648, were taken near Presidio, Mazatlan, Sinaloa, July 22, 1934. Smith {loc. cit.) has commented on this series. 518 The University Science Bulletin Uta tuberculata Schmidt Uta tuberculata Schmidt, Amer. Mus. Nov., No. 22, 1921, p. 4. (Type description; type locality, Colima, state of Colima, Mexico; Riithling, collector.) Smith, Univ. Kan. Sci. Bull., XXII, April 15, 1935, pp. 171-172, pi. XXVI, fig. 1 (photograph of E. H. T. Collection, No. 552, male). A single specimen, No. 552, was collected about fifteen miles south of Presidio, Mazatlan, Sinaloa. Smith {loc. cit.) has reported on this specimen. Anolis nebidosus (Wiegraann) Three specimens, Nos. 553, 554, taken near Presidio, and No. 761, near Mazatlan, are referred to this species. Body slightly compressed, the forehead concave ; supraorbitals are in contact and continued forward as two divergent frontal series, but not forming a frontal keel; normally three supraoculars sepa- rated from the supraorbitals by a row of granular scales; four rows of loreals ; four canthals ; a slight nuchal crest ; occipital large, very much larger than auricular opening; six or seven labials to below middle of eye; gulars slightly keeled; ventrals keeled, slightly smaller than the dorsals which pass rather gradually into the granular lateral scales; enlarged postanals; fourteen scales under the second and third phalanges of fourth toe; gular appendage large, reaching beyond the thorax, grayish or with a pinkish tinge. Markings very indistinct save that radiating lines about the eye are evident. Sceloporus nelsoni Cochran Sceloporus nelsoni Cochran, Jour. Washington Acad. Sci. XIII, 1923, May 4, pp. 185-186. (Type description; type locality, Plomosas, Sinoloa, Mexico. Nelson and Goldman, collectors.) The following specimens collected in Sinaloa are referred to this species: Nos. 577, 578 about two miles east of Mazatlan, July 20, 1934; Nos. 755-760 about four miles southeast of Mazatlan, July 25, 1934. The relationship of this form is obviously with pyrocephalus, but the differences pointed out by Doctor Cochran, while small, are numerous and certain of them rather constant. The males of my series have, usually, a well-defined black shoul- der spot extending somewhat on the arm, while on the breast on either side is an orange or brick-red spot touching the black spot and partially surrounded by darker color. The anterior part of the abdomen is whitish or yellowish. The chin is whitish and bluish- gray, the white forming a number of converging narrow lines usually continuous with lines extended across labials and side of head. These lines are narrower and more numerous than those in pyro- Taylor: Fauna of Sinaloa 519 cephalus. The belly is black medially, bordered by bluish color on each side, this fading to dim yellowish-brown which appears laven- der in preserved specimens; a darker lateral band with a few yel- lowish flecks usually present. The femoral pores vary between 15 and 18, 16 and 17 occurring most frequently. The dorsal color is grayish-brown, with two olive, dorsolateral lines, the edges of which are not clearly defined; tail with fourteen narrow light annulations, immediately in front of which the ground color is darker than remaining interspace; a dark light-ringed spot on posterior part of occipital scale. Females are grayish-brown, the dorsolateral stripes scarcely or not discernible and with small, paired, indefinite, dark spots pres- ent; belly immaculate whitish (of a greenish cast in one) ; chin less distinctly striped, the dark shoulder spot and the orange breast spot less distinct than in males (or obsolete). The tail is much more strongly banded, the whitish or cream rings widening dorsally. Sceloporus utiformis Cope Sceloporus utiformis Cope, Proc. Acad. Nat. Sci. Phila., 1864, pp. 177. (Type descrip- tion; type locality Colima, Colinia, Mexico); Bocourt, Miss. Sci. au Mexique; Kept. Batr., Liv. 4, 1874, pp. 208-210, pi. XVIII bis. fig. 6, Ga, 6b (dorsal view of head, detail of ear, and dorsal scale) (Colima). A single specimen (No. 555) of this very well-defined species was taken about twelve miles south of Presidio, July 19, 1934. The specimen was discovered in the bed of a dry creek and was the only one seen during my two days collecting near Presidio. Compared with Cope's type description the number of femoral pores is larger, 16-17 in this specimen while the type has but 13. The description states, 'Ten longitudinal rows of large, highly keeled, shortly mucronate dorsal scales, separated by many lateral series of minute flat scales, from the smaller entire edged abdomi- nals.'' This specimen has approximately eight rows on the neck, ten on the posterior part of the body, while near the middle there are fourteen rows, the large scales pushing down on the sides. It is probable that this condition may also obtain in the type. The ear opening is very large and the tympanum relatively super- ficial ; the lateral nuchal pocket is especially deep, and the preauric- ular lobules much reduced. The head scales are distinctly rugose. Strongly reddish-brown above, with ten narrow, irregular trans- verse dark bands, edged posteriorly with greenish or yellowish-white, the light edging formed by small white spots that are not continuous; this light color appears on the sides as numerous irregular white flecks each involving several granular scales. The tail is completely 520 The University Science Bulletin encircled by brownish bands four scales wide, separated by wider lighter bands; head variegated brow^n. Upper labials traversed by six greenish- white, vertical stripes which continue on the throat and chin, those below eye strongest and separated by deeper brown interspaces. Limbs barred with wide dark and narrow greenish or yellowish-white bands. Throat gray, reticulated with white; belly and under side of limbs dirty white. Sceloporus clarkii boulengeri (Stejneger) Sceloporus boulengeri Stejneger, North Amer. Fauna, No. 7, May 1, 1893, p. 180, pi. 1, fig. 5a-c. (Type description; type locality, Presidio, fifty miles from Mazatlan, Sin. Forrer, collector.) Sceloporus clarkii boulengeri Burt, Trans. Micros. Soc, Vol. LIV, No. 2, April 1935, pp. 171, 172 (part) (confuses S. horridus oligoporus and S. melanorhirms with boulengeri). The following specimens were obtained: Nos. 528-534 ten miles south of Presidio (sixty miles south of Mazatlan), July 19; No. 587, two miles east of Mazatlan, July 21; Nos. 667-669 (skeletons) and 670-683 about two miles north of Presidio, July 23. The specimens of this series, with the exception of No. 587, are topotypes. The femoral pores in twenty specimens (40 counts) have the following frequency: 8 occurs seven times; 9, seventeen times; 10, fourteen times and 11 twice, or an average of 9.02. Burt's {loc. cit.) comments on the femoral pores are due to his confusion of Sceloporus horridus oligoporus (Cope) (Klauber No. 7335 Zi- huatenejo, Guerrero, with two femoral pores) and Sceloporus melanorhinus (Klauber No. 7336 Zihuatenejo, Guerrero, with twenty- one femoral pores, and No. 10153 Petatlan, Guerrero, with a high number of pores). The femoral pores are not more variable in Sceloporus than in other forms. All the specimens were shot from large trees that stood isolated in pasture fields. Each of these scattered trees harbored rather large colonies. Very large series could have been taken had I so desired, as many specimens were left unmolested. Cnemidophorus sacki Wiegmann Cnemidophorus sexlitwattis gularis Burt, U. S. Nat. Mas. Bull., No. 154, 1931, pp. 97-122 (part). Cnemidophorus sexlineatus sackii Burt, Proc. Biol. Soc. Wash., 44, June 29, 1931, pp. 73-78. In endeavoring to determine the proper name for the Cnemidoph- orus from southern Sinaloa, I have found it necessary to review the specimens of Cnemidophorus gidaris and Cnemidophorus sexlineatus in the collections of Kansas University (about 200 of the former and 500 of the latter) and especially specimens from Texas and Taylor: Fauna of Sinaloa 521 Oklahoma, where the ranges occupied by these species overlap a known distance of more than five hundred miles. Special attention was given those specimens from Oklahoma and Texas with a view of discovering intergradation between these forms, a condition which I had not observed during extensive collecting in Texas. I have had available about 200 specimens from these states. I am wholly un- able to discern intergradation in the pertinent characters (which may be used to separate the species) that suggests in any way a crossing of these species. There obtains in adults of both sexlineatus and gularis strong sexual dimorphism in color and markings. The male sexlineatus has the ventral surface colored a very light blue in life, the color not or but slightly more intense on the throat; however, this blue color is lost in preservation and various changes result, depending upon the preservative used. The throat usually becomes a different shade of blue and the abdomen seems to be darker blue on the sides. In formalin the throat may become blackish and the abdomen like- wise. The females of this form are rather creamy yellow ventrally, which color remains or becomes whitish in alcohol. In formalin the sides of the abdomen are a little darker. The maximum size in Texas and Oklahoma probably rarely exceeds 75 mm. snout to vent. The gularis of this region have a strongly-defined pinkish to red- dish-pink coloration on chin and throat often extending to the en- larged scutes on the breast; the ventral surface of the body of adult males is dark blue-black anteriorly, becoming somewhat bluish posteriorly, and with white, cream or bluish-white spots present, es- pecially on the outer part of abdomen. Undersides of the hind limbs (partially), anal region and often the underside of tail are immaculate cream. The females, however, are without either the pink throats or bluish bellies and as such are very often confused with sexlineatus. Data from these misidentified females taken with data from sex- lineatus bring about a disheartening confusion which suggests in- tergradation. While it is not certain, it seems highly probable that Burt (loc. cit., 1931) has confused the females of gularis with both sexes of sexlinatus, since he does not note this sexual dimorphism. He states (p. 83) : "At the point of intergradation with gularis there is a complete transition from the characteristic granules on the post-antebrachium of sexlineatus to the larger, better developed, polygons that are usually found in that form" {gularis). This statement is wholly true when applied to series containing both the 522 The University Science Bulletin ventrally unmarked females of gularis and males and females of sexUneatus, and is not true when applied to sexlineatus males or females of the material available in the Kansas University collec- tions. In consequence, I believe that gularis should not be associated with sexlineatus as a subspecies as Bocourt proposes (Mission Sci. Mexique 1874), Cope (1900) and followed by Burt (1931). The application of the name C. sexlineatus I believe should be limited to the small Cnemidophorus described by Linnaeus, occupying ter- ritory over a wide area in the eastern and central United States and as far west as Texas, Oklahoma, Kansas, Colorado and New Mex- ico. It may occur in northern Mexico. The cnemidophori obtained in Sinaloa include the following: Nos. 535-551, ten miles south of Presidio, July 19; 581, 588-590, along edge of tide flats near Mazatlan, July 20 and 21; 684-688, in the vicinity of Presidio, July 23; 750-754, near Mazatlan, July 24. It is with some hesitancy that I am associating the above name with the present species. Burt (Proc. Biol. Soc. Wash., Vol. 44, June 29, 1931, pp. 73-78) has recently published additional details of the type of C. saoki which Parker had observed on the type. These details apply to certain of the specimens before me. If these are not typical they would certainly have a subspecific relationship with sacki. The specimens present the following characters: A six-lined form, the lateral line extending to the tip of the snout, crossing the sub- oculars, passing above the ear and terminating in the groin. The dorsolateral line arises at the last superciliary and continues a considerable distance on the tail. The dorsal lines are less dis- tinct and are separated by a ground color much lighter than the dark- brown color between the other lines. There are small dots between the lateral and dorsolateral lines, males and females showing little or no sexual dimorphism. In larger specimens the lines break up into series of white dots which with the intervening dots tend to form light, broken, transverse bands. In still older specimens the area occupied by the longitudinal lines in the younger specimens now appears as the ground color, with a tendency to form inter- rupted dark transverse bands. The ventral surface of chin and throat is bluish-gray in adult males and females, rarely with a trace of pinkish. The bellies are deep blue-black with numerous bluish- white spots. The tail beneath has some bluish or blackish spotting. The specimens from the tidewater flats have somewhat less dark Taylor: Fauna of Sinaloa 523 color below, but it is strongly evident on the sides. The dorsal linear markings do not break up into dots; the upper surface of the hind limbs are uniformly colored. The numerous spots on the Presidio specimens are lacking. The anterior continuation of the lateral line across the loreal region is strongly pronounced in young and old. The chin and throats are more strongly contrasting black and yellow. The enlarged scales on the gular fold are yellow-cream in all. The largest size noted in the series is 119 mm. snout to vent, while several specimens measure 110 mm, or more. Measurements (in mm.) and scale counts of Cnemidophorus sacki Wieg mann Number, E. H. T. Collection 540 536 539 535 589 548 541 Snout to vent 119 28 30 21 13 76 25 25 34 20, 22 113 36 114 24 25.5 20 11.2 76 24 25 34 20, 21 118 37 110 28 28 20 12 77 25 27 29 20, 20 89 33 no 24 26 18 11 73 22 25 31 18,19 107 35 no 24 25 20 12 72 21 22 34 20, 21 100 34 108 23 24 15 10 74 23 26 35 18, 19 91 35 105 Snout to occiput 23.5 Snout to ear 25 Breadth of head 17 Interorbital width Hind leg 11 74 Tibia 23 Fourth toe 25 Lamellae, fourth toe 31 Femoral pores 17,17 Dorsal scale rows 105 Ventrals (long) 36 Constrictor constrictor imperator (Linnaeus) Two specimens were taken: No. 691, near Presidio, and No. 711, two miles east of Mazatlan. The latter specimen is alive at this date in the laboratory of zoology at University of Kansas. Masticophis semilineatus (Cope) Bascanium semilineatum Cope, Proc. U. S. Nat. Mus., XIV, pp. 622, 626. tion; type locality Colorado river bottom. Schott, collector. Type descrip- A large specimen of this species was collected about two miles east of Mazatlan, July 20. It was observed crawling in sparse brush, and when sighting me, it stopped, reared its head and re- mained motionless for some time. Certain other specimens seen, apparently of this species, escaped in brushy ground. 524 The University Science Bulletin Ortenburger (1928) cites previous records for Mazatlan by Van Denburgh (1897), and for Presidio by Giinther (1894). My specimen, a male, presents the following characters: Ven- trals, 192; caudals, 109+ ; scale formula, 19-17-17-13-13; upper labials, 8-8; lower labials, 9-10; preoculars, 2-2, touching frontal; postoculars, 2-2; temporals, 2+2+3+2; loreal, 1-1 (fused partially with preocular) ; head length to jaw angle, 42 mm.; to end of parie- tals, 33 mm.; supraorbital width, 15 mm.; total length, 1685+ mm.; tail 470+ mm.; maxillary teeth, 20-20; mandibular, 21-22; nasal divided; frontal equal to distance to end of snout; eye di- ameter reaches edge of posterior nasal. In general, the color is typical. The ventrals have series of punc- tate spots bordering their edges, absent from a few. On the chin and throat the spots are much larger and darker. Masticophis lineatus (Bocourt) Bascanion lineatum Bocourt, Miss. Sci. au Mexique, Rept., Liv. 12, 1890, pp. 697, and /00-701, pi. XL VIII, fig., 1, la, lb, Ic. (T>'pe description; type locality, "Mexico." Col- lectors, Alfredo Duges and Boucard.) Masticophis lineatus Ortenburger, Mem. Univ. Michigan Mus., I, 1928, pp. 134-138, pi. XXV, figs. 3, 4, 5. This little-known form is represented by three specimens, collected vTuly 23 at a small pool beside the railway, about one mile north of Presidio (fifty miles south of Mazatlan). A fourth specimen, col- lected at the same time, escaped from my hotel room in Mazatlan when I was preparing to preserve it, and was crushed in the street a few moments later. It was not preserved. No. 663 was found in a tree swallowing a specimen of Hyla baudinii. The frog was recovered alive. The three specimens, Nos. 662 2 , 663 S , 664 $ , present the fol- lowing characters, respectively: Ventrals, 184, 184, 185; sub- caudals, ?, 112, 120; upper labials, 8-8, 8-8, 8-8; lower labials, 10-11, 10-10, 10-10; preoculars, 2-2, 2-2, 2-2 (the large upper scale of each snake shows a strong suture partially dividing the scale) ; post- oculars, 2-2, 2-2, 2-2; scale formulae, 20-17-17-13-13, 19-17-17-13- 13, 19-17-17-13-13; mandibular teeth, 18-19, 18-18, 18-18; maxillary teeth, 18-18, 18-19, 18-18; head length to angle of jaw, 42, 42, 40 mm.; length to end of parietal, 28, 30.5, 27.5 mm.; interorbital width, 14, 15, 14 mm.; total length, 960 (incomplete), 1346, 1376 mm.; tail, 180 (incomplete), 380, 385 mm. All three have four scales touching the first pair of chinshields which are slightly longer than second pair and separated by a few small scales; a single loreal, nearly twice as long as high; preoculars Taylor: Fauna of Sinaloa 525 separated from frontal; the seventh labial exceeds but little the size of the eighth (in one, smaller) ; the temporals are very irregular, but only a single anterior present, followed by two or three; the frontal is as long as, or only minutely less than, its distance to the tip of the snout; the parietals usually about one fifth longer than frontal; the eye equals its distance from nostril or is but slightly less; internasals are from two thirds to three fifths as long as the prefrontals. The specimens agree in colors and markings. Above they are grayish-tan, save on the anterior part of body where the color is a strong lavender- or orchid-gray for a distance of six or eight inches behind head. There are faint suggestions of very narrow pinkish- white bars with some darker dots bordering them. The top of head is uniform dark amber; a cream spot present on the preocular; most of the upper labials are yellowish; a faint vertical light line crosses the seventh labial and extends above it; dorsal scales with fine black dots on apex, missing on some lateral scales, and absent on tail. Below yellow, the gray of sides encroaching on the ven- trals; edges of ventrals on neck region, rosy, and on caudals a rosy or pinkish line, becoming lavender towards tip. Natrix valida (Kennicott) Regina valida Kennicott, Proc. Acad. Nat. Sci. Phila., 1860, p. 334. (Type de.scription ; type locality, Durango, Mexico.) Tropidonotus validus Glinthpr, Biologia Centrali-Americana, Reptilia and Batrachia, July 1894, p. 134. (Presidio and Mazatlan, etc.) Four specimens were collected in the vicinity of Presidio, July 19 and 22. All were in the immediate vicinity of water. The specimens Nos. 563, 564, 640, 641, present the following char- acters respectively: Sex, 5 , ^ , ^ , 5 ; ventrals, 141, 143, 142, 146; subcaudals, 72, 76, 77, 73; supralabials, all 8-8; infralabials, nil 10-10; preoculars, 1-1, 1-1, 1-1, 2-2; postoculars, 3-3, 3-3, 3-3, 2-2; temporals, 1+3+4, 1+2+4 : 1+3+4, 1+2+3 : 1+3 : 1+2+3; labials touch chinshields, all 5; loreal, 0-0, 1-1, 1-1, 1-1; anal, all, divided; scale fcjrmulae, 19-19-17-17, 20-19-17-17, 20-19- 17-16, 19-19-17-17; (measurements in millimeters) head length, 35, 13.2, 24, 26; length to parietal, 25, 12, 18, 18; supraorbital width, 8, 5, 6, 8; length, 870, 308, 605, 635; tail, 207, 80, 155, 156; maxillary teeth number, of Nos. 563 and 640, 24-23, 24-24; mandibular teeth, 25, 25. The three smaller specimens are uniform gray in color, and show two alternating rows of black spots on each side anteriorly. There 526 The University Science Bulletin is a dim lateral line on second and third scale rows, below which the color is slightly darker; first six supralabials with black posterior edges. Lcptodeira maculata (Hallowell) Leptodira personata Cope, Proc. Acad. Nat. Sci. Phila., 1868 (1869), p. 310. (Type de- scription; type locality, "Mazatlan, Western Mexico"); GUnther, Biologia Centrali -Americana, Reptiles, May, 1895, pp. 171, 172, pi. LIV, figs. A (adult, natural size) and B (young). Two young specimens, No. 566 (ten miles south of Presidio, July 19) and No. 666 (one mile north of Presidio, July 23), were taken. Each was found hidden under a log in relatively dry situations. These present the following characters: Ventrals, 165, 167, sub- caudals, 72, 68; supralabials, 8-8, 8-8; infralabials, 9-10, 10-10; preoculars, 1-1, 2-2 (the lower very small) ; temporals, l-|-2+3, l-|-l-f2-|-3 (the anterior temporal segmented); anal, 2, 2; length, 280, 249 mm.; tail, 60, 54 mm.; spots on body, 24, 25; on tail, 12, 13; scale rows, 21-21-21-17; 21-23-23-17. They agree in the following characters: Fourth and fifth subocu- lars enter the orbit ; length of eye equal to its distance from center of nostril; loreal slightly longer than high; posterior chinshields about as long but somewhat more slender than anterior ; five labials touch anterior chinshields; frontal with sides nearly parallel, in contact with the preocular (on one side in No. 666 it is minutely separated) , equal to its distance to end of snout; scales with two apical pits. In both specimens the blotches are somewhat diagonal across the back, and black in color. The dorsal ground color is faun. In No. 666 there is a foreshadowing of the darkening of the ground color, as occurs in the adult, by the appearance of black flecks low on the sides. The head is dark, and the parietals are partially outlined with yellowish-white, and somewhat lighter along the sutures of the other head scales; labials and ventral surface white. The nuchal collar is four or five scales wide, followed by the widest dorsal blotch cover- ing nine to eleven scale rows medially. The type has the preoculars, 2-1 (suggesting as do these speci- mens that these scales are variable), and the upper preocular is sep- arated from the frontal. Leptodeira -punctata (Peters) Crotaphopeltis punctatus Peters, Mon. Ber. Akad. Wiss. Berlin, 1866, p. 93. (Type de- scription; type locality? South Africa.) Leptodira pacifica Cope, Proc. Acad. Nat. Sci. Phila., 1868 (1869) p. 310. (Type de- scription; type locality, Mazatlan; Bishoff, collector); Glinther, Biologia Centrali-Americana, Reptilia and Batrachia, 1895, p. 169; Boulenger, Cat. Snakes Brit. Mus., Ill, 1896, p. 19 (Presidio, near Mazatlan). Sibon pacificum Cope, Bull. U. S. Nat. Mus. No. 32, 1887, p. 67 ; Proc. U. S. Nat. Mus., XIV, 1892, p. 678. Leptodira punctata Boulenger, The Zool., 1887, p. 178. Taylor: Fauna of Sinaloa 527 A single male specimen of this rare snake was captured late at night near a small railway bridge about a mile east of Mazatlan. The specimen was crawling along the bank of a small rivulet which held water from a rain of the previous night. It presents the following characters: Portion of rostral visible above very narrow; frontal longer than its distance from the end of the snout, shorter than the parietals; nostril very large, pierced chiefly in the anterior part of the divided nasal ; loreal small, as high as wide; two preoculars, the upper very high, the lower minute; two postoculars, both in contact with the single large anterior temporal; posterior temporals two; diameter of eye equal to its distance from the middle of the nostril. Upper labials 7-7, the sixth extremely large, the third and fourth entering the eye; anterior chinshields slightly wider but no longer than the posterior; latter scales sepa- rated from the first widened ventral by two pairs of small scales and two single enlarged scales ; lower labials, 9-9, the first four touch the chinshields. Ventrals, 149; anal divided (preceded by a very small median scale); caudals, 70; terminal scale elongate, conical, with slight, lateral grooves. Length, 516 mm.; tail, 130 mm.; head width, 13 mm.; length to angle of jaw, 19 mm. Color in life. Above slightly reddish-brown, with a series of black spots on either side of the median line extending to the tail ; and on the side one or two indefinite rows of irregular black flecks tending to form angular reticulations. Head brown; four small dark spots on the parietals; a small median black spot borders the parietals and on either side of the nape are two large black spots narrowly sepa- rated by a yellowish area; labials very light tan; ventral surface cream. Trimorphodon paucimaculatus sp. nov. (Plate XLVI, fig. 1) Holotype. E. H. T. No. 709, collected at Mazatlan, Sin., Mexico, July 24, 1934; E. H. Taylor, collector. Diagnosis. A species related to T. bi-sciitatus Dumeril and Bib- ron, but not having the preoculars touching frontal, and with the dorsal spots greatly elongated and fewer in number. Frontal as long as parietal; prefrontals as wide as long. Ventrals, 253; anal divided; subcaudals, 76. Description of the type. Part of rostral visible above equal to about one third its distance from the rostral; frontonasals a third wider than long; prefrontals very large, their greatest width about the same as their greatest length or slightly greater; frontal not 528 The University Science Bulletin angular anteriorly, but with an acute angle posteriorly; length of frontal equal to that of the parietal, a little longer than its distance to the end of the snout; the width of the parietal about three fourths of its length. Nostril pierced in the nasal near the supranasal border, and a suture partially dividing the scale runs from nostril to the first labial; the scale is undivided above; three loreals, the anterior upper largest, higher than long, the posterior longer than high, the third loreal lies below and almost wholly posterior to the second loreal, and appears to be formed by a segmenting of the upper part of the third labial ; three pre- and three postoculars, the upper preocular not especially large and well separated from the frontal; temporal formulae, 3+5+4+5, 3+4+5+4; upper labials, 9-9, the fourth and fifth entering orbit. The diameter of the eye equals the distance from posterior edge of the nasal; lower labials, 12-13, five touching the anterior chinshields; mental triangular; posterior chinshields less than half the anterior, and separated from the first ventral by five rows of scales ; eight or nine scales between the first ventral and the last labial; ventrals angular, 253; anal divided; 76 subcaudals, the terminal scute with a dorsal, a ventral and two lateral grooves. Scale formula, 25-23-25-25-25-19-17 (last count made in front of anus). The teeth of this form seem fewer than those in typical bi-scutatiis Dumeril and Bibron. The number present is eight, the first two much enlarged, the first a little smaller than the second; the next five are subequal, rather widely spaced, but no teeth appear to be missing; and then after a space are tw^o grooved teeth a little thicker but scarcely higher than the second tooth. Color in life. A dim olive band, slightly black-edged, crosses snout slightly in advance of the eyes; followed by a lighter band that crosses head between, but curves back behind eyes to the angle of the jaw; this is followed by a broad, black-edged, arched, lavender-gray band nearly severed medially by a projection from the lighter arched mark following the preceding; behind this another wide arched band confluent with the first dorsal spot. General color light, grayish-lavender; on body, 20, on the tail 10 brownish- lavender, darker-edged, saddlelike spots, each with a lighter, dorsal, central portion and separated from the following spot by four or five scale rows, its length involving 11-12 transverse scale rows; a few small scattered spots along side of body; a few dark lavender flecks on the ventrals; body below dirty cream. Measurements. Length, 880 mm.; tail, 142 mm.; head length to jaw angle, 25 mm.; width, 14 mm. Taylor: Fauna of Sixaloa 529 Remarks. This species is, as has been stated, related to T. bi- scutatus. The type locality for the latter is "Mexique," but the high ventral and subcaudal counts suggest a southern specimen. Boulenger (Cat. Snakes III, p. 54) lists a series of seven specimens from Mazatlan and Presidio (fifty miles south of Mazatlan?) which show a relatively uniform scale formula, 25 (24) ; ventral scales, 237-251 (average 247); subcaudals, 78 9 and 84-87 £, average for both sexes being 83. Giinther (Biol. Centrali-Amer., ]\lay, 1895, p. 174), writing of these and certain other specimens, states that they do not agree with the forms included in certain synopses presented by Cope. Neither Boulenger nor Giinther comment on the markings or the relation of the upper preocular to the frontal. It seems likely that the bi-scutatiis as used by Giinther and Boulenger is a composite and it is (luite probable that the lot mentioned above from South- ern Sinaloa actually should be associated with this form. Kinosternon hirtipes Wagler Three specimens (Nos. 560, f>89, (i!)()i, which I collected at Presidio, are in the hands of Dr. Norman Hartweg, who will include data on them in his study of the genus Kinosternon. I am indebted to him for this identification. bibli()(;raphy Bailey. John W. 192S. A revision of the lizards of the genus Ctenosaura. Proc. U. S. Nat. Mu?.. 73, 1928. p]). 1-55, plates 1-30. Barbour, Thomas. 1926. Reptiles and amphibians, liieir habits and adapta- tions. Houghton Mifflin Co. Boulenger, G. A. 1882. De.scription of a new genus and species of frogs of the family Hylidae. Ann. Mag. Nat. Hist.. (5), X, 1882. pp. 326-328. 1883. Description.s of new si)ecies of lizards and fiogs collected by Herr A. Forrcr in Mexico. Ann. Mag. Nat. Hist., (5), XI, 1883, pp. 342-344. Burt, C. E. 1931. A study of the Teiid lizards of the genus Cnemidophorus with special reference to their phylogenetic relationships. Bull. U. S. Nat. Mus., No. 154, 1931, pp. 1-286. 1931a. The status of the spotted race-runner Cne>niduphorws scvlineatus gidaris (Baird and Ciirard). Proc. Biol. Soc. Wa.sh., p. 44, June 29, 1931, pp. 73-78. 1935. Notes on a collection of lizards from Western Mexico and tropi- cal America. Trans. Mic. Soc, LIV, No. 2, April 1935, pp. 167-178. CocHR.\N, Doris. 1923. A new lizard of the genus Sceloporus. Journ. Wash. Acad. Sci., Ill, No. 9, May 4, 1923. 34—6037 530 The University Science Bulletin GuNTHER, Albert. 1882. Notice of a .second .species of Trijirion. Ann. Mag. Xat. Hist.. Scr. 5. X. No. 58. 1882, p. 279. 1900-1902. Biologia Centrali-Americana, Reptilia and Batrachia. Feb. 1900 to May 1902. j.p. 197-326, pis. 60-76. Kellogg, Remixgtox. 1932. Mexican tailless amphibians in the United States National Museum. U. S. Nat. Mus. Bull.. No. 160, 1932, pp. 1-224. Schmidt, K.\rl P. 1931. New species of North American lizards of the genus Holbrokia and Uta. Amer. Mus. Nov.. No. 22. Dec. 1, 1931, pp. 1-6. Stejneger, Leonh.\hd. 1893. North American fauna. No. 7. May 1, 1893, pp. 159-224. Smith, Hob.art. 1935. Miscellaneous notes on Mexican lizards. Univ. Kan. Sci. Bull.. XXII, No. 6, April 15, 1935, pp. 119-155, plates XXII to XXV. 1935a. Descriptions of new species of lizards from Mexico of the genus Uta, with notes on other Mexican species. Univ. Kan. Sci. Bull., XXII, No. 7. April 15. 1935. pj.. 157-183, plate XXVI. 19351). Notes on some Mexican lizards of the genus Holbrookia, with the description of a new species. T^niv. Kan. Sci. Bull.. XXII. No. 8, April 15. 1935, ])p. 185-201, plates XXVII-XXVIII. T.WLOR, E. H. 1935. Colconyx fasciatus, a neglected species of gecko. Univ. Kan. Sci. Bull.. XXII, No. 9, April 15, 1935. pp. 203-205. 1937. New species of Mexican amphibia. Trans. Kan. Acad. Sci.. vol. 39, 1937. pp. . V.A.X Denburgh, John. 1897. Reptiles from Sonora, Sinaloa, and Jalisco, Mexico, with a description of a new species of Sceloporus. Proc. Acad. Nat. Sci. Phila., 1897, pp. 460-464. 532 The University Science Bulletin PLATE XLIV Figure 1. Scaphinpu.s couchii No. 2915 E. H. T. & H. M. S. Mazatlan, Sinaloa. 2. Scophiopit>< couchii No. 2916 E.H.T. it H.M.S. Mazatlan. Sinaloa. 3. Scaphiopus couchii No. 2911 E.H.T. it H. M.S. Giiaymas, Sonera. 4. Scuphinpiis couchii No. 2912 E.H.T. it H.M.S. C;uayma.s. Sonora. Taylor: Fauna of Sixaloa 533 PLATE XLIV 2. f 534 The University Science Bulletin PLATE XLV Figure 1. Bufo kcUoggi No. 42 E.H.T. ct H. M.S. yomio-. Mazatlan, Sinaloa. 2. Bufo kcUoggi No. 27 E.li.T. & B.. M.S. S- Mazatlan, Sinaloa. 3. Biij,, kiUoggi No. 21 E.H.T. it H. M.S. Type. ?. Mazath'm, Sinaloa. 4. Bujo dcbilis No. 11530 K. U. young. San Diego county, Texas. 5. Bufo dcbilis No. 21524 K. U. S . Benton, Atascosa county. Texas. 6. Bufo (Icbili.^ No. 21526 K. U. 9 . Benton, Atascosa county, Texas. 7. Bufo iu.^idior No. 64 E. H. T. & H. M. S. young. Two miles south Majoma, Zacatecas. 8. Bufo inddior No. 87 E. H. T. .k H. M. S. i . Two miles south Majoma, Zacatecas. 9. Bufo iuddior No. 123 E. H. T. & H. M. S. $. Fifteen miles south of Zacatecas, Zacatecas. Taylor: Fauna of Sinaloa 535 PLATE XLV ^.^A^ 7 "- u :) o s ■- - - >■ " v,^. 536 The University Sciexce Bulletin' PLATE XLVI Figure 1. Trimorphodon -pauchnaculatus, sp. nov. Xo. 709, E. H. T. Type. Head, enlarged. Actual size, length to angle of jaw, 25 mm., width, 14 mm. Mazatlan, Sinaloa. 2. Coleonyx ja^^ciatus Xo. 556, E. H. T. Fifteen niilef 80uth of Presidio, Mazatlan, Sinaloa. Taylor: Fauna of Sixaloa 537 PLATE XLVI 'rx '<■ \ z. ! THE UNIYERSITY OF KANSAS SCIENCE BULLETIN Vol. XXIV.] July 15, 1936 [No. 21. The Lizards of the Torquatus Group of the Genus Sceloporus Wiegmann, 1828* HOBART M. SMITH Department of Zoology, University of Kansas Ab.stract: Complete synonymies, descriptions and a key are gi\'en for the recognized species and subspecies of the lurquaiiis group, in which are inckided serrifer, t. (orqndlu.^, I. melanogasler, hullerl, m. 7nucroiiatus, m. omiltemanus, cynn()a)-iabilis, utiformis, pyrocephnlus, macu- losus, microlepldutus, jorrnoaus, spinosus, undulaLus and torquatu-s) of the genus, and of the species and subspecies of the torquatus group. It is sug- gested that the genus Utn is dcMived from ancestral forms of SceIoj)orus whose most closely related lixing s])eci(>s arc in I he variabilis group. INTRODUCTION S("l']LOP()lvUS is one of the largest, most progressive and most recent of the new world lizard genera. Its extreme genetic activity has resulted in the appearance of numerous variations, many of which have not become fixed, partially because of an insufficiency of the time element, an inadequacy of their survi\-al value, and unfavorable environmental conditions. A thorough re- *After this manuscript was in press, Mr. .Joseph R. Bailey kindly pointed out to me that, unfortunately, the name Sce!oporu!< iorqiintus Wiegmann is not tenable. In 1820 Wied de- scribed Stellii) tor([uatuf:, which was placed by Wied in his new genus Tropidurus rlescribed ill 1824 (Al)l)ild. Nat. Bras.). In 1828 Wiegmann described Sceloporus torquatus, which the same author designated as the genotype of Sceloporus in 1834. In 1830 Wagler plaicetl, Sceloporus torquatus (also S. spinosus and (/rammicus) in the genus Tropidurus, together with Tropidurus torciuatus. Thereby Sceloporus torc/uatus is suppressed as a homonym and cannot he resurrected. The only names synonymous with torciuatus are mehuwgaster Cope and jerrariperezi Cope, both described in 188') in the same article. S. melaiiofiaster is very definitely typical of the northern subspecies, while jerrariperezi is based upon intergrades between the northern and southern subspecies, definitely approaching the southern subspecies more closely than the noi"thern. There seems to be no alternative for the selection of jerrariperezi as the name to replace torquatus. The specific and subsjiecific names of the races of the species previously known as Sceloporus torquatus Wiegmann should therefore lie Sceloporus jerrariperezi jer- rariperezi Cope and Sceloporus jerrariperezi mekinogaster Cope. As Sceloporus poinsettii Baird and Girard (1854) is the next oldest name in the group, I propose that the group previously known as the torquatus group be called the poinsettii group. (539) 540 The University Science Bulletin vision which describes and analyzes to a reasonable extent these extraordinarily numerous variations has long been one of the major desiderata in the study of American herpetology. Few American genera of lizards present to students of herpetology a greater array of taxonomic difRculties than this. The greatest difficulty encoun- tered is the determination of the extent of fixation of the sundry variations; while a second difficulty is that of determining what degree of fixation is required that a variant population be given taxonomic recognition. The several authors who have dealt with the genus have dis- agreed with each other in a surprising number of cases. They have expressed the result of their studies in one of two extremes — either the unwarranted recognition of individual or aberrant variations, or equally unwarranted synonymizing under one name of distinct species or subspecies which have in common certain obvious simi- larities but which also have dissimilar characters, overlooked or not well appreciated, which do define distinct and separate genetic entities. This paper is presented as the first part of a proposed revision of the genus Sceloporus. It deals with one unit — the torquatus group. Of this group, about 1,800 specimens have been examined. Most of the Mexican material and many of the specimens from the United States are in the collections at the University of Kansas; and most of the specimens of this group in other museums of the United States have also been examined. Studies of this nature must of a necessity be somewhat incom- plete, because of the ina(lec[uacy of available material, the lack of more direct evidence of relationships, and the lack of absolute knowledge of the methods of speciation. Since such data are not available, in many cases the only recourse is to speculation, with a varying amount of probability of truth in each case. For these reasons many of the conclusions presented hei'cwith must be con- sidered tentative. ACKXOWLEDGM EXTS It has been possible to complete the present work only through the invaluable courtesies of numerous individuals who have gener- ously afforded aid. Among these I wish to express my indebtedness in particular to Dr. Edward H. Taylor, who has been of direct aid in many ways, and whose concepts of speciation and related phe- nomena have assisted in arriving at the conclusions here presented. I am indebted also to Dr. H. H. Lane, of the University of Kansas, Smith: Genus Scelopokus 541 for much invaluable advice and assistance in other respects; to Mr. C. D. Bunker, curator of Dyche Museum of Birds and Mammals, for permission to study specimens under his care; to Dr. K. P. Schmidt, for permission to examine specimens in the Field Museum ; to Mrs. Helen T. Gaige, for permission to examine specimens in the Museum of Zoology of the University of Michigan ; to Dr. Howard K. Gloyd, of the same institution, for the opportunity of making field studies under his direction; to Dr. Thomas Barl:iour and Mr. Arthur Loveridge, for permission to examine specimens in the Mu- seum of Comparative Zoology; to Mr. David H. Dunkle, of Harvard ITniversity; to Dr. G. K. Noble, for permission to study specimens in the American Museum of Natural History; to Mr. C. F. Kauffeld, of the same institution, for numerous courtesies; to Drs. Leonhard Stejneger and Doris Cochran, for permission to examine specimens in the United States National Museum; to Dr. H. ^^■. Fowler, for permission to examine specimens in the Phihidelphia Academy of Natural Sciences; to Mr. L. M. Klauber, for pcrinis>i()ii to examine specimens in his personal collection, and for \-nrious data; to Sefiores Dr. Isaac Ochoterena and Rafael Martin del Canipo, of tlie Tnstituto de Biologia in Mexico City, for numerous courtesies; and to Dr R. H. Painter, of Kansas State College of Manhattan, for numerous courtesies. The study has been aided financially by grants from the graduate research fund of the University of Kansas. The drawings have been made by Miss Myra Wildish, Mr. Russell Chezem, Mr. Carol Johnson and Mr. Maxim Eliaslievich, all of the University of Kan- sas. TUc photographs are tlie woik of Mr. Oi'cn Bingham, also of the Un '.versify of Kansas. EXPLORATION A difficulty frequently encountered by specialists in attempting to loosen taxonomic knots is that of securing an acquaintance with their subjects in their natural state. It has been my good fortune to observe in the field most of the species dealt with herein. My first opportunity was during the summer of 1930, collecting in Texas, New Mexico and Arizona with Mr. Howard K. Gloyd and his wife. In the summer of 1931 I collected in Texas and New Mexico with Dr. and Mrs. R. H. Painter, of Kansas State College at Manhattan. The introduction to Mexican Scelopori was made possible by Dr. Edward H. Taylor, whom I accompanied during the summer of 1932 on an expedition covering seventeen central and northern Mexican states. In the summer of 1934 Mr. David H. Dunkle, now at Har- 542 The University Science Bulletin vard University, and myself, ventured again into northern Mexico. A third trip, which covered most of the other states of Mexico, was undertaken during the summer of 1935. METHOD OF MEASUREMENT AND DESCRIPTION OF DL^GNOSTIC CHARACTERS For each specimen studied in detail, forty-seven characters of proportion and scalation were recorded. Certain measurements in- cluded in the tables presented herewith may be explained. The dis- tance from snout to occiput is measured from the posterior edge of the interparietal to the end of the snout, in a plane parallel with the edge of the mouth. The snout to ear measurement is taken from the posterior border of the ear, in a line parallel to the median vertical axis of the head. The hind leg is measured along its pos- terior border, from the insertion of the leg to the tip of the fourth toe. excluding the claw. The fourth toe is measured from the base of the fifth, and does not include tlie claw. Tlie tibia is measured along its anterior border, flexing the tibia back against the femur, holding the metatarsus at right angles to the tibia, and measuring from the inner angle of the tibiometatarsal joint to the proximal end of the tibia. The scales of importance in descriptions of species of the tor- quatus group are as follows: Interparietal (occipital). A large scale in the middle at the pos- terior edge of the head, always single, with a spot marking the posi- tion of the parietal foramen. The scale is usually more or less square, and its relative size is of some importance. At birth of the animal the interparietal is cfuite large, and gradually decreases in relative size as the animal attains greater age. In species wliich attain greatest size, the interparietal is relatively the smallest. Parietah. A single, usually more or less triangular parietal bor- ders the interparietal on either side. I do not consider the smaller, smooth scales occasionally present behind this scale as parietals. Frontoparietals. Normally a single pair of small, rectangular frontoparietal precede the parietals, separating the interparietal from the row of scales about the supraoculars. The frontoparietals may contact each other medially, or may be separated either by an azygous scale or by contact of the frontal with the interparietal. In some species the frontoparietals are usually divided into four scales. Smith: Genus Sceloporus 543 Frontal. This is a large scale between the orbits, normally trans- versely divided into two sections, the anterior being somewhat larger than the posterior. In ornatus the tendency is to lose the posterior portion of the frontal, probably by fusion with adjacent scales. In koculars _^,_.-^ superolianes ■::,'^i^''^'// xpreocular "'' ' y /r